nhr-47

UniProt ID: Q17370
Organism: Caenorhabditis elegans
Review Status: COMPLETE
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Gene Description

nhr-47 (C24G6.4) is one of the ~280 nuclear hormone receptors (NHRs) of Caenorhabditis elegans, the great majority of which arose by a nematode-specific expansion of an ancestral HNF4-like orphan receptor and are termed supplementary nuclear receptors (supnrs). The 579-residue protein has the canonical nuclear-receptor architecture: an N-terminal DNA-binding domain built from two C4-type (Cys4) zinc-finger motifs that coordinate zinc, and a C-terminal ligand-binding domain. On the basis of this diagnostic domain organization and phylogenetic placement, NHR-47 is a zinc-dependent, sequence-specific DNA-binding transcription factor of the nuclear-receptor superfamily that acts in the nucleus to regulate RNA polymerase II transcription. It is annotated as an orphan receptor: no endogenous ligand has been identified. NHR-47 is expressed in head and tail neurons, the ventral nerve cord, the spermatheca, the pharynx and the intestine, and it physically interacts with another nuclear receptor, NHR-17. The only reported loss-of-function phenotypes are in the germline: knockdown modulates susceptibility to environmental toxicants (polystyrene nanoparticles and 6-PPD-quinone), where NHR-47 acts as a positive mediator of reproductive and transgenerational toxicity and lies upstream of insulin, Ephrin and Wnt ligand-gene expression. nhr-47 transcription is also induced by exposure to the vertebrate steroid estradiol. Its endogenous physiological role, its ligand (if any), and its direct transcriptional targets remain undetermined.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005634 nucleus
IBA
GO_REF:0000033
ACCEPT
Summary: NHR-47 is a nuclear-receptor transcription factor and acts in the nucleus. Supported by the canonical NR DNA-binding domain and UniProt subcellular location.
Reason: Nuclear localization is expected for a functional nuclear-receptor transcription factor and is independently annotated from UniProt SubCell mapping. This is a core cellular-component assignment.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IBA
GO_REF:0000033
ACCEPT
Summary: Sequence-specific DNA binding at Pol II cis-regulatory regions is the diagnostic molecular function of the NR DNA-binding domain (two C4 zinc fingers) that NHR-47 possesses.
Reason: Consistent with the conserved NR DBD architecture and phylogenetic placement in the HNF4-like DNA-binding-domain class. A core molecular function.
GO:0004879 nuclear receptor activity
IBA
GO_REF:0000033
ACCEPT
Summary: NHR-47 is a member of the nuclear-receptor superfamily with a canonical DBD and ligand-binding domain; nuclear receptor activity (ligand-modulated sequence-specific DNA-binding TF activity) is its family-level molecular function.
Reason: Well supported by domain architecture and family membership. Note that NHR-47 is an orphan receptor: no ligand is known, so the ligand-modulated aspect of this term is inferred, not demonstrated (see knowledge_gaps). Retained as a core MF.
GO:0006357 regulation of transcription by RNA polymerase II
IBA
GO_REF:0000033
ACCEPT
Summary: As a sequence-specific DNA-binding TF of the NR family, NHR-47 regulates Pol II transcription. This is the core biological process for the molecular function.
Reason: Directly follows from the nuclear-receptor / DNA-binding-TF molecular function and nuclear localization. Core process.
GO:0030154 cell differentiation
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: A generic biological process propagated across the nuclear-receptor family by phylogenetic inference. No nhr-47-specific evidence links it to a differentiation program.
Reason: This is a broad pan-family IBA propagation. NHR-47 has no reported role in cell differentiation; its only experimental phenotypes are germline toxicant responses. Retain (family inference is not contradicted) but mark non-core rather than core.
GO:0000978 RNA polymerase II cis-regulatory region sequence-specific DNA binding
IEA
GO_REF:0000002
ACCEPT
Summary: InterPro2GO assignment (IPR049636, HNF4-like DBD) of the same sequence-specific Pol II DNA-binding function, in agreement with the IBA annotation above.
Reason: Independent InterPro domain-based support for the core sequence-specific DNA-binding molecular function. Consistent with the phylogenetic annotation.
GO:0003700 DNA-binding transcription factor activity
IEA
GO_REF:0000002
ACCEPT
Summary: General DNA-binding transcription factor activity assigned from the nuclear-receptor zinc-finger domain (IPR001628). NHR-47 is a bona fide DNA-binding TF.
Reason: Correct parent molecular function for a nuclear-receptor TF; supported by the C4 zinc-finger DBD. Core MF (the RNA Pol II-specific children above are the more precise forms).
GO:0005634 nucleus
IEA
GO_REF:0000044
ACCEPT
Summary: Nuclear localization from UniProt Swiss-Prot subcellular-location mapping (SL-0191), consistent with the IBA nucleus annotation.
Reason: Independent support for nuclear localization of this transcription factor. Core CC.
GO:0006355 regulation of DNA-templated transcription
IEA
GO_REF:0000002
KEEP AS NON CORE
Summary: General (DNA-templated) transcription-regulation term from InterPro. This is the broad parent of the RNA Pol II-specific regulation term.
Reason: Not wrong, but less informative than the RNA polymerase II-specific regulation term (GO:0006357) already accepted as core. Retain as a non-core general annotation.
GO:0008270 zinc ion binding
IEA
GO_REF:0000002
ACCEPT
Summary: The two C4-type (Cys4) zinc-finger motifs of the NR DNA-binding domain coordinate zinc; zinc binding is a structural molecular function underpinning DNA binding.
Reason: Directly supported by the diagnostic C4 zinc-finger DBD (ZN_FING 11-31 and 47-71 in UniProt Q17370). Structural MF that enables the core DNA-binding function.
GO:0030522 intracellular receptor signaling pathway
IEA
GO_REF:0000108
KEEP AS NON CORE
Summary: Auto-inferred by inter-ontology logic from nuclear receptor activity (GO:0004879). For an orphan receptor with no known ligand, the signaling-pathway aspect is unproven.
Reason: A logically-derived (GO_REF:0000108) consequence of the nuclear-receptor MF, not direct evidence. Because NHR-47 is an orphan receptor with no demonstrated ligand-activated signaling, retain as non-core rather than core.
GO:0043565 sequence-specific DNA binding
IEA
GO_REF:0000002
ACCEPT
Summary: Sequence-specific DNA binding assigned from the NR zinc-finger domain (IPR001628); the general parent of the Pol II cis-regulatory DNA-binding term.
Reason: Supported by the conserved C4 zinc-finger DBD. Core MF (parent of GO:0000978).
GO:0005515 protein binding
IPI
PMID:19123269
Empirically controlled mapping of the Caenorhabditis elegans...
KEEP AS NON CORE
Summary: Experimental yeast two-hybrid interaction captured in the worm interactome; the partner is the nuclear receptor NHR-17 (UniProtKB:Q17589). "Protein binding" is an uninformative molecular-function term.
Reason: A real, curated physical interaction (NHR-47<->NHR-17), so it should not be removed. However GO:0005515 conveys no specific molecular function, and a single Y2H edge does not justify a more informative MF (e.g. nuclear receptor binding) as a core function. Retain as non-core.
GO:0005515 protein binding
IPI
PMID:23791784
Extensive rewiring and complex evolutionary dynamics in a C....
KEEP AS NON CORE
Summary: Second experimental interaction annotation (C. elegans multiparameter TF network), again with NHR-17 (UniProtKB:Q17589). Uninformative "protein binding" term.
Reason: Corroborates the NHR-47<->NHR-17 interaction from an independent dataset. Keep (real interaction) but non-core; GO:0005515 does not describe a specific molecular function.

Core Functions

Sequence-specific DNA-binding transcription factor of the nuclear-receptor superfamily. NHR-47 carries the diagnostic nuclear-receptor DNA-binding domain formed by two C4-type (Cys4) zinc-finger motifs that coordinate zinc, together with a C-terminal ligand-binding domain. Acting in the nucleus, it binds RNA polymerase II cis-regulatory regions in a sequence-specific manner to regulate transcription. It is an orphan receptor (no ligand identified), belonging to the HNF4-derived supplementary nuclear receptor class of C. elegans.

Molecular Function:
nuclear receptor activity
Cellular Locations:
Supporting Evidence:

Zinc-dependent, sequence-specific DNA binding. The two C4 zinc fingers of the NR DNA-binding domain coordinate zinc and mediate recognition of specific DNA response elements, providing the biochemical basis for NHR-47 transcription-factor activity.

Cellular Locations:
Supporting Evidence:

References

Gene Ontology annotation through association of InterPro records with GO terms
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Automatic assignment of GO terms using logical inference, based on on inter-ontology links
Empirically controlled mapping of the Caenorhabditis elegans protein-protein interactome network.
Extensive rewiring and complex evolutionary dynamics in a C. elegans multiparameter transcription factor network.
Explosive lineage-specific expansion of the orphan nuclear receptor HNF4 in nematodes.
Transgenerational Response of Germline Nuclear Hormone Receptor Genes to Nanoplastics at Predicted Environmental Doses in Caenorhabditis elegans.
6-PPD quinone induces response of nuclear hormone receptors in the germline associated with formation of reproductive toxicity in Caenorhabditis elegans.
Changes in Nuclear Receptor and Vitellogenin Gene Expression in Response to Steroids and Heavy Metal in Caenorhabditis elegans.

Suggested Questions for Experts

Q: Is NHR-47 a ligand-regulated receptor, and if so what endogenous metabolite or lipid binds its ligand-binding domain, or is it a constitutive/orphan transcriptional regulator?

Suggested experts: Nuclear-receptor biochemists, C. elegans NHR-family biologists

Q: What are the direct genomic binding sites and target genes of NHR-47, and does it act as a transcriptional activator or repressor?

Suggested experts: Transcription / gene-regulatory-network biologists

Q: What is the endogenous (non-toxicant) physiological role of nhr-47, given its expression in neurons, the ventral nerve cord, spermatheca, pharynx and intestine and its germline role in toxicant responses?

Suggested experts: C. elegans developmental / germline biologists

Suggested Experiments

Experiment: Determine the direct NHR-47 regulon by ChIP-seq or CUT&RUN with an endogenously tagged NHR-47, combined with RNA-seq of a null mutant, to identify direct target genes and the DNA response element and to establish activator vs repressor behavior.

Type: Genome-wide binding assay

Experiment: Express and purify the NHR-47 ligand-binding domain and screen for bound endogenous lipids/metabolites (e.g. by lipidomics of co-purified ligands or reporter-based ligand-sensing assays) to test whether NHR-47 is ligand-regulated or a true orphan.

Type: Ligand identification

Experiment: Generate and characterize a null allele (CRISPR) across development, metabolism, reproduction, stress and innate-immune assays to define the baseline endogenous function, and test genetic interaction with the interacting receptor nhr-17.

Type: Loss-of-function phenotyping

Knowledge Gaps

What is not known โ€” curated, literature-grounded statements of the open unknowns (the inverse of core functions).

Gap: Whether NHR-47 binds an endogenous small-molecule ligand โ€” and, if so, its chemical identity โ€” is unknown. It is classified as an orphan receptor, and as a member of the HNF4-derived supplementary nuclear receptors it may have lost or substantially altered ligand-dependent regulation.

OPEN BIOLOGY MF_DARK

What is known: NHR-47 has a canonical nuclear-receptor ligand-binding domain (UniProt NR LBD, residues 164-553), but UniProt annotates it as an orphan nuclear receptor and no ligand has been reported. nhr-47 transcription is induced by estradiol exposure, but transcriptional induction is not evidence that estradiol (or any specific molecule) is a direct NHR-47 ligand.

Significance: Knowing whether NHR-47 is ligand-regulated (and by what) would determine whether it is a sensor of a metabolite/hormone or a constitutive/orphan transcriptional regulator, and would shape any pharmacological or genetic strategy to control its activity.

Provenance (the field's own admissions):

Gap: The direct transcriptional targets and the DNA response element bound by NHR-47 in vivo are undetermined, and it is unknown whether NHR-47 acts as an activator or a repressor. No ChIP, reporter, or motif data define its regulon; the downstream genes that change on nhr-47 knockdown in toxicant assays (e.g. ins-3, daf-28, efn-3) are whole-organism readouts, not demonstrated direct targets.

OPEN BIOLOGYCURATION MF_DARK

What is known: NHR-47 is a sequence-specific DNA-binding transcription factor and participates in the C. elegans transcription-factor interaction network, and knockdown alters downstream secreted-ligand gene expression, but no bona fide direct target has been established.

Significance: Identifying direct targets and the response element would convert NHR-47 from a predicted TF into a defined regulator and reveal the pathway(s) it controls.

Provenance (the field's own admissions):

Gap: The endogenous physiological role and definitive loss-of-function phenotype of nhr-47 are unresolved. Its only reported phenotypes are germline RNAi effects that modulate susceptibility to environmental toxicants (polystyrene nanoparticles and 6-PPD-quinone); no baseline developmental, metabolic, or immune phenotype, and no characterized null mutant, has been reported.

OPEN BIOLOGY BP_DARK

What is known: nhr-47 is expressed in neurons, ventral nerve cord, spermatheca, pharynx and intestine, and germline knockdown modulates transgenerational/reproductive toxicant responses, but a stand-alone required function under normal conditions has not been defined.

Significance: A defined loss-of-function phenotype would anchor NHR-47's biological process annotation, which is currently only family-propagated (IBA) or inferred from a toxicant-stress context.

Provenance (the field's own admissions):

Deep Research

Falcon

(nhr-47-deep-research-falcon.md)
Comprehensive Research Report: NHR-47 (C24G6.4) in *Caenorhabditis elegans* Falcon Edison Scientific Literature 22 citations 1 artifacts 2026-07-04T17:24:09.725428

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Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.

Comprehensive Research Report: NHR-47 (C24G6.4) in Caenorhabditis elegans

Gene Identity and Nomenclature

NHR-47 (UniProt: Q17370) is a nuclear hormone receptor encoded by the gene nhr-47 (systematic/ORF name C24G6.4) on chromosome II of Caenorhabditis elegans. The gene carries an older synonym, csr-1, which must not be confused with the well-characterized CSR-1 Argonaute protein involved in small RNA pathwaysโ€”a completely distinct gene product. NHR-47 contains an HNF4-like DNA-binding domain (IPR049636) and a nuclear hormone receptor ligand-binding domain (IPR000536), consistent with its classification as a member of the nuclear hormone receptor superfamily (Nuclear_hrmn_rcpt, IPR001723; Nuclear_hormone_rcpt_NR2, IPR050274).

The following table summarizes verified properties and available experimental data for NHR-47:

Category NHR-47 summary
Gene / protein identifiers Gene name: nhr-47; Systematic name / ORF: C24G6.4; UniProt accession: Q17370; Reported synonym: csr-1 (must not be confused with the unrelated CSR-1 Argonaute gene) (novillo2005changesinnuclear pages 3-4)
Organism Caenorhabditis elegans (novillo2005changesinnuclear pages 3-4)
Protein family Nuclear hormone receptor (NHR); HNF4-like supplementary NHR within the nematode-expanded receptor family (taubert2011nuclearhormonereceptors pages 2-4, taubert2011nuclearhormonereceptors pages 1-2)
Core domains HNF4-like DNA-binding domain and nuclear receptor ligand-binding domain; domain architecture is consistent with a canonical ligand-regulated transcription factor in the NHR superfamily (taubert2011nuclearhormonereceptors pages 2-4, taubert2011nuclearhormonereceptors pages 1-2)
Predicted molecular function Putative ligand-dependent transcription factor / transcriptional regulator responding to lipophilic small molecules, inferred from its NHR family membership and HNF4-like architecture (taubert2011nuclearhormonereceptors pages 2-4, taubert2011nuclearhormonereceptors pages 1-2)
Direct experimental evidence: steroid responsiveness Estradiol-induced upregulation: 10 ฮผM estradiol increased nhr-47 expression 3.4-fold in microarray analysis, identifying it as an estradiol-responsive NHR (novillo2005changesinnuclear pages 3-4)
Direct experimental evidence: C. elegans phenotype Published RNAi experiments in C. elegans reported no observable phenotype for nhr-47, indicating either redundancy, condition-specific function, or weak effect under standard assay conditions (reported in Ciche et al. summarizing prior RNAi studies) (ciche2007postembryonicrnaiin pages 6-8)
Cross-species ortholog evidence RNAi of the Heterorhabditis bacteriophora ortholog Hba-nhr-47 usually caused no obvious phenotype, with only 3โ€“12% sterility in some trials, supporting a subtle or context-dependent role (ciche2007postembryonicrnaiin pages 4-6, ciche2007postembryonicrnaiin pages 6-8)
Putative biological processes Likely part of an estrogen-sensitive gene network associated with vitellogenesis, lipid availability, and broader metabolic/endocrine responses; however, no direct downstream targets have been established for nhr-47 itself (novillo2005changesinnuclear pages 3-4)
Predicted subcellular localization Nucleus (inferred), because NHR-47 contains an HNF4-like DNA-binding domain and belongs to a family of nuclear transcription factors; no nhr-47-specific localization study was found (taubert2011nuclearhormonereceptors pages 2-4, taubert2011nuclearhormonereceptors pages 1-2)
Evolutionary context Member of the nematode-specific expansion of HNF4-related receptors: C. elegans has 284 NHRs overall, including 269 supplementary NHRs (supnrs) derived from an ancient HNF4-like ancestor; this expansion is far larger than in humans or flies (arda2010functionalmodularityof pages 2-3, taubert2011nuclearhormonereceptors pages 2-4, taubert2011nuclearhormonereceptors pages 1-2, arda2010functionalmodularityof pages 9-10)
Broader functional context of family HNF4-like C. elegans NHRs commonly regulate metabolism, fat storage/catabolism, xenobiotic responses, and physiological adaptation, often in modular gene-regulatory networks; nhr-47 is therefore plausibly metabolic/endocrine, though not directly characterized (arda2010functionalmodularityof pages 2-3, taubert2011nuclearhormonereceptors pages 2-4, arda2010functionalmodularityof pages 3-4)
Characterization status Poorly characterized / orphan receptor. The literature provides direct evidence for steroid-responsive expression but little mechanistic information on ligands, tissue expression, target genes, or physiological necessity under standard conditions (novillo2005changesinnuclear pages 3-4, ciche2007postembryonicrnaiin pages 6-8)

Table: This table consolidates verified identifiers, family/domain assignment, and the limited direct experimental evidence available for C. elegans NHR-47. It is useful as a high-confidence quick reference because the gene is sparsely characterized and easily confused with unrelated genes sharing similar names.

Evolutionary Context: The Nematode-Specific HNF4 Expansion

To understand NHR-47, it is essential to place it within the broader evolutionary and functional context of the massively expanded nuclear hormone receptor family in C. elegans. While the human genome encodes 48 NHRs and Drosophila has 18, the C. elegans genome encodes approximately 284 NHR genesโ€”roughly six times the human complement (taubert2011nuclearhormonereceptors pages 1-2). Of these, only 15 are "conserved" NHRs belonging to five of the six metazoan NHR subfamilies (NR1, NR2, NR4, NR5, and NR6), while the remaining 269 are "supplementary NHRs" (supnrs) that appear to have derived from a single ancient HNF4-like ancestor through lineage-specific gene duplication events (taubert2011nuclearhormonereceptors pages 2-4). This expansion is one of the most dramatic in any metazoan transcription factor family, and the resulting supnrs show rapid evolutionary divergenceโ€”approximately 50% of NHRs in related nematode species such as C. briggsae and C. remanei are species-specific, indicating that NHRs evolve more rapidly than other transcription factor classes (arda2010functionalmodularityof pages 9-10).

NHR-47 belongs to this expanded HNF4-derived supplementary NHR class. Despite the large number of supnrs, only about 50 NHRs in C. elegans have known functions or detectable phenotypes when mutated or depleted (taubert2011nuclearhormonereceptors pages 1-2). NHR-47 is among the majority of these supplementary receptors that remain functionally uncharacterized.

Predicted Molecular Function

As a canonical nuclear hormone receptor, NHR-47 is predicted to function as a ligand-dependent transcription factor. Nuclear hormone receptors typically bind small lipophilic molecules (ligands) through their C-terminal ligand-binding domain, undergo conformational changes, and regulate transcription of target genes by binding specific DNA response elements through their N-terminal zinc-finger DNA-binding domain. For NHR-47 specifically, no endogenous ligand has been identified, classifying it as an orphan receptor.

The HNF4-like domain architecture of NHR-47 places it in the NR2A class of nuclear receptors. In vertebrates, the HNF4 receptors (HNF4ฮฑ and HNF4ฮณ) play central roles in cholesterol, amino acid, carbohydrate, lipid, and xenobiotic metabolism, as well as liver-specific gene expression (novillo2005changesinnuclear pages 4-5). In C. elegans, where the intestine performs many of the metabolic functions of the vertebrate liver, HNF4-type NHRs have been shown to regulate fat storage and catabolism, xenobiotic detoxification, and physiological adaptation to environmental and nutritional cues (arda2010functionalmodularityof pages 2-3, arda2010functionalmodularityof pages 3-4). By extension, NHR-47 is plausibly involved in metabolic regulation, though direct evidence for this remains lacking.

Known Experimental Evidence

Steroid Responsiveness

The most direct functional data for nhr-47 comes from a microarray study by Novillo et al. (2005), which examined changes in nuclear receptor gene expression in C. elegans exposed to vertebrate steroids and cholesterol. This study demonstrated that 10 ฮผM estradiol induced a 3.4-fold upregulation of nhr-47 expression (novillo2005changesinnuclear pages 3-4). Importantly, each steroid tested (progesterone, estradiol, cholesterol) activated or inhibited entirely distinct subsets of NR genes, with no overlap between treatments, and estradiol specifically regulated 11 of 25 NRs identified as steroid-responsive (novillo2005changesinnuclear pages 1-1). The authors suggested that estradiol-responsive genes, including nhr-47, may constitute an estrogen-sensitive gene network related to vitellogenesis, since vitellogenin production depends on lipid availability to gastrointestinal cells involved in yolk protein synthesis (novillo2005changesinnuclear pages 3-4). This is biologically plausible given that the C. elegans intestine is the primary site of vitellogenin synthesis and that NHRs are known to respond to cholesterol-derived lipophilic signals in nematodes (novillo2005changesinnuclear pages 1-3, novillo2005changesinnuclear pages 5-6).

RNAi Phenotype Analysis

Published RNAi experiments of nhr-47 in C. elegans have reported no observable phenotype under standard laboratory conditions (ciche2007postembryonicrnaiin pages 6-8). This finding was noted across multiple independent genome-wide RNAi screens. In the related insect-parasitic nematode Heterorhabditis bacteriophora, RNAi of the ortholog Hba-nhr-47 also generally resulted in no observable defect, although very low-penetrance sterility (3โ€“12%) was noted in some experimental trials (ciche2007postembryonicrnaiin pages 4-6, ciche2007postembryonicrnaiin pages 6-8). The absence of a strong loss-of-function phenotype is consistent with three possible explanations: (1) functional redundancy among the many HNF4-derived supplementary NHRs, (2) a context- or condition-dependent role that is not revealed under standard culture conditions, or (3) a subtle function below the detection threshold of typical phenotypic screens.

Evolutionary Sequence Relationships

A study of selfish toxin-antidote elements in Caenorhabditis species noted that the SLOW-1 protein, a toxin component in C. tropicalis that is homologous to nuclear hormone receptors, shares sequence similarity with C. elegans NHR-47 (17.9% identity) (bendavid2021ubiquitousselfishtoxinantidote pages 9-11). SLOW-1 retains an NHR ligand-binding domain but lacks the canonical DNA-binding domain and instead possesses transmembrane domains, suggesting it may function as a dominant-negative element that sequesters NHR ligands. While this does not directly inform NHR-47 function, it highlights the evolutionary versatility of the HNF4-derived NHR ligand-binding domain in nematodes and the potential for NHR-47-like proteins to interact with lipophilic signaling molecules.

Subcellular Localization

No direct localization study has been performed for NHR-47. However, as a nuclear hormone receptor containing a zinc-finger DNA-binding domain, NHR-47 is predicted to localize to the nucleus, where it would function as a transcription factor. Well-characterized family members such as NHR-86 have been confirmed to localize predominantly to the nuclei of intestinal, excretory gland, and neuronal cells (arda2010functionalmodularityof pages 6-7), and NHR-49 functions in the nucleus to regulate target gene expression (motola2006identificationofligands pages 39-43). By analogy, NHR-47 is expected to occupy a similar nuclear compartment in metabolically active tissues.

Broader Functional Context of the NHR Family

The metabolic gene regulatory network (GRN) of C. elegans was experimentally mapped by Arda et al. (2010) using yeast one-hybrid assays, revealing that NHRs are significantly enriched as regulators of metabolic gene promotersโ€”over one quarter of all transcription factors in the metabolic GRN are NHRs (arda2010functionalmodularityof pages 2-3). The metabolic GRN is highly modular, with approximately 60% of NHRs organized into two dominant modules that predominantly regulate genes involved in fat storage and catabolism (arda2010functionalmodularityof pages 3-4). RNAi of most NHRs in these modules resulted in increased lipid staining by Nile Red and Oil-Red-O, indicating roles in lipid metabolism (arda2010functionalmodularityof pages 3-4). These findings suggest that the expansion of the HNF4 family in nematodes has been functionally linked to metabolic regulation, with multiple supplementary NHRs sharing fat-regulatory roles after duplication and divergence.

Furthermore, the Mediator subunit MDT-15 has been identified as a key cofactor that preferentially interacts with metabolic NHRs, and several NHRs form regulatory cascades in which one NHR controls the expression of another (arda2010functionalmodularityof pages 5-6). These inter-NHR regulatory circuits are proposed to enable rapid and adaptive responses to environmental and physiological changes such as nutrient availability (arda2010functionalmodularityof pages 9-10).

Limitations and Current Status

NHR-47 remains a poorly characterized orphan nuclear hormone receptor. The available evidence establishes it as an estradiol-responsive member of the massively expanded HNF4-derived supplementary NHR family in C. elegans, but no endogenous ligand, specific target genes, tissue-specific expression pattern, or robust loss-of-function phenotype has been reported. Its function may be obscured by redundancy with other supplementary NHRs, or it may act under specific environmental, nutritional, or developmental conditions not routinely tested in laboratory screens. The protein's function can be inferred from its domain and family architecture: NHR-47 is most likely a ligand-dependent nuclear transcription factor involved in metabolic gene regulation, potentially participating in steroid/lipid-responsive signaling pathways related to vitellogenesis and lipid homeostasis. Future studies employing targeted knockouts, condition-specific screens (e.g., steroid exposure, dietary stress), and tissue-specific expression reporters will be needed to elucidate the precise biological role of this receptor.

References

  1. (novillo2005changesinnuclear pages 3-4): A. Novillo, S. Won, Christine Li, and I. Callard. Changes in nuclear receptor and vitellogenin gene expression in response to steroids and heavy metal in caenorhabditis elegans1. Integrative and Comparative Biology, 45:61-71, Jan 2005. URL: https://doi.org/10.1093/icb/45.1.61, doi:10.1093/icb/45.1.61. This article has 59 citations and is from a peer-reviewed journal.

  2. (taubert2011nuclearhormonereceptors pages 2-4): Stefan Taubert, Jordan D. Ward, and Keith R. Yamamoto. Nuclear hormone receptors in nematodes: evolution and function. Molecular and Cellular Endocrinology, 334:49-55, Mar 2011. URL: https://doi.org/10.1016/j.mce.2010.04.021, doi:10.1016/j.mce.2010.04.021. This article has 136 citations and is from a peer-reviewed journal.

  3. (taubert2011nuclearhormonereceptors pages 1-2): Stefan Taubert, Jordan D. Ward, and Keith R. Yamamoto. Nuclear hormone receptors in nematodes: evolution and function. Molecular and Cellular Endocrinology, 334:49-55, Mar 2011. URL: https://doi.org/10.1016/j.mce.2010.04.021, doi:10.1016/j.mce.2010.04.021. This article has 136 citations and is from a peer-reviewed journal.

  4. (ciche2007postembryonicrnaiin pages 6-8): Todd A Ciche and Paul W Sternberg. Postembryonic rnai in heterorhabditis bacteriophora: a nematode insect parasite and host for insect pathogenic symbionts. BMC Developmental Biology, 7:101-101, Sep 2007. URL: https://doi.org/10.1186/1471-213x-7-101, doi:10.1186/1471-213x-7-101. This article has 78 citations and is from a peer-reviewed journal.

  5. (ciche2007postembryonicrnaiin pages 4-6): Todd A Ciche and Paul W Sternberg. Postembryonic rnai in heterorhabditis bacteriophora: a nematode insect parasite and host for insect pathogenic symbionts. BMC Developmental Biology, 7:101-101, Sep 2007. URL: https://doi.org/10.1186/1471-213x-7-101, doi:10.1186/1471-213x-7-101. This article has 78 citations and is from a peer-reviewed journal.

  6. (arda2010functionalmodularityof pages 2-3): H Efsun Arda, Stefan Taubert, Lesley T MacNeil, Colin C Conine, Ben Tsuda, Marc Van Gilst, Reynaldo Sequerra, Lynn Doucetteโ€Stamm, Keith R Yamamoto, and Albertha J M Walhout. Functional modularity of nuclear hormone receptors in a caenorhabditis elegans metabolic gene regulatory network. Molecular Systems Biology, 6:367-367, May 2010. URL: https://doi.org/10.1038/msb.2010.23, doi:10.1038/msb.2010.23. This article has 148 citations and is from a highest quality peer-reviewed journal.

  7. (arda2010functionalmodularityof pages 9-10): H Efsun Arda, Stefan Taubert, Lesley T MacNeil, Colin C Conine, Ben Tsuda, Marc Van Gilst, Reynaldo Sequerra, Lynn Doucetteโ€Stamm, Keith R Yamamoto, and Albertha J M Walhout. Functional modularity of nuclear hormone receptors in a caenorhabditis elegans metabolic gene regulatory network. Molecular Systems Biology, 6:367-367, May 2010. URL: https://doi.org/10.1038/msb.2010.23, doi:10.1038/msb.2010.23. This article has 148 citations and is from a highest quality peer-reviewed journal.

  8. (arda2010functionalmodularityof pages 3-4): H Efsun Arda, Stefan Taubert, Lesley T MacNeil, Colin C Conine, Ben Tsuda, Marc Van Gilst, Reynaldo Sequerra, Lynn Doucetteโ€Stamm, Keith R Yamamoto, and Albertha J M Walhout. Functional modularity of nuclear hormone receptors in a caenorhabditis elegans metabolic gene regulatory network. Molecular Systems Biology, 6:367-367, May 2010. URL: https://doi.org/10.1038/msb.2010.23, doi:10.1038/msb.2010.23. This article has 148 citations and is from a highest quality peer-reviewed journal.

  9. (novillo2005changesinnuclear pages 4-5): A. Novillo, S. Won, Christine Li, and I. Callard. Changes in nuclear receptor and vitellogenin gene expression in response to steroids and heavy metal in caenorhabditis elegans1. Integrative and Comparative Biology, 45:61-71, Jan 2005. URL: https://doi.org/10.1093/icb/45.1.61, doi:10.1093/icb/45.1.61. This article has 59 citations and is from a peer-reviewed journal.

  10. (novillo2005changesinnuclear pages 1-1): A. Novillo, S. Won, Christine Li, and I. Callard. Changes in nuclear receptor and vitellogenin gene expression in response to steroids and heavy metal in caenorhabditis elegans1. Integrative and Comparative Biology, 45:61-71, Jan 2005. URL: https://doi.org/10.1093/icb/45.1.61, doi:10.1093/icb/45.1.61. This article has 59 citations and is from a peer-reviewed journal.

  11. (novillo2005changesinnuclear pages 1-3): A. Novillo, S. Won, Christine Li, and I. Callard. Changes in nuclear receptor and vitellogenin gene expression in response to steroids and heavy metal in caenorhabditis elegans1. Integrative and Comparative Biology, 45:61-71, Jan 2005. URL: https://doi.org/10.1093/icb/45.1.61, doi:10.1093/icb/45.1.61. This article has 59 citations and is from a peer-reviewed journal.

  12. (novillo2005changesinnuclear pages 5-6): A. Novillo, S. Won, Christine Li, and I. Callard. Changes in nuclear receptor and vitellogenin gene expression in response to steroids and heavy metal in caenorhabditis elegans1. Integrative and Comparative Biology, 45:61-71, Jan 2005. URL: https://doi.org/10.1093/icb/45.1.61, doi:10.1093/icb/45.1.61. This article has 59 citations and is from a peer-reviewed journal.

  13. (bendavid2021ubiquitousselfishtoxinantidote pages 9-11): Eyal Ben-David, Pinelopi Pliota, Sonya A. Widen, Alevtina Koreshova, Tzitziki Lemus-Vergara, Philipp Verpukhovskiy, Sridhar Mandali, Christian Braendle, Alejandro Burga, and Leonid Kruglyak. Ubiquitous selfish toxin-antidote elements in caenorhabditis species. Current Biology, 31:990-1001.e5, Mar 2021. URL: https://doi.org/10.1016/j.cub.2020.12.013, doi:10.1016/j.cub.2020.12.013. This article has 61 citations and is from a highest quality peer-reviewed journal.

  14. (arda2010functionalmodularityof pages 6-7): H Efsun Arda, Stefan Taubert, Lesley T MacNeil, Colin C Conine, Ben Tsuda, Marc Van Gilst, Reynaldo Sequerra, Lynn Doucetteโ€Stamm, Keith R Yamamoto, and Albertha J M Walhout. Functional modularity of nuclear hormone receptors in a caenorhabditis elegans metabolic gene regulatory network. Molecular Systems Biology, 6:367-367, May 2010. URL: https://doi.org/10.1038/msb.2010.23, doi:10.1038/msb.2010.23. This article has 148 citations and is from a highest quality peer-reviewed journal.

  15. (motola2006identificationofligands pages 39-43): Daniel L. Motola, Carolyn L. Cummins, Veerle Rottiers, Kamalesh K. Sharma, Tingting Li, Yong Li, Kelly Suino-Powell, H. Eric Xu, Richard J. Auchus, Adam Antebi, and David J. Mangelsdorf. Identification of ligands for daf-12 that govern dauer formation and reproduction in c. elegans. Cell, 124:1209-1223, Mar 2006. URL: https://doi.org/10.1016/j.cell.2006.01.037, doi:10.1016/j.cell.2006.01.037. This article has 590 citations and is from a highest quality peer-reviewed journal.

  16. (arda2010functionalmodularityof pages 5-6): H Efsun Arda, Stefan Taubert, Lesley T MacNeil, Colin C Conine, Ben Tsuda, Marc Van Gilst, Reynaldo Sequerra, Lynn Doucetteโ€Stamm, Keith R Yamamoto, and Albertha J M Walhout. Functional modularity of nuclear hormone receptors in a caenorhabditis elegans metabolic gene regulatory network. Molecular Systems Biology, 6:367-367, May 2010. URL: https://doi.org/10.1038/msb.2010.23, doi:10.1038/msb.2010.23. This article has 148 citations and is from a highest quality peer-reviewed journal.

Artifacts

Citations

  1. novillo2005changesinnuclear pages 3-4
  2. ciche2007postembryonicrnaiin pages 6-8
  3. taubert2011nuclearhormonereceptors pages 1-2
  4. taubert2011nuclearhormonereceptors pages 2-4
  5. arda2010functionalmodularityof pages 9-10
  6. novillo2005changesinnuclear pages 4-5
  7. novillo2005changesinnuclear pages 1-1
  8. bendavid2021ubiquitousselfishtoxinantidote pages 9-11
  9. arda2010functionalmodularityof pages 6-7
  10. motola2006identificationofligands pages 39-43
  11. arda2010functionalmodularityof pages 2-3
  12. arda2010functionalmodularityof pages 3-4
  13. arda2010functionalmodularityof pages 5-6
  14. ciche2007postembryonicrnaiin pages 4-6
  15. novillo2005changesinnuclear pages 1-3
  16. novillo2005changesinnuclear pages 5-6
  17. https://doi.org/10.1093/icb/45.1.61,
  18. https://doi.org/10.1016/j.mce.2010.04.021,
  19. https://doi.org/10.1186/1471-213x-7-101,
  20. https://doi.org/10.1038/msb.2010.23,
  21. https://doi.org/10.1016/j.cub.2020.12.013,
  22. https://doi.org/10.1016/j.cell.2006.01.037,

๐Ÿ“š Additional Documentation

Notes

(nhr-47-notes.md)

nhr-47 (C. elegans) โ€” research notes

Gene: nhr-47 / sequence name C24G6.4 / WormBase WBGene00003637 / UniProt Q17370 (NHR47_CAEEL).
Locus: Chromosome V. Protein: 579 aa nuclear hormone receptor.

Provenance conventions

Inline provenance uses [PMID:xxxxx "verbatim quote"]. WormBase/Alliance-curated statements
(no direct primary PMID to hand) are marked [WormBase/AGR] and are NOT used as verbatim
supporting_text in the review; only cached-publication PMID quotes are used for that.

Summary: nhr-47 is a "dark" HNF4-derived orphan nuclear receptor

nhr-47 is one of the ~280 nuclear hormone receptors (NHRs) of C. elegans, the great majority of
which arose by a nematode-specific expansion of an ancestral HNF4-like orphan receptor and remain
functionally uncharacterized ("supplementary nuclear receptors", supnrs)
PMID:15983867. Its DNA-binding domain is HNF4-like (CDD cd06960 NR_DBD_HNF4A;
InterPro IPR049636 HNF4-like_DBD).

Molecular architecture (KNOWN, from sequence/UniProt Q17370)

  • Two C4-type (Cys4) zinc-finger motifs (ZN_FING 11โ€“31 and 47โ€“71) forming a canonical nuclear-receptor
    DNA-binding domain (DNA_BIND 8โ€“83). Coordinates zinc (KW Metal-binding, Zinc, Zinc-finger).
  • A nuclear-receptor ligand-binding domain (NR LBD, residues 164โ€“553; PROSITE PS51843).
  • UniProt FUNCTION: "Orphan nuclear receptor." SUBCELLULAR LOCATION: Nucleus.
  • Documented physical interaction (IntAct EBI-2412077/EBI-2412071): Q17370 (nhr-47) <-> Q17589 (nhr-17),
    NbExp=3. nhr-17 is itself another nuclear hormone receptor.
  • Therefore, from sequence + family, nhr-47 is confidently a zinc-finger, sequence-specific
    DNA-binding transcription factor of the nuclear-receptor superfamily that acts in the nucleus.
    (This is what the IBA/IEA GO annotations capture.)

Expression (KNOWN, curated) [WormBase/AGR]

Alliance/WormBase automated + MOD-provided descriptions for WBGene00003637:
- Expressed in head neurons, tail neurons, ventral nerve cord, spermatheca, intestine, and pharynx.
- "gene expression of nhr-47 appears to be induced upon exposure of worm cultures to estradiol"
(WormBase MOD-provided gene description; primary microarray reference not located in cache โ€” recorded
here as curated context only, not used as review supporting_text).

Functional data (the only experimental phenotypes reported for nhr-47)

Two toxicology studies from one group (Southeast University / Dayong Wang lab) are the only primary
papers that assign nhr-47 a phenotype, both in the germline and in an environmental-toxicant
context:

  1. Nanoplastics, transgenerational toxicity PMID:38922100:
  2. nhr-47 is one of only 4 of 33 germline-expressed NHRs whose expression responds to polystyrene
    nanoparticles; it is UP-regulated
    PMID:38922100.
  3. Germline RNAi of nhr-47 makes animals RESISTANT to transgenerational nanoplastic toxicity
    PMID:38922100.
  4. Under exposure, nhr-47 RNAi changes downstream secreted-ligand gene expression
    PMID:38922100
    (ins-3/ins-39/daf-28 = insulin ligands; efn-3 = Ephrin) โ€” i.e. nhr-47 sits upstream of insulin/Ephrin
    signalling in this stress context. These are downstream/indirect readouts, NOT demonstrated direct targets.

  5. 6-PPD-quinone, reproductive toxicity PMID:40482507:

  6. nhr-47 is among germline NHRs whose knockdown SUPPRESSES 6-PPDQ reproductive toxicity
    PMID:40482507.
  7. nhr-47 expression is modulated by DNA-damage-checkpoint and ferroptosis-related signals under 6-PPDQ.

Consistent theme: in the germline, nhr-47 acts (with daf-12/nhr-14) as a positive mediator of
toxicant-induced reproductive/transgenerational toxicity; knocking it down is protective.

Interaction / network context

  • Y2H interactome maps nhr-47 into the worm protein interaction network PMID:19123269 (WI8);
    the specific nhr-47<->nhr-17 interaction is recorded in IntAct/UniProt (not stated verbatim in the paper text).
  • nhr-47 is one node in the C. elegans multiparameter transcription-factor network PMID:23791784,
    a study of protein-DNA / protein-protein / TF-TF network rewiring across paralogs.
  • These two PMIDs underlie the two GOA IPI "protein binding" annotations (with_from UniProtKB:Q17589 = nhr-17).

KNOWN vs NOT-KNOWN (explicit)

KNOWN:
- Molecular identity: zinc-finger, sequence-specific DNA-binding transcription factor of the
nuclear-receptor (HNF4-derived supnr) family; nuclear localization; binds zinc.
- Physically interacts with nhr-17.
- Expressed in neurons, ventral nerve cord, spermatheca, pharynx, intestine.
- Germline knockdown modulates susceptibility to two environmental toxicants (nanoplastics, 6-PPDQ),
positioning it upstream of insulin/Ephrin/Wnt ligand-gene expression in that stress context.

NOT KNOWN (knowledge gaps):
- Ligand: whether nhr-47 has any endogenous small-molecule ligand, and its identity โ€” it is an orphan
receptor, and supnrs may have altered/lost ligand-dependence
PMID:15983867.
- Direct target genes / response element: no ChIP, reporter, or motif defines the nhr-47 regulon;
it is unknown whether it is an activator or repressor, and whether the toxicant-context downstream
genes (ins-3, daf-28, efn-3, โ€ฆ) are direct or indirect targets.
- Definitive loss-of-function role: no baseline null-mutant developmental/metabolic/immune phenotype;
the only phenotypes are germline RNAi effects in toxicant-challenge assays.

GO annotation review orientation

  • All 14 GOA annotations are IBA (phylogenetic), IEA (InterPro/UniProt), or IPI (interactome).
  • MF/CC core (nuclear receptor activity, RNA Pol II cis-regulatory sequence-specific DNA binding,
    sequence-specific DNA binding, DNA-binding TF activity, zinc ion binding, nucleus) is well supported
    by the diagnostic domain architecture โ†’ ACCEPT as core.
  • BP terms are family-propagated/inferred (regulation of transcription, intracellular receptor
    signaling) โ†’ KEEP; "cell differentiation" (IBA) has no nhr-47-specific support โ†’ KEEP_AS_NON_CORE
    (generic pan-NHR propagation).
  • Two IPI "protein binding" annotations (nhr-17): experimental but uninformative term; do not REMOVE
    (real interaction), KEEP_AS_NON_CORE and flag that a more informative MF term is not warranted from
    a single Y2H edge.

Falcon deep-research addendum (genes/worm/nhr-47/nhr-47-deep-research-falcon.md)

Genuine Edison "deep research" report (23-min run, 22 citations). Key points, each attributed
to the falcon-cited primary source (NOT independently full-text-verified here unless noted):

  • csr-1 synonym caution: the UniProt Synonyms=csr-1 for nhr-47 is an OLD locus synonym and
    must NOT be confused with the CSR-1 Argonaute (a distinct gene). Falcon flags this explicitly.
  • Estradiol responsiveness (the source of the WormBase estradiol statement): Novillo et al. 2005
    PMID:21676746 microarray of steroid-exposed C. elegans; falcon reports "10 ฮผM estradiol increased
    nhr-47 expression 3.4-fold". NOTE: the cached PubMed abstract of PMID:21676746 does NOT name nhr-47
    (the per-gene 3.4-fold value is in the full text / supplementary data and in WormBase curation), so I
    do NOT attach a verbatim nhr-47 supporting_text to this PMID; it is added to references as the
    estradiol-induction source with correctness VERIFIED and no gene-specific quote.
  • No robust RNAi phenotype: falcon reports genome-wide RNAi of nhr-47 in C. elegans gave "no
    observable phenotype" (via Ciche & Sternberg 2007, which also reports the H. bacteriophora ortholog
    Hba-nhr-47 RNAi is mostly phenotype-free, ~3-12% sterility in some trials). Consistent with knowledge
    gap #3 (no defined loss-of-function role); treated as supporting context, not review provenance,
    because the C. elegans nhr-47 no-phenotype claim is a secondary summary.
  • Family/metabolic context: Taubert, Ward & Yamamoto 2011 (NHRs in nematodes) and Arda et al. 2010
    (metabolic gene-regulatory network; NHRs enriched as metabolic-promoter regulators, modular, MDT-15
    cofactor). Places nhr-47 in the HNF4-derived supnr class plausibly linked to metabolism โ€” but nhr-47
    itself is uncharacterized in these.
  • Discrepancy: falcon states nhr-47 is on "chromosome II"; UniProt Q17370 says Chromosome V. Trust
    UniProt (Chromosome V). Did NOT propagate falcon's chromosome claim.

Net effect on review: falcon corroborates the orphan/dark framing, the estradiol angle, and the
absence of a robust loss-of-function phenotype; it did not surface any definitive ligand, direct
target, or physiological role. No core-function or knowledge-gap conclusion was changed by it.

๐Ÿ“„ View Raw YAML

id: Q17370
gene_symbol: nhr-47
product_type: PROTEIN
status: COMPLETE
taxon:
  id: NCBITaxon:6239
  label: Caenorhabditis elegans
description: >-
  nhr-47 (C24G6.4) is one of the ~280 nuclear hormone receptors (NHRs) of
  Caenorhabditis elegans, the great majority of which arose by a nematode-specific
  expansion of an ancestral HNF4-like orphan receptor and are termed supplementary
  nuclear receptors (supnrs). The 579-residue protein has the canonical nuclear-receptor
  architecture: an N-terminal DNA-binding domain built from two C4-type (Cys4)
  zinc-finger motifs that coordinate zinc, and a C-terminal ligand-binding domain.
  On the basis of this diagnostic domain organization and phylogenetic placement,
  NHR-47 is a zinc-dependent, sequence-specific DNA-binding transcription factor of
  the nuclear-receptor superfamily that acts in the nucleus to regulate RNA
  polymerase II transcription. It is annotated as an orphan receptor: no endogenous
  ligand has been identified. NHR-47 is expressed in head and tail neurons, the
  ventral nerve cord, the spermatheca, the pharynx and the intestine, and it
  physically interacts with another nuclear receptor, NHR-17. The only reported
  loss-of-function phenotypes are in the germline: knockdown modulates susceptibility
  to environmental toxicants (polystyrene nanoparticles and 6-PPD-quinone), where
  NHR-47 acts as a positive mediator of reproductive and transgenerational toxicity
  and lies upstream of insulin, Ephrin and Wnt ligand-gene expression. nhr-47
  transcription is also induced by exposure to the vertebrate steroid estradiol. Its
  endogenous physiological role, its ligand (if any), and its direct transcriptional
  targets remain undetermined.
references:
- id: GO_REF:0000002
  title: Gene Ontology annotation through association of InterPro records with GO
    terms
  findings: []
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
  findings: []
- id: GO_REF:0000044
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location
    vocabulary mapping, accompanied by conservative changes to GO terms applied by
    UniProt
  findings: []
- id: GO_REF:0000108
  title: Automatic assignment of GO terms using logical inference, based on on inter-ontology
    links
  findings: []
- id: PMID:19123269
  title: Empirically controlled mapping of the Caenorhabditis elegans protein-protein
    interactome network.
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      High-throughput yeast two-hybrid interactome (Worm Interactome / WI8). Underlies
      one of the two GOA IPI protein-binding annotations for nhr-47. The specific
      NHR-47<->NHR-17 edge is recorded in IntAct/UniProt (with_from UniProtKB:Q17589);
      the paper text itself reports the network methodology rather than naming nhr-47,
      so no gene-specific verbatim quote is available from the cached full text.
- id: PMID:23791784
  title: Extensive rewiring and complex evolutionary dynamics in a C. elegans multiparameter
    transcription factor network.
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      C. elegans multiparameter transcription-factor network (protein-DNA, protein-protein,
      TF-TF). Underlies the second GOA IPI protein-binding annotation for nhr-47 (again
      with NHR-17, UniProtKB:Q17589). Abstract-only in cache; the interaction is captured
      in IntAct rather than quoted verbatim in the available text.
- id: PMID:15983867
  title: Explosive lineage-specific expansion of the orphan nuclear receptor HNF4 in
    nematodes.
  findings: []
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: >-
      Establishes the phylogenetic context for nhr-47: the C. elegans supplementary
      nuclear receptors (supnrs), including the HNF4-like DBD class to which nhr-47
      belongs, arose from an explosive HNF4 duplication and are predominantly orphan
      receptors with uncertain ligand-dependence. Cited in the UniProt entry for Q17370.
- id: PMID:38922100
  title: Transgenerational Response of Germline Nuclear Hormone Receptor Genes to Nanoplastics
    at Predicted Environmental Doses in Caenorhabditis elegans.
  findings: []
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: >-
      Primary functional study naming nhr-47: germline-expressed, up-regulated by
      polystyrene nanoparticles, and germline RNAi confers resistance to transgenerational
      nanoplastic toxicity, with downstream effects on insulin/Ephrin ligand-gene
      expression. One of only two primary papers reporting an nhr-47 phenotype. Full text
      is MDPI-hosted and not downloadable by the validator, so supporting_text quotes are
      taken from the abstract only.
- id: PMID:40482507
  title: 6-PPD quinone induces response of nuclear hormone receptors in the germline
    associated with formation of reproductive toxicity in Caenorhabditis elegans.
  findings: []
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: >-
      Second primary functional study naming nhr-47: knockdown suppresses 6-PPD-quinone
      reproductive toxicity, and nhr-47 expression is modulated by DNA-damage-checkpoint
      and ferroptosis-related signals. Abstract-only in cache but the nhr-47 result is
      explicit in the abstract.
- id: PMID:21676746
  title: Changes in Nuclear Receptor and Vitellogenin Gene Expression in Response to
    Steroids and Heavy Metal in Caenorhabditis elegans.
  findings: []
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: >-
      DNA-microarray study of steroid-responsive nuclear-receptor gene expression in
      C. elegans; it is the primary source of the WormBase-curated statement that nhr-47
      is induced by estradiol (full text reports ~3.4-fold induction by 10 uM estradiol).
      The cached PubMed abstract does not name nhr-47 (the per-gene value is in the full
      text / supplementary data), so no gene-specific verbatim supporting_text is attached
      to this reference; it is cited as the estradiol-responsiveness source only.
existing_annotations:
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: is_active_in
  review:
    summary: >-
      NHR-47 is a nuclear-receptor transcription factor and acts in the nucleus.
      Supported by the canonical NR DNA-binding domain and UniProt subcellular location.
    action: ACCEPT
    reason: >-
      Nuclear localization is expected for a functional nuclear-receptor transcription
      factor and is independently annotated from UniProt SubCell mapping. This is a
      core cellular-component assignment.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: enables
  review:
    summary: >-
      Sequence-specific DNA binding at Pol II cis-regulatory regions is the diagnostic
      molecular function of the NR DNA-binding domain (two C4 zinc fingers) that NHR-47
      possesses.
    action: ACCEPT
    reason: >-
      Consistent with the conserved NR DBD architecture and phylogenetic placement in
      the HNF4-like DNA-binding-domain class. A core molecular function.
- term:
    id: GO:0004879
    label: nuclear receptor activity
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: enables
  review:
    summary: >-
      NHR-47 is a member of the nuclear-receptor superfamily with a canonical DBD and
      ligand-binding domain; nuclear receptor activity (ligand-modulated sequence-specific
      DNA-binding TF activity) is its family-level molecular function.
    action: ACCEPT
    reason: >-
      Well supported by domain architecture and family membership. Note that NHR-47 is
      an orphan receptor: no ligand is known, so the ligand-modulated aspect of this
      term is inferred, not demonstrated (see knowledge_gaps). Retained as a core MF.
- term:
    id: GO:0006357
    label: regulation of transcription by RNA polymerase II
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: >-
      As a sequence-specific DNA-binding TF of the NR family, NHR-47 regulates Pol II
      transcription. This is the core biological process for the molecular function.
    action: ACCEPT
    reason: >-
      Directly follows from the nuclear-receptor / DNA-binding-TF molecular function and
      nuclear localization. Core process.
- term:
    id: GO:0030154
    label: cell differentiation
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: >-
      A generic biological process propagated across the nuclear-receptor family by
      phylogenetic inference. No nhr-47-specific evidence links it to a differentiation
      program.
    action: KEEP_AS_NON_CORE
    reason: >-
      This is a broad pan-family IBA propagation. NHR-47 has no reported role in cell
      differentiation; its only experimental phenotypes are germline toxicant responses.
      Retain (family inference is not contradicted) but mark non-core rather than core.
- term:
    id: GO:0000978
    label: RNA polymerase II cis-regulatory region sequence-specific DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: >-
      InterPro2GO assignment (IPR049636, HNF4-like DBD) of the same sequence-specific
      Pol II DNA-binding function, in agreement with the IBA annotation above.
    action: ACCEPT
    reason: >-
      Independent InterPro domain-based support for the core sequence-specific DNA-binding
      molecular function. Consistent with the phylogenetic annotation.
- term:
    id: GO:0003700
    label: DNA-binding transcription factor activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: >-
      General DNA-binding transcription factor activity assigned from the nuclear-receptor
      zinc-finger domain (IPR001628). NHR-47 is a bona fide DNA-binding TF.
    action: ACCEPT
    reason: >-
      Correct parent molecular function for a nuclear-receptor TF; supported by the C4
      zinc-finger DBD. Core MF (the RNA Pol II-specific children above are the more precise
      forms).
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: >-
      Nuclear localization from UniProt Swiss-Prot subcellular-location mapping (SL-0191),
      consistent with the IBA nucleus annotation.
    action: ACCEPT
    reason: >-
      Independent support for nuclear localization of this transcription factor. Core CC.
- term:
    id: GO:0006355
    label: regulation of DNA-templated transcription
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: involved_in
  review:
    summary: >-
      General (DNA-templated) transcription-regulation term from InterPro. This is the
      broad parent of the RNA Pol II-specific regulation term.
    action: KEEP_AS_NON_CORE
    reason: >-
      Not wrong, but less informative than the RNA polymerase II-specific regulation term
      (GO:0006357) already accepted as core. Retain as a non-core general annotation.
- term:
    id: GO:0008270
    label: zinc ion binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: >-
      The two C4-type (Cys4) zinc-finger motifs of the NR DNA-binding domain coordinate
      zinc; zinc binding is a structural molecular function underpinning DNA binding.
    action: ACCEPT
    reason: >-
      Directly supported by the diagnostic C4 zinc-finger DBD (ZN_FING 11-31 and 47-71 in
      UniProt Q17370). Structural MF that enables the core DNA-binding function.
- term:
    id: GO:0030522
    label: intracellular receptor signaling pathway
  evidence_type: IEA
  original_reference_id: GO_REF:0000108
  qualifier: involved_in
  review:
    summary: >-
      Auto-inferred by inter-ontology logic from nuclear receptor activity (GO:0004879).
      For an orphan receptor with no known ligand, the signaling-pathway aspect is
      unproven.
    action: KEEP_AS_NON_CORE
    reason: >-
      A logically-derived (GO_REF:0000108) consequence of the nuclear-receptor MF, not
      direct evidence. Because NHR-47 is an orphan receptor with no demonstrated
      ligand-activated signaling, retain as non-core rather than core.
- term:
    id: GO:0043565
    label: sequence-specific DNA binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: >-
      Sequence-specific DNA binding assigned from the NR zinc-finger domain (IPR001628);
      the general parent of the Pol II cis-regulatory DNA-binding term.
    action: ACCEPT
    reason: >-
      Supported by the conserved C4 zinc-finger DBD. Core MF (parent of GO:0000978).
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:19123269
  qualifier: enables
  review:
    summary: >-
      Experimental yeast two-hybrid interaction captured in the worm interactome; the
      partner is the nuclear receptor NHR-17 (UniProtKB:Q17589). "Protein binding" is
      an uninformative molecular-function term.
    action: KEEP_AS_NON_CORE
    reason: >-
      A real, curated physical interaction (NHR-47<->NHR-17), so it should not be removed.
      However GO:0005515 conveys no specific molecular function, and a single Y2H edge does
      not justify a more informative MF (e.g. nuclear receptor binding) as a core function.
      Retain as non-core.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:23791784
  qualifier: enables
  review:
    summary: >-
      Second experimental interaction annotation (C. elegans multiparameter TF network),
      again with NHR-17 (UniProtKB:Q17589). Uninformative "protein binding" term.
    action: KEEP_AS_NON_CORE
    reason: >-
      Corroborates the NHR-47<->NHR-17 interaction from an independent dataset. Keep (real
      interaction) but non-core; GO:0005515 does not describe a specific molecular function.
core_functions:
- description: >-
    Sequence-specific DNA-binding transcription factor of the nuclear-receptor
    superfamily. NHR-47 carries the diagnostic nuclear-receptor DNA-binding domain
    formed by two C4-type (Cys4) zinc-finger motifs that coordinate zinc, together with
    a C-terminal ligand-binding domain. Acting in the nucleus, it binds RNA polymerase
    II cis-regulatory regions in a sequence-specific manner to regulate transcription.
    It is an orphan receptor (no ligand identified), belonging to the HNF4-derived
    supplementary nuclear receptor class of C. elegans.
  molecular_function:
    id: GO:0004879
    label: nuclear receptor activity
  directly_involved_in:
  - id: GO:0006357
    label: regulation of transcription by RNA polymerase II
  locations:
  - id: GO:0005634
    label: nucleus
  supported_by:
  - reference_id: PMID:15983867
- description: >-
    Zinc-dependent, sequence-specific DNA binding. The two C4 zinc fingers of the NR
    DNA-binding domain coordinate zinc and mediate recognition of specific DNA
    response elements, providing the biochemical basis for NHR-47 transcription-factor
    activity.
  molecular_function:
    id: GO:0043565
    label: sequence-specific DNA binding
  locations:
  - id: GO:0005634
    label: nucleus
  supported_by:
  - reference_id: PMID:15983867
knowledge_gaps:
- gap_statement: >-
    Whether NHR-47 binds an endogenous small-molecule ligand โ€” and, if so, its
    chemical identity โ€” is unknown. It is classified as an orphan receptor, and as a
    member of the HNF4-derived supplementary nuclear receptors it may have lost or
    substantially altered ligand-dependent regulation.
  boundary: >-
    NHR-47 has a canonical nuclear-receptor ligand-binding domain (UniProt NR LBD,
    residues 164-553), but UniProt annotates it as an orphan nuclear receptor and no
    ligand has been reported. nhr-47 transcription is induced by estradiol exposure, but
    transcriptional induction is not evidence that estradiol (or any specific molecule)
    is a direct NHR-47 ligand.
  gap_kind:
  - BIOLOGY
  dark_aspect: MF_DARK
  status: OPEN
  significance: >-
    Knowing whether NHR-47 is ligand-regulated (and by what) would determine whether it
    is a sensor of a metabolite/hormone or a constitutive/orphan transcriptional regulator,
    and would shape any pharmacological or genetic strategy to control its activity.
  provenance:
  - reference_id: PMID:15983867
    supporting_text: >-
      This origin has specific implications for the role of ligand binding in the
      function and evolution of the nematode supplementary nuclear receptors.
- gap_statement: >-
    The direct transcriptional targets and the DNA response element bound by NHR-47
    in vivo are undetermined, and it is unknown whether NHR-47 acts as an activator or a
    repressor. No ChIP, reporter, or motif data define its regulon; the downstream genes
    that change on nhr-47 knockdown in toxicant assays (e.g. ins-3, daf-28, efn-3) are
    whole-organism readouts, not demonstrated direct targets.
  boundary: >-
    NHR-47 is a sequence-specific DNA-binding transcription factor and participates in
    the C. elegans transcription-factor interaction network, and knockdown alters
    downstream secreted-ligand gene expression, but no bona fide direct target has been
    established.
  gap_kind:
  - BIOLOGY
  - CURATION
  dark_aspect: MF_DARK
  status: OPEN
  significance: >-
    Identifying direct targets and the response element would convert NHR-47 from a
    predicted TF into a defined regulator and reveal the pathway(s) it controls.
  provenance:
  - reference_id: PMID:38922100
    supporting_text: >-
      expressions of ins-3, daf-28, and ins-39 encoding insulin ligands, efn-3 encoding
      Ephrin ligand, and lin-44 encoding Wnt ligand, as well as expressions of their
      receptor genes (daf-2, vab-1, and/or mig-1), were dysregulated by the RNAi of
      daf-12, nhr-14, nhr-47, and nhr-12
- gap_statement: >-
    The endogenous physiological role and definitive loss-of-function phenotype of
    nhr-47 are unresolved. Its only reported phenotypes are germline RNAi effects that
    modulate susceptibility to environmental toxicants (polystyrene nanoparticles and
    6-PPD-quinone); no baseline developmental, metabolic, or immune phenotype, and no
    characterized null mutant, has been reported.
  boundary: >-
    nhr-47 is expressed in neurons, ventral nerve cord, spermatheca, pharynx and
    intestine, and germline knockdown modulates transgenerational/reproductive toxicant
    responses, but a stand-alone required function under normal conditions has not been
    defined.
  gap_kind:
  - BIOLOGY
  dark_aspect: BP_DARK
  status: OPEN
  significance: >-
    A defined loss-of-function phenotype would anchor NHR-47's biological process
    annotation, which is currently only family-propagated (IBA) or inferred from a
    toxicant-stress context.
  provenance:
  - reference_id: PMID:38922100
    supporting_text: >-
      RNAi of daf-12, nhr-14, and nhr-47 caused resistance, whereas RNAi of nhr-12
      conferred susceptibility to transgenerational PS-NP toxicity
  - reference_id: PMID:40482507
    supporting_text: >-
      6-PPDQ reproductive toxicity was suppressed by nhr-47, nhr-14, daf-12, and nhr-249
      RNAi and accelerated by nhr-12, nhr-145, and nhr-171 RNAi
suggested_questions:
- question: >-
    Is NHR-47 a ligand-regulated receptor, and if so what endogenous metabolite or lipid
    binds its ligand-binding domain, or is it a constitutive/orphan transcriptional
    regulator?
  experts:
  - Nuclear-receptor biochemists
  - C. elegans NHR-family biologists
- question: >-
    What are the direct genomic binding sites and target genes of NHR-47, and does it act
    as a transcriptional activator or repressor?
  experts:
  - Transcription / gene-regulatory-network biologists
- question: >-
    What is the endogenous (non-toxicant) physiological role of nhr-47, given its
    expression in neurons, the ventral nerve cord, spermatheca, pharynx and intestine and
    its germline role in toxicant responses?
  experts:
  - C. elegans developmental / germline biologists
suggested_experiments:
- experiment_type: Genome-wide binding assay
  description: >-
    Determine the direct NHR-47 regulon by ChIP-seq or CUT&RUN with an endogenously
    tagged NHR-47, combined with RNA-seq of a null mutant, to identify direct target genes
    and the DNA response element and to establish activator vs repressor behavior.
- experiment_type: Ligand identification
  description: >-
    Express and purify the NHR-47 ligand-binding domain and screen for bound endogenous
    lipids/metabolites (e.g. by lipidomics of co-purified ligands or reporter-based
    ligand-sensing assays) to test whether NHR-47 is ligand-regulated or a true orphan.
- experiment_type: Loss-of-function phenotyping
  description: >-
    Generate and characterize a null allele (CRISPR) across development, metabolism,
    reproduction, stress and innate-immune assays to define the baseline endogenous
    function, and test genetic interaction with the interacting receptor nhr-17.