WD Repeat Domain 6 (WDR6) – Structure, Function, and Biological Roles OpenAI o3-deep-research-2025-06-26 120 citations 2025-12-27T18:06:46.092475

WD Repeat Domain 6 (WDR6) – Structure, Function, and Biological Roles

Introduction and Key Characteristics

WDR6 (WD repeat domain 6) is a human protein encoded by the WDR6 gene, belonging to the WD-repeat protein family. WD-repeat proteins are defined by repeating ~40–60 amino acid motifs typically ending in a tryptophan-aspartate (W-D) dipeptide (pmc.ncbi.nlm.nih.gov). These repeats fold into β-propeller structures that serve as scaffolds for protein–protein interactions, enabling assembly of multi-protein complexes (pmc.ncbi.nlm.nih.gov). Consistent with this role, WDR6 is composed of 11 WD-repeat units and is ~1121 amino acids in length (pubmed.ncbi.nlm.nih.gov). Notably, the repeats of WDR6 are arranged in two clusters separated by a hydrophobic segment that was initially predicted as a transmembrane region (pubmed.ncbi.nlm.nih.gov). Despite this predicted domain, experiments indicate WDR6 is predominantly an intracellular protein localized to the cytoplasm, similar to its interaction partner LKB1 (pubmed.ncbi.nlm.nih.gov). Early northern blot analyses showed WDR6 is ubiquitously expressed in adult and fetal human tissues (pubmed.ncbi.nlm.nih.gov), suggesting it serves fundamental cellular functions. Indeed, WDR6 has emerged as a multifaceted regulatory adaptor protein implicated in diverse processes including RNA modification, cell cycle control, metabolism, and intracellular signaling.

Structurally, WDR6 is a member of a highly conserved subfamily of WD proteins. It shows homology to yeast proteins like Trm734, a factor required for tRNA modification (pubmed.ncbi.nlm.nih.gov). In fact, WDR6 is the human homolog of yeast Trm734, which foreshadows one of WDR6’s key roles in RNA biology. No close paralogs of WDR6 exist in humans beyond the general WD-repeat motif (WDR1–WDR5 are distinct proteins) (pubmed.ncbi.nlm.nih.gov). The WD β-propeller domain of WDR6 (InterPro: IPR015943) suggests it provides a platform for binding other macromolecules, consistent with WDR6’s role in multi-protein complexes. Below, we discuss the current understanding of WDR6’s functions, drawing on recent research (primarily 2020–2023) and how this protein contributes to specific biochemical pathways, cellular localization of its activity, and potential real-world applications.

Role in tRNA Anticodon Loop Modification and Translation

One well-characterized function of WDR6 is as a co-factor in tRNA modification. WDR6 is annotated as a “tRNA (34-2'-O)-methyltransferase regulator,” referring to position 34 in the tRNA anticodon loop (www.ncbi.nlm.nih.gov) (pubchem.ncbi.nlm.nih.gov). Specifically, WDR6 associates with the methyltransferase enzyme FTSJ1 (also known as Trm7 in other organisms) to catalyze 2′-O-methylation of ribose at the wobble position (nucleotide 34) of specific tRNAs (pmc.ncbi.nlm.nih.gov). FTSJ1 is an S-adenosylmethionine-dependent methyltransferase, and WDR6 serves as an essential auxiliary subunit for its activity (pmc.ncbi.nlm.nih.gov). A 2020 study by Li et al. reconstituted the human FTSJ1–WDR6 complex in vitro and demonstrated that it methylates the 2’-OH of nucleotide 34 (forming Cm34 or Gm34) on target tRNAs (pmc.ncbi.nlm.nih.gov). This modification occurs only on tRNAs that also carry a methylated guanosine at position 37 (m¹G37), indicating that WDR6–FTSJ1-mediated 2′-O-methylation at position 34 is tightly linked to other anticodon-loop modifications (pmc.ncbi.nlm.nih.gov). In cells lacking functional FTSJ1 (such as patient cells with FTSJ1 mutations), certain tRNAs – for example, tRNA^Phe(GAA) – were found to completely lack 2′-O-methylation at C32 and G34 (pmc.ncbi.nlm.nih.gov). This corresponds to the yeast paradigm where Trm7 (FTSJ1 homolog) requires two accessory proteins (Trm732 and Trm734) to modify positions 32 and 34, respectively (pubmed.ncbi.nlm.nih.gov). WDR6 is the higher eukaryote homolog of yeast Trm734, the factor specifically needed for the position 34 modification (pubmed.ncbi.nlm.nih.gov). Consistently, WDR6 is required for proper positioning of the tRNA substrate in the FTSJ1 active site and for full methyltransferase activity (www.genecards.org).

Functionally, these tRNA modifications are critical for accurate decoding during protein synthesis. The anticodon-loop 2′-O-methylations enhance tRNA structure and ribosome binding, thereby influencing translational fidelity and efficiency (pmc.ncbi.nlm.nih.gov). In humans, loss of the FTSJ1/WDR6-mediated modifications has been linked to disease: mutations in FTSJ1 (an X-linked gene) cause a form of non-syndromic intellectual disability, presumably due to downstream effects on protein synthesis in neurons (pubmed.ncbi.nlm.nih.gov). While germline mutations in WDR6 itself have not been reported in this disorder, the requirement of WDR6 for FTSJ1 activity suggests that WDR6 is crucial for the biogenesis of properly modified tRNAs, especially in the brain. Indeed, one analysis found that ~40% of the genes with a strong codon bias for UUU (phenylalanine codon decoded by tRNA^Phe) are related to brain and nervous system functions (pmc.ncbi.nlm.nih.gov). This underscores the idea that WDR6, by enabling specific tRNA modifications, indirectly supports the translation of mRNAs vital for neural development and function.

At the cellular level, WDR6 and FTSJ1 are cytoplasmic enzymes (pmc.ncbi.nlm.nih.gov), acting on cytosolic tRNA pools. WDR6 does not itself catalyze methylation (it lacks a catalytic domain), but serves as a scaffold or guide – likely binding the tRNA and/or the FTSJ1 enzyme – to ensure the correct tRNA conformation for methyl transfer (www.genecards.org). This has parallels to other tRNA-modifying complexes (e.g., the METTL1–WDR4 complex for tRNA m^7G46, where WDR4 is a non-catalytic WD-repeat subunit that orientates the tRNA) (www.ncbi.nlm.nih.gov) (www.nature.com). In summary, WDR6’s primary biochemical function identified so far is in the tRNA epitranscriptomic pathway: it partners with the FTSJ1 methyltransferase to modify the anticodon loop, thereby safeguarding translational accuracy. This role is supported by strong experimental evidence (pmc.ncbi.nlm.nih.gov) and highlights WDR6’s broader theme of action – acting as a non-enzymatic regulator within larger protein complexes.

Interaction with LKB1 and Cell Cycle Regulation

Beyond RNA biology, WDR6 has been linked to tumor suppressor signaling and cell cycle control. In a yeast two-hybrid screen reported in 2007, WDR6 was identified as a binding partner of the serine/threonine kinase LKB1 (also known as STK11) (pubmed.ncbi.nlm.nih.gov). LKB1 is a master kinase that activates AMP-activated protein kinase (AMPK) and other kinases, and germline LKB1 mutations cause Peutz-Jeghers cancer syndrome. The WDR6–LKB1 interaction is cytoplasmic (both proteins colocalize in the cytosol (pubmed.ncbi.nlm.nih.gov)), and functionally WDR6 appears to enhance LKB1’s growth-inhibitory effects. Co-expression of WDR6 with LKB1 in cell culture significantly increased LKB1’s ability to arrest cell proliferation and induce a G₁ cell-cycle block (pubmed.ncbi.nlm.nih.gov). In these experiments, WDR6 synergized with LKB1 to elevate levels of the CDK inhibitor p27^Kip1, a key enforcer of the G₁ checkpoint (pubmed.ncbi.nlm.nih.gov). Reporter assays showed that WDR6 boosted LKB1-dependent transcription from the p27^Kip1 promoter, suggesting WDR6 helps transmit LKB1’s signal to cell cycle regulatory genes (pubmed.ncbi.nlm.nih.gov).

Collectively, these findings provide initial evidence that WDR6 participates in the LKB1 tumor-suppressor pathway (pubmed.ncbi.nlm.nih.gov). Specifically, WDR6 acts as a positive regulator or co-factor of LKB1, aiding it to induce cell-cycle arrest and growth suppression (pubmed.ncbi.nlm.nih.gov). The mechanism is not fully determined, but given LKB1’s role in activating AMPK and related kinases, one hypothesis is that WDR6 might anchor LKB1 near specific substrates or assist in assembling complexes (for example, linking LKB1 to transcriptional co-regulators of the p27 gene) (pubmed.ncbi.nlm.nih.gov). Notably, LKB1 also influences cellular metabolism and polarity; whether WDR6 contributes to those aspects is not yet clear. Importantly, from a functional annotation perspective, this interaction positions WDR6 in a cellular stress and growth checkpoint pathway. It also connects WDR6 to cancer biology: loss of LKB1 is common in lung adenocarcinoma and other cancers, and WDR6’s ability to augment LKB1’s function suggests it may have tumor-suppressive effects when LKB1 is present (pubmed.ncbi.nlm.nih.gov). Indeed, the RefSeq gene summary (2016) for WDR6 highlights that WDR6 is “implicated in cell growth arrest” via its interaction with LKB1 (www.genecards.org). This role exemplifies WDR6 functioning as an adapter protein in signaling pathways – in this case, linking a kinase (LKB1) to the cell cycle machinery.

Involvement in Autophagy and Nutrient Signaling

Another emerging role for WDR6 is in autophagy regulation, particularly during nutrient stress. Evidence for this came from a genetic screening study in 2012 that pointed to WDR6 as a factor in starvation-induced autophagy (pmc.ncbi.nlm.nih.gov). Autophagy is a catabolic process activated during amino acid starvation to recycle nutrients, and its regulation involves nutrient-sensing pathways (e.g. mTOR and AMPK). WDR6 has been reported as necessary to suppress autophagy when cells are starved for amino acids (www.genecards.org). In other words, in the absence of WDR6, cells may undergo heightened autophagy upon nutrient deprivation. This was initially a somewhat counterintuitive finding – since starvation normally induces autophagy – but it suggests WDR6 might normally act to fine-tune or limit the autophagic response (pmc.ncbi.nlm.nih.gov). One possibility is that WDR6 interfaces with the AMPK–mTOR pathway (LKB1 is an upstream kinase for AMPK, and AMPK activation promotes autophagy). The WDR6–LKB1 connection hints that WDR6 could modulate AMPK or mTOR signaling, thereby affecting autophagy induction (pmc.ncbi.nlm.nih.gov). In support of this notion, WDR6 was found to co-immunoprecipitate with LKB1 (upstream of AMPK) and to appear in an autophagy-focused interaction screen, although those reports did not fully clarify the mechanism (pmc.ncbi.nlm.nih.gov).

Recent research has provided more direct insight into how WDR6 can regulate autophagy and how this ties into disease contexts. A 2023 study in EMBO Molecular Medicine examined WDR6’s function in liver cancer and uncovered a mechanism connecting WDR6 to the autophagy machinery (pmc.ncbi.nlm.nih.gov). The authors found that WDR6 binds to UVRAG (UV radiation resistance-associated gene protein), which is a key component of the autophagy initiation complex (part of the Beclin1-PI3KC3/Vps34 pathway). WDR6 targets UVRAG for degradation by recruiting it to a Cullin-4A E3 ubiquitin ligase complex (CUL4A–DDB1–ROC1) (pmc.ncbi.nlm.nih.gov). WDR6 contains a unique “WDxR” motif – a short sequence within a WD-repeat blade – that mediates this recruitment to the E3 ligase (pmc.ncbi.nlm.nih.gov). Through this mechanism, WDR6 promotes UVRAG’s ubiquitination and proteolysis, effectively blunting autophagy initiation (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Consequences of UVRAG loss were observed as an accumulation of the transcription factor NF-κB p65, which under normal conditions can be degraded via autophagy (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). In WDR6-proficient cells, UVRAG is degraded, autophagic turnover of p65 is blocked, and NF-κB (p65) remains active, driving robust production of TNFα (tumor necrosis factor) (pmc.ncbi.nlm.nih.gov). Thus, WDR6 creates a link between autophagy and inflammatory signaling: by suppressing autophagic clearance of p65, it sustains NF-κB/TNFα signaling.

This WDR6-NF-κB connection has significant effects on the immune environment, especially in tumors (detailed in the next section). From a cell biology standpoint, it solidifies WDR6’s role as a regulator of autophagy and stress responses. WDR6’s inhibition of autophagy (via UVRAG degradation) aligns with earlier observations that WDR6 is required to suppress autophagy during starvation (www.genecards.org). It appears WDR6 might ensure that autophagy doesn’t overshoot or that certain autophagy-related proteins are kept in check. The discovery of the WDxR motif as a functional element also suggests WDR6 may belong to a class of DDB1–CUL4-associated factors (DCAFs) – substrate adaptors that present specific targets (like UVRAG) to the CUL4 E3 ubiquitin ligase (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Indeed, other WD-repeat proteins are known to serve as substrate receptors for CUL4-based E3 complexes (pmc.ncbi.nlm.nih.gov), and WDR6 now appears to be one of them.

In summary, WDR6 modulates autophagy by controlling the stability of an autophagy initiator (UVRAG). This places WDR6 at a crossroads between nutrient-sensing pathways, autophagic flux, and inflammatory signaling. The autophagy-suppressing function of WDR6 could have broad implications: excessive or dysregulated autophagy is involved in neurodegeneration and cancer; WDR6 might normally help restrain autophagy to balanced levels, though in pathological contexts this restraint can be double-edged (as discussed below for tumors). These insights, mostly uncovered in the last few years, highlight WDR6’s expanding functional repertoire beyond its original identification as a tRNA methylation cofactor.

Metabolic Functions in Insulin Signaling and Lipogenesis

Emerging evidence indicates that WDR6 also plays a role in metabolic regulation, particularly in the context of insulin signaling and lipid metabolism. A 2023 study in Nature Metabolism found that WDR6 is upregulated in the livers of insulin-resistant mice and drives hepatic de novo lipogenesis (DNL) – the synthesis of fatty acids in the liver (www.nature.com). Under normal physiology, insulin promotes lipogenesis via transcription factors like SREBP1c. In insulin resistance (IR), certain insulin signals fail (e.g. glucose uptake), but paradoxically the lipogenic pathway often remains active, contributing to fatty liver (hepatic steatosis) (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Investigating this phenomenon, researchers discovered that WDR6 levels increase in the liver during high-fat-diet induced insulin resistance, and that WDR6 is required for the continued activation of lipogenic gene expression in this state (pmc.ncbi.nlm.nih.gov).

Mechanistically, WDR6 was shown to bind the β-catalytic subunit of protein phosphatase 1 (PPP1CB) (www.nature.com). Protein phosphatase-1 is a key serine/threonine phosphatase whose activity can be modulated by regulatory subunits. WDR6 appears to function as a targeting/regulatory subunit for PP1β. Specifically, WDR6 binding facilitates dephosphorylation of PP1β at Thr316 (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Dephosphorylation at this site is thought to enhance PP1β activity (Thr316 phosphorylation can negatively regulate PP1 activity (pmc.ncbi.nlm.nih.gov)). Active PP1β, in turn, was found to influence a signaling axis involving DNA-dependent protein kinase (DNA-PK) and the transcription factor USF1 (pmc.ncbi.nlm.nih.gov) (www.nature.com). The outcome of this cascade is increased transcription of FASN, the gene encoding fatty acid synthase, which is a rate-limiting enzyme in fatty acid production (pmc.ncbi.nlm.nih.gov) (www.nature.com). In WDR6 knockout liver cells, FASN expression was significantly reduced, confirming WDR6’s role in upregulating lipogenic enzymes (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Conversely, expressing a mutant PP1β that mimics the dephosphorylated state (Thr316Ala) elevated FASN levels, whereas a phosphomimetic PP1β (Thr316Asp) depressed FASN, matching the effect of WDR6 on PP1β regulation (pmc.ncbi.nlm.nih.gov). These findings suggest that WDR6 promotes lipogenesis by activating PP1β, which then triggers a pro-lipogenic gene transcription program (pmc.ncbi.nlm.nih.gov).

Importantly, this study also demonstrated a potential therapeutic angle: using molecular modeling and screening, the authors identified a small natural compound (referred to as XLIX) that disrupts the WDR6–PPP1CB interaction, thereby keeping PP1β in a phosphorylated (less active) state (www.nature.com). Treatment with this compound in insulin-resistant mice reduced hepatic FASN expression and lipid accumulation, effectively mitigating fatty liver development (www.nature.com). This proof-of-concept highlights WDR6 as a promising drug target for metabolic diseases like non-alcoholic fatty liver disease (NAFLD) associated with insulin resistance (www.nature.com). In summary, WDR6 has a critical role in a novel insulin-regulated pathway: by acting as a PP1 regulatory subunit, it connects insulin signaling to the control of a major metabolic enzyme. This function again reflects WDR6’s general theme of action as a scaffold protein – here, bringing together a phosphatase (PP1β) with its target pathways to modulate phosphorylation-dependent signals.

It is intriguing that WDR6 interacts with an insulin receptor substrate in other contexts as well. An earlier proteomics study in rat brain found WDR6 as a binding partner of IRS4 (insulin receptor substrate-4) (www.ncbi.nlm.nih.gov), which may hint at additional links between WDR6 and insulin/IGF signaling pathways. While the full scope of WDR6’s metabolic involvement is not yet known, the current data firmly establish WDR6 as an insulin-responsive regulator of lipid metabolism. Given the rise in metabolic syndrome and NAFLD cases, the discovery of WDR6’s role in lipogenesis is a significant development that occurred in 2023. It provides a specific biochemical function (regulation of PP1β activity) for WDR6 in the metabolic context, expanding the protein’s functional annotation.

Implications in Cancer and Immunology

Considering WDR6’s roles in cell growth (LKB1 pathway), autophagy, and metabolism, it is perhaps not surprising that dysregulation of WDR6 has been linked to cancer. However, until recently WDR6 was a relatively little-explored gene in oncology (pmc.ncbi.nlm.nih.gov). New research has begun to clarify WDR6’s impact on tumor biology and the tumor immune microenvironment. The 2023 EMBO Molecular Medicine study on hepatocellular carcinoma (HCC) is particularly illuminating. This study found that WDR6 promotes an immunosuppressive, tumor-promoting microenvironment in HCC (pmc.ncbi.nlm.nih.gov). In a series of experiments, the authors showed that knocking out WDR6 in mouse HCC cells had minimal effect on their growth in vitro, but drastically inhibited tumor growth and lung metastasis in vivo when these cells were implanted into immune-competent mice (pmc.ncbi.nlm.nih.gov). In immune-deficient mice, WDR6 status had little impact on tumor growth (pmc.ncbi.nlm.nih.gov), indicating that WDR6’s pro-tumor effects are largely mediated through the immune system rather than cell-intrinsic proliferation.

Mechanistically, as discussed, WDR6 drives NF-κB/TNFα signaling by blocking autophagic degradation of p65 through UVRAG ubiquitination (pmc.ncbi.nlm.nih.gov). The consequence in tumors is that high WDR6 expression leads to chronically elevated TNFα levels, which reshape the immune cell population in the tumor (pmc.ncbi.nlm.nih.gov). TNFα is a cytokine that can recruit immunosuppressive myeloid cells. Indeed, WDR6-proficient HCC tumors had a higher infiltration of MDSCs (myeloid-derived suppressor cells) and a lower presence of activated CD8⁺ T cells (pmc.ncbi.nlm.nih.gov). MDSCs are known to suppress T-cell responses, and their accumulation creates a microenvironment conducive to tumor progression. The study demonstrated that these effects could be partly reversed by blocking TNFα – inhibiting TNFα signaling led to improved T-cell infiltration and tumor control in WDR6-high tumors (pmc.ncbi.nlm.nih.gov). Furthermore, TNFα itself was found to feed back and increase WDR6 transcription via NF-κB, establishing a vicious positive feedback loop (WDR6 → ↑TNFα → ↑WDR6) (pmc.ncbi.nlm.nih.gov). This loop helps explain how WDR6 may become upregulated in cancers and sustain an immunosuppressive niche. Clinically, analysis of HCC patient samples showed that the WDR6/UVRAG/NF-κB axis is hyperactivated in a subset of HCC and correlates with poor prognosis (pmc.ncbi.nlm.nih.gov). High WDR6 expression in tumors could therefore serve as a marker of aggressive disease.

Excitingly, the HCC study also provided a therapeutic proof-of-concept: a peptide that mimics the WDxR motif was used to disrupt the WDR6–UVRAG interaction, thereby preventing UVRAG degradation (pmc.ncbi.nlm.nih.gov). In preclinical models, this WDxR-mimetic peptide restored autophagy, reduced TNFα levels, and enhanced the efficacy of anti-PD-L1 immunotherapy against WDR6-driven tumors (pmc.ncbi.nlm.nih.gov). Anti-PD-L1 (checkpoint blockade) is more effective when immunosuppression in the tumor microenvironment is relieved, so breaking the WDR6-TNFα loop made the tumors more responsive. This finding suggests that WDR6 could be a novel immuno-oncology target: patients with WDR6-overexpressing tumors might benefit from therapies that inhibit WDR6 function or its downstream effects. This is a striking example of how deep functional research on a gene can directly inform potential translational strategies.

Apart from HCC, WDR6’s role in other cancers is also being investigated. A bioinformatics-driven study in 2022 analyzed WDR6 expression across cancers, with a focus on lung cancer. Interestingly, in contrast to HCC, higher WDR6 expression was associated with better survival in non-small cell lung cancer patients (pmc.ncbi.nlm.nih.gov). In lung adenocarcinoma (LUAD), WDR6 levels correlated positively with advanced tumor stage (higher in later stages) (pmc.ncbi.nlm.nih.gov), yet paradoxically patients with WDR6-high tumors had longer overall and relapse-free survival compared to WDR6-low patients (pmc.ncbi.nlm.nih.gov). The same study reported that WDR6 expression in lung cancer correlates with markers of immune infiltration – including levels of various lymphocytes, immunomodulatory genes, and chemokines (pmc.ncbi.nlm.nih.gov). One interpretation is that WDR6, being tied to immune pathways, might reflect an active immune environment in lung tumors that helps control the cancer (hence high WDR6 linking to better outcomes) (pmc.ncbi.nlm.nih.gov). This is somewhat at odds with the HCC findings (where WDR6 promoted an immune-suppressive environment), suggesting that WDR6’s impact may be context-dependent or differ by tissue. It’s possible that in lung cancers, WDR6’s association with LKB1 status is a factor – for instance, LKB1-inactivated lung tumors are known for poor immune surveillance, and if WDR6 requires LKB1 to execute certain functions, the net effect of WDR6 could differ. More research is needed to reconcile these observations. Nonetheless, the use of WDR6 as a prognostic biomarker has been proposed: one study concludes WDR6 could serve as a novel marker for survival and immune infiltration status in lung cancer (pmc.ncbi.nlm.nih.gov).

In summary, WDR6’s involvement in cancer spans multiple dimensions: cell-intrinsic roles (affecting cell cycle via p27 and metabolism via FASN) and cell-extrinsic roles (shaping the tumor immune microenvironment via NF-κB/TNFα and autophagy pathways). The current consensus is that WDR6, while not a classic oncogene or tumor suppressor by itself, functions as a modifier of signaling pathways that cancer cells hijack. Its ubiquitous expression and ability to interact with key players (kinases, phosphatases, ubiquitin ligases) put it in a position to modulate cancer-related phenotypes. As research from 2020–2023 has shown, WDR6 can both positively and negatively influence cancer progression, depending on the context, by rewiring metabolic and immune signals.

Conclusion and Future Perspectives

WDR6 is a prime example of a multi-functional scaffold protein whose importance has become clearer in recent years. Initially characterized simply as a novel WD-repeat protein with no known function, WDR6 is now recognized to play critical regulatory roles in:

These diverse functions are unified by a theme: WDR6 serves as a molecular adaptor or co-factor, bringing enzymes or signaling molecules into the correct complexes or conformations to execute their functions. Through its WD-repeat domains, WDR6 can simultaneously bind multiple partners – a property that underlies its role in assembling the FTSJ1-tRNA complex, the LKB1 signaling complex, the PP1-DNA-PK-USF1 phosphatase complex, and the CUL4A-DDB1-UVRAG ubiquitin ligase complex. Its subcellular localization is mainly cytosolic (pubmed.ncbi.nlm.nih.gov), but it likely shuttles or localizes to where its partner proteins are (e.g., at the ribosome/tRNA interface, in the nucleus at gene promoters, or at autophagosome assembly sites) as needed.

From a functional annotation standpoint, WDR6 can be described as a scaffolding protein that regulates enzyme specificity and signal transduction. It has no known enzymatic activity of its own; instead, it confers substrate specificity or stability to enzymes like methyltransferases and phosphatases (www.genecards.org). Loss-of-function of WDR6 could thus have pleiotropic consequences, as seen in experimental models (e.g., impaired tRNA modification and codon translation, increased autophagy, reduced lipogenesis, and changes in cell cycle dynamics). However, because WDR6 is not absolutely essential for viability (mice and cells can survive without it, albeit with specific defects (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov)), its functions seem to fine-tune cellular processes rather than being wholly indispensable for basal survival.

In terms of real-world applications and ongoing research: WDR6 is gaining interest as a potential therapeutic target. The discovery of compounds and peptides that can inhibit WDR6 interactions (as shown for metabolic and cancer contexts) opens avenues for drug development (www.nature.com) (pmc.ncbi.nlm.nih.gov). For instance, a WDR6 inhibitor could be explored to treat fatty liver disease by reducing FASN-driven lipogenesis, or as an adjuvant in cancer immunotherapy to prevent WDR6-mediated immune evasion. Additionally, WDR6 expression levels might serve as biomarkers for certain diseases – high WDR6 could indicate a tRNA modification deficiency if FTSJ1 is mutated (helping in the diagnosis of related intellectual disability) or might inform prognosis and treatment choices in cancers (as suggested for lung cancer and HCC) (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov).

Going forward, there are still open questions. It will be important to delineate how WDR6 is regulated (what controls its expression and activity) and whether any post-translational modifications on WDR6 affect its function. Some data suggest WDR6 itself might be subject to regulation by major pathways (for example, NF-κB can induce WDR6, and insulin can upregulate WDR6) (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Understanding these regulatory loops could provide further insight into how WDR6 integrates into cellular homeostasis. Moreover, the full set of WDR6-interacting proteins is likely broader than currently known; a systematic proteomic analysis could reveal additional partners and pathways involving WDR6.

In conclusion, WDR6 has evolved from an uncharacterized gene into a protein of significant interest due to its role in fundamental cellular processes and disease-related pathways. Its WD-repeat structure underpins versatile binding capabilities, allowing WDR6 to act as a critical node linking enzymes to their substrates or signaling molecules to their pathways. The latest research (2020–2024) has shed light on WDR6’s functions in tRNA modification, tumor suppression, autophagy, immunity, and metabolism, making it a compelling subject for further investigation in both basic biology and translational medicine (pmc.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov) (www.nature.com). As we continue to unravel WDR6’s interactome and regulatory mechanisms, we gain not only a deeper understanding of cellular complexity but also new strategies to combat diseases where WDR6’s pathways are dysregulated.

References: (Publication dates and sources for key studies)
- Li et al., 2020, EMBO Reports – Defined WDR6’s role with FTSJ1 in tRNA 2′-O-methylation (pmc.ncbi.nlm.nih.gov).
- Xie et al., 2007, Biochem. Biophys. Res. Commun. – Discovered WDR6’s interaction with LKB1 and effect on p27^Kip1 (pubmed.ncbi.nlm.nih.gov).
- McKnight et al., 2012, EMBO J. – Implicated WDR6 in amino-acid starvation induced autophagy (screening study) (pmc.ncbi.nlm.nih.gov).
- Zhang et al., 2023, EMBO Mol. Medicine – Elucidated WDR6–UVRAG–NFκB mechanism in HCC and immune evasion (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov).
- Meng et al., 2023, Nature Metabolism – Demonstrated WDR6’s role in insulin resistance and lipogenesis, and identified a WDR6-inhibiting compound (www.nature.com) (www.nature.com).
- Lv et al., 2022, Immun. Inflamm. Dis. – Correlated WDR6 expression with prognosis and immune infiltration in lung cancer (bioinformatics study) (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov).
- Bi et al., 2000, Biochem. Biophys. Res. Commun. – Cloned human WDR6, reported its sequence, chromosomal location, and ubiquitous expression (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). (Foundational genomic data)

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  6. AnnotationURLCitation(end_index=2058, start_index=1911, title='Molecular cloning, expression analysis, and chromosome mapping of WDR6, a novel human WD-repeat gene - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/10903905/#:~:text=since%20its%2011%20WD%20repeats,probably%20belongs%20to%20a%20highly')
  7. AnnotationURLCitation(end_index=2611, start_index=2479, title="Intellectual disability-associated gene ftsj1 is responsible for 2'-O-methylation of specific tRNAs - PubMed", type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/32558197/#:~:text=Figure%20EV2,or%20Trm734%20from%20different%20species')
  8. AnnotationURLCitation(end_index=2999, start_index=2838, title='Molecular cloning, expression analysis, and chromosome mapping of WDR6, a novel human WD-repeat gene - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/10903905/#:~:text=fluorescence%20in%20situ%20hybridization,are%20its%20distantly%20related%20members')
  9. AnnotationURLCitation(end_index=3820, start_index=3706, title='Wdr6 WD repeat domain 6 [Mus musculus (house mouse)] - Gene - NCBI', type='url_citation', url='https://www.ncbi.nlm.nih.gov/gene/83669#:~:text=...from%20OrthoDB%20,Go%20to%20the%20top')
  10. AnnotationURLCitation(end_index=3914, start_index=3821, title="tRNA (34-2'-O)-methyltransferase regulator WDR6 (human) | Protein Target - PubChem", type='url_citation', url='https://pubchem.ncbi.nlm.nih.gov/protein/Q9NNW5#:~:text=website')
  11. AnnotationURLCitation(end_index=4248, start_index=4126, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  12. AnnotationURLCitation(end_index=4501, start_index=4379, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  13. AnnotationURLCitation(end_index=4807, start_index=4685, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  14. AnnotationURLCitation(end_index=5161, start_index=5039, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  15. AnnotationURLCitation(end_index=5518, start_index=5350, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=tRNA%20modifications%20at%20the%20anti%E2%80%90codon,1%7DG37%20as%20a%20prerequisite')
  16. AnnotationURLCitation(end_index=5818, start_index=5682, title="Defects in tRNA Anticodon Loop 2'-O-Methylation Are Implicated in Nonsyndromic X-Linked Intellectual Disability due to Mutations in FTSJ1 - PubMed", type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/26310293/#:~:text=Nonsyndromic%20X,FTSJ1%2C%20the%20likely%20TRM7%20homolog')
  17. AnnotationURLCitation(end_index=6074, start_index=5942, title="Intellectual disability-associated gene ftsj1 is responsible for 2'-O-methylation of specific tRNAs - PubMed", type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/32558197/#:~:text=Figure%20EV2,or%20Trm734%20from%20different%20species')
  18. AnnotationURLCitation(end_index=6368, start_index=6221, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=Together%20with%20methyltransferase%20FTSJ1%2C%20methylates,')
  19. AnnotationURLCitation(end_index=6777, start_index=6611, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=We%20find%20that%20modifications%20at,FTSJ1%20and%20nervous%20system%20development')
  20. AnnotationURLCitation(end_index=7164, start_index=7028, title="Defects in tRNA Anticodon Loop 2'-O-Methylation Are Implicated in Nonsyndromic X-Linked Intellectual Disability due to Mutations in FTSJ1 - PubMed", type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/26310293/#:~:text=Nonsyndromic%20X,FTSJ1%2C%20the%20likely%20TRM7%20homolog')
  21. AnnotationURLCitation(end_index=7739, start_index=7573, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=We%20find%20that%20modifications%20at,FTSJ1%20and%20nervous%20system%20development')
  22. AnnotationURLCitation(end_index=8153, start_index=7978, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=FTSJ1%2C%20an%20intellectual%20disability%20associated,UUU%20codon%20usage%20biased%20genes')
  23. AnnotationURLCitation(end_index=8557, start_index=8410, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=Together%20with%20methyltransferase%20FTSJ1%2C%20methylates,')
  24. AnnotationURLCitation(end_index=8868, start_index=8734, title='Wdr6 WD repeat domain 6 [Rattus norvegicus (Norway rat)] - Gene - NCBI', type='url_citation', url='https://www.ncbi.nlm.nih.gov/gene/301007#:~:text=NCBI%20www.ncbi.nlm.nih.gov%20%20...GeneRIF%20,Norway%20rat')
  25. AnnotationURLCitation(end_index=8996, start_index=8869, title='Structures and mechanisms of tRNA methylation by METTL1–WDR4 | Nature', type='url_citation', url='https://www.nature.com/articles/s41586-022-05565-5#:~:text=Nature%20www,Thus%2C%20our%20structural%20models')
  26. AnnotationURLCitation(end_index=9411, start_index=9289, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  27. AnnotationURLCitation(end_index=9974, start_index=9830, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Germline%20mutations%20of%20the%20serine%2Fthreonine,Consistently')
  28. AnnotationURLCitation(end_index=10355, start_index=10223, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=been%20fully%20elucidated,dependent%20kinase%20%28CDK')
  29. AnnotationURLCitation(end_index=10754, start_index=10586, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  30. AnnotationURLCitation(end_index=11015, start_index=10893, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=colony%20formation%20of%20Hela%20cells,Kip1')
  31. AnnotationURLCitation(end_index=11309, start_index=11191, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=WDR6%20was%20able%20to%20synergize,Kip1')
  32. AnnotationURLCitation(end_index=11546, start_index=11428, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=WDR6%20was%20able%20to%20synergize,Kip1')
  33. AnnotationURLCitation(end_index=11851, start_index=11683, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  34. AnnotationURLCitation(end_index=12304, start_index=12136, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  35. AnnotationURLCitation(end_index=12961, start_index=12793, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  36. AnnotationURLCitation(end_index=13230, start_index=13104, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=,provided%20by%20RefSeq%2C%20Feb%202016')
  37. AnnotationURLCitation(end_index=13836, start_index=13669, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=match%20at%20L830%20screening%20suggesting,WDR6%20in%20autophagy%2C%20these%20data')
  38. AnnotationURLCitation(end_index=14239, start_index=14109, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=the%20substrate%20tRNA%20for%20methylation,')
  39. AnnotationURLCitation(end_index=14707, start_index=14538, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=screening%20suggesting%20a%20possible%20role,WDR6%20in%20autophagy%2C%20these%20data')
  40. AnnotationURLCitation(end_index=15142, start_index=14973, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=screening%20suggesting%20a%20possible%20role,WDR6%20in%20autophagy%2C%20these%20data')
  41. AnnotationURLCitation(end_index=15525, start_index=15356, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=screening%20suggesting%20a%20possible%20role,WDR6%20in%20autophagy%2C%20these%20data')
  42. AnnotationURLCitation(end_index=15986, start_index=15811, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  43. AnnotationURLCitation(end_index=16483, start_index=16308, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  44. AnnotationURLCitation(end_index=16794, start_index=16619, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  45. AnnotationURLCitation(end_index=17095, start_index=16920, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  46. AnnotationURLCitation(end_index=17286, start_index=17096, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=Since%20UVRAG%20is%20a%20critical,enhancing%20CUL4A%E2%80%90mediated%20ubiquitination%20and%20degradation')
  47. AnnotationURLCitation(end_index=17649, start_index=17447, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=Ubiquitination%20and%20autophagy%20regulate%20protein,blocking%20autophagy%E2%80%90dependent%20degradation%20of%20p65')
  48. AnnotationURLCitation(end_index=17796, start_index=17650, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=autophagy%E2%80%90dependent%20degradation%20of%20p65%20in,HCC')
  49. AnnotationURLCitation(end_index=18108, start_index=17976, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=mice,HCC%2C%20predicting%20a%20poor%20prognosis')
  50. AnnotationURLCitation(end_index=18788, start_index=18658, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=the%20substrate%20tRNA%20for%20methylation,')
  51. AnnotationURLCitation(end_index=19337, start_index=19147, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=match%20at%20L538%20Since%20UVRAG,enhancing%20CUL4A%E2%80%90mediated%20ubiquitination%20and%20degradation')
  52. AnnotationURLCitation(end_index=19528, start_index=19338, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=Since%20UVRAG%20is%20a%20critical,enhancing%20CUL4A%E2%80%90mediated%20ubiquitination%20and%20degradation')
  53. AnnotationURLCitation(end_index=19823, start_index=19633, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=Since%20UVRAG%20is%20a%20critical,enhancing%20CUL4A%E2%80%90mediated%20ubiquitination%20and%20degradation')
  54. AnnotationURLCitation(end_index=21137, start_index=20994, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  55. AnnotationURLCitation(end_index=21578, start_index=21423, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=The%20human%20WDR6%20protein%20is,relevant%20effects%20of%20WDR6%20and')
  56. AnnotationURLCitation(end_index=21709, start_index=21579, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=tissues,to%20reduce%20fatty%20liver%20disease')
  57. AnnotationURLCitation(end_index=22085, start_index=21955, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=tissues,to%20reduce%20fatty%20liver%20disease')
  58. AnnotationURLCitation(end_index=22333, start_index=22190, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  59. AnnotationURLCitation(end_index=22763, start_index=22599, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=match%20at%20L639%20PPP1CB%29%2C%20phospho,in%20the%20two%20distinct%20datasets')
  60. AnnotationURLCitation(end_index=22925, start_index=22764, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=match%20at%20L649%20interacted%20with,in%20order%20to%20perform%20reciprocal')
  61. AnnotationURLCitation(end_index=23203, start_index=23059, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=PPP1CB%29%2C%20phospho,in%20the%20two%20distinct%20datasets')
  62. AnnotationURLCitation(end_index=23507, start_index=23358, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=phosphatase%201%20,compound%2C%20XLIX%2C%20that%20inhibits%20the')
  63. AnnotationURLCitation(end_index=23651, start_index=23508, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  64. AnnotationURLCitation(end_index=23949, start_index=23819, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=tissues,to%20reduce%20fatty%20liver%20disease')
  65. AnnotationURLCitation(end_index=24093, start_index=23950, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  66. AnnotationURLCitation(end_index=24389, start_index=24225, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=To%20clarify%20the%20physiological%20function,of%20mutant%20PPP1CB%20in%20which')
  67. AnnotationURLCitation(end_index=24546, start_index=24390, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=Thr316%20was%20replaced%20with%20either,Thr316Ala%20and%2C%20conversely')
  68. AnnotationURLCitation(end_index=24927, start_index=24763, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=To%20clarify%20the%20physiological%20function,of%20mutant%20PPP1CB%20in%20which')
  69. AnnotationURLCitation(end_index=25234, start_index=25070, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=To%20clarify%20the%20physiological%20function,of%20mutant%20PPP1CB%20in%20which')
  70. AnnotationURLCitation(end_index=25674, start_index=25534, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=phosphatase%201%20,the%20treatment%20of%20hepatic%20steatosis')
  71. AnnotationURLCitation(end_index=25983, start_index=25834, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,the%20treatment%20of%20hepatic%20steatosis')
  72. AnnotationURLCitation(end_index=26328, start_index=26159, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=protein%20kinase%20and%20upstream%20stimulatory,the%20treatment%20of%20hepatic%20steatosis')
  73. AnnotationURLCitation(end_index=27049, start_index=26935, title='Wdr6 WD repeat domain 6 [Rattus norvegicus (Norway rat)] - Gene - NCBI', type='url_citation', url='https://www.ncbi.nlm.nih.gov/gene/301007#:~:text=Bibliography%20Go%20to%20the%20top,PMID')
  74. AnnotationURLCitation(end_index=28068, start_index=27900, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=and%20triple%E2%80%90A%20syndrome.,immunological%20role%20for%20LUAD%C2%A0and%20LUSC')
  75. AnnotationURLCitation(end_index=28542, start_index=28385, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=promotes%20its%20degradation,against%20HCC%20with%20WDR6%20dysregulation')
  76. AnnotationURLCitation(end_index=28971, start_index=28808, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=The%20WD%E2%80%90repeat%20,degradation%20of%20p65%2C%20elevates%20intratumoral')
  77. AnnotationURLCitation(end_index=29205, start_index=29045, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=The%20WD%E2%80%90repeat%20,chromatin%20accessibility%20at%20the%20TNF%CE%B1')
  78. AnnotationURLCitation(end_index=29649, start_index=29474, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  79. AnnotationURLCitation(end_index=29939, start_index=29807, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=mice,HCC%2C%20predicting%20a%20poor%20prognosis')
  80. AnnotationURLCitation(end_index=30327, start_index=30170, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=promotes%20its%20degradation,against%20HCC%20with%20WDR6%20dysregulation')
  81. AnnotationURLCitation(end_index=30800, start_index=30643, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=promotes%20its%20degradation,against%20HCC%20with%20WDR6%20dysregulation')
  82. AnnotationURLCitation(end_index=31145, start_index=30970, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=myeloid%E2%80%90derived%20suppressor%20cell%20,against%20HCC%20with%20WDR6%20dysregulation')
  83. AnnotationURLCitation(end_index=31587, start_index=31412, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=myeloid%E2%80%90derived%20suppressor%20cell%20,against%20HCC%20with%20WDR6%20dysregulation')
  84. AnnotationURLCitation(end_index=32047, start_index=31876, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=are%20reversed%20by%20TNF%CE%B1%20blockade,against%20HCC%20with%20WDR6%20dysregulation')
  85. AnnotationURLCitation(end_index=32397, start_index=32226, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=are%20reversed%20by%20TNF%CE%B1%20blockade,against%20HCC%20with%20WDR6%20dysregulation')
  86. AnnotationURLCitation(end_index=33384, start_index=33227, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=Our%20results%20showed%20WDR6%20was,and%20chemokines%20in%20lung%20cancer')
  87. AnnotationURLCitation(end_index=33659, start_index=33502, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=Our%20results%20showed%20WDR6%20was,and%20chemokines%20in%20lung%20cancer')
  88. AnnotationURLCitation(end_index=33946, start_index=33789, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=Our%20results%20showed%20WDR6%20was,and%20chemokines%20in%20lung%20cancer')
  89. AnnotationURLCitation(end_index=34332, start_index=34141, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=postprogression%20survival%2C%20and%20Relapse%E2%80%90free%20survival,and%20chemokines%20in%20lung%20cancer')
  90. AnnotationURLCitation(end_index=34688, start_index=34531, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=Our%20results%20showed%20WDR6%20was,and%20chemokines%20in%20lung%20cancer')
  91. AnnotationURLCitation(end_index=35543, start_index=35385, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=found%20the%20expression%20of%20WDR6,and%20chemokines%20in%20lung%20cancer')
  92. AnnotationURLCitation(end_index=36860, start_index=36738, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  93. AnnotationURLCitation(end_index=37131, start_index=36963, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  94. AnnotationURLCitation(end_index=37469, start_index=37294, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  95. AnnotationURLCitation(end_index=37731, start_index=37588, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  96. AnnotationURLCitation(end_index=38401, start_index=38269, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=been%20fully%20elucidated,dependent%20kinase%20%28CDK')
  97. AnnotationURLCitation(end_index=39046, start_index=38899, title='WDR6 Gene - GeneCards | WDR6 Protein | WDR6 Antibody', type='url_citation', url='https://www.genecards.org/cgi-bin/carddisp.pl?gene=WDR6#:~:text=Together%20with%20methyltransferase%20FTSJ1%2C%20methylates,')
  98. AnnotationURLCitation(end_index=39559, start_index=39415, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=PPP1CB%29%2C%20phospho,in%20the%20two%20distinct%20datasets')
  99. AnnotationURLCitation(end_index=39670, start_index=39560, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=PPP1CB%20,These%20results')
  100. AnnotationURLCitation(end_index=40205, start_index=40065, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=phosphatase%201%20,the%20treatment%20of%20hepatic%20steatosis')
  101. AnnotationURLCitation(end_index=40377, start_index=40206, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=are%20reversed%20by%20TNF%CE%B1%20blockade,against%20HCC%20with%20WDR6%20dysregulation')
  102. AnnotationURLCitation(end_index=41061, start_index=40903, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=found%20the%20expression%20of%20WDR6,and%20chemokines%20in%20lung%20cancer')
  103. AnnotationURLCitation(end_index=41237, start_index=41062, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=myeloid%E2%80%90derived%20suppressor%20cell%20,against%20HCC%20with%20WDR6%20dysregulation')
  104. AnnotationURLCitation(end_index=41789, start_index=41614, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=myeloid%E2%80%90derived%20suppressor%20cell%20,against%20HCC%20with%20WDR6%20dysregulation')
  105. AnnotationURLCitation(end_index=41920, start_index=41790, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10590755/#:~:text=tissues,to%20reduce%20fatty%20liver%20disease')
  106. AnnotationURLCitation(end_index=42970, start_index=42848, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  107. AnnotationURLCitation(end_index=43139, start_index=42971, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  108. AnnotationURLCitation(end_index=43297, start_index=43140, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=promotes%20its%20degradation,against%20HCC%20with%20WDR6%20dysregulation')
  109. AnnotationURLCitation(end_index=43441, start_index=43298, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  110. AnnotationURLCitation(end_index=43949, start_index=43827, title='Intellectual disability‐associated gene ftsj1 is responsible for 2′‐O‐methylation of specific tRNAs - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC7403668/#:~:text=owing%2C%20in%20part%2C%20to%20the,Our')
  111. AnnotationURLCitation(end_index=44245, start_index=44077, title='Association of LKB1 with a WD-repeat protein WDR6 is implicated in cell growth arrest and p27(Kip1) induction - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/17216128/#:~:text=Immunofluorescence%20staining%20revealed%20that%20WDR6,that%20WDR6%20is%20implicated%20in')
  112. AnnotationURLCitation(end_index=44538, start_index=44369, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=screening%20suggesting%20a%20possible%20role,WDR6%20in%20autophagy%2C%20these%20data')
  113. AnnotationURLCitation(end_index=44831, start_index=44656, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=HCC%20cells%2C%20its%20deficiency%20in,recruits%20NF%E2%80%90%CE%BAB%20to%20activate%20the')
  114. AnnotationURLCitation(end_index=44989, start_index=44832, title='Targeting WDxR motif reprograms immune microenvironment and inhibits hepatocellular carcinoma progression - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC10165360/#:~:text=promotes%20its%20degradation,against%20HCC%20with%20WDR6%20dysregulation')
  115. AnnotationURLCitation(end_index=45291, start_index=45148, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=containing%20protein%206%20,for%20the%20treatment%20of%20hepatic')
  116. AnnotationURLCitation(end_index=45432, start_index=45292, title='Upregulation of WDR6 drives hepatic de novo lipogenesis in insulin resistance in mice | Nature Metabolism', type='url_citation', url='https://www.nature.com/articles/s42255-023-00896-7#:~:text=phosphatase%201%20,the%20treatment%20of%20hepatic%20steatosis')
  117. AnnotationURLCitation(end_index=45747, start_index=45590, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=Our%20results%20showed%20WDR6%20was,and%20chemokines%20in%20lung%20cancer')
  118. AnnotationURLCitation(end_index=45906, start_index=45748, title='WD repeat domain 6 as a novelty prognostic biomarker correlates with immune infiltration in lung cancer: A preliminary study - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC9382870/#:~:text=found%20the%20expression%20of%20WDR6,and%20chemokines%20in%20lung%20cancer')
  119. AnnotationURLCitation(end_index=46256, start_index=46064, title='Molecular cloning, expression analysis, and chromosome mapping of WDR6, a novel human WD-repeat gene - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/10903905/#:~:text=transduction%2C%20transcription%2C%20and%20proliferation,significant%20sequence%20similarity%20with%20Arabidopsis')
  120. AnnotationURLCitation(end_index=46404, start_index=46257, title='Molecular cloning, expression analysis, and chromosome mapping of WDR6, a novel human WD-repeat gene - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/10903905/#:~:text=since%20its%2011%20WD%20repeats,probably%20belongs%20to%20a%20highly')