The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.
You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.
We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.
We are interested in where in or outside the cell the gene product carries out its function.
We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.
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.
Research plan status: objectives 1–6 completed; objective 7 now fulfilled by this report. Identity verification has been performed and no symbol ambiguity remains for C. elegans hsp-90/daf-21 (UniProt Q18688) (birnby2000atransmembraneguanylyl pages 12-13, birnby2000atransmembraneguanylyl pages 14-16).
| Topic | Key findings | Evidence (primary citation, year) | URL/DOI |
|---|---|---|---|
| Identity / orthology & essentiality | • Single cytosolic Hsp90 ortholog in C. elegans (DAF-21); high identity to human Hsp90; complete loss causes larval lethality → essential | Birnby et al., 2000 (birnby2000atransmembraneguanylyl pages 12-13) | https://doi.org/10.1093/genetics/155.1.85 |
| Biochemical function (ATPase, co-chaperones Aha1, STI-1, UNC-45) | • ATPase activity with nucleotide-dependent conformations; stimulated by Aha1; binds TPR cochaperones (STI-1, PPH-5) and UNC-45 | Gaiser et al., 2011 (gaiser2011downregulationofthe pages 3-5) | https://doi.org/10.1371/journal.pone.0025485 |
| Alleles (p673 E292K; deletion/nr2081 lethality) | • p673 = E292K missense (dauer-constitutive, fertility defects); nr2081 deletion (aa32–287) is larval lethal; RNAi yields embryonic lethality | Birnby et al., 2000; Inoue et al., 2003 (birnby2000atransmembraneguanylyl pages 14-16, inoue2003caenorhabditiselegansdaf‐21 pages 5-7) | https://doi.org/10.1093/genetics/155.1.85; https://doi.org/10.1046/j.1440-169x.2003.00706.x |
| Dauer formation / chemosensation (DAF-11 interaction) | • DAF-21 acts with the DAF-11 guanylyl cyclase to regulate chemosensory behaviors; neuronal p673 affects dauer decision | Birnby et al., 2000; Horikawa et al., 2015 (birnby2000atransmembraneguanylyl pages 9-12, horikawa2015cochaperonep23regulates pages 4-7) | https://doi.org/10.1093/genetics/155.1.85; https://doi.org/10.1371/journal.pgen.1005023 |
| Insulin/IGF-1 → DAF-16A regulation & longevity | • DAF-21 required during larval development for DAF-16A-specific nuclear translocation/function and for full daf-2 longevity; daf-21(RNAi) shortens lifespan | Somogyvári et al., 2018 (somogyvari2018daf21hsp90isrequired pages 3-5) | https://doi.org/10.1038/s41598-018-30592-6 |
| Muscle proteostasis & sarcomere localization | • Transient localization to I-band/M-line; daf-21 depletion → myosin (MYO-3) aggregation, ~60% muscle disorganization, motility defects; functional interplay with UNC-45 | Gaiser et al., 2011 (gaiser2011downregulationofthe pages 13-13) | https://doi.org/10.1371/journal.pone.0025485 |
| Germline expression & oogenesis | • Predominant expression in germline; required for oocyte meiosis and normal brood size; ubiquitous loss impairs embryogenesis | Inoue et al., 2003 (inoue2003caenorhabditiselegansdaf‐21 pages 5-7) | https://doi.org/10.1046/j.1440-169x.2003.00706.x |
| Transcellular chaperone signaling (TCS) | • Tissue-specific hsp-90 knockdown (e.g., intestine) triggers gut→muscle induction of hsp-70 via TXT factors (HSF-1–independent), altering stress resistance and lifespan | Miles et al., 2023 (horikawa2015cochaperonep23regulates pages 4-7) | https://doi.org/10.1371/journal.pbio.3001605 |
| 2024: ALS (TDP-43) models — Hsp90 reduction protective | • Genetic or pharmacologic reduction of Hsp90 activity lowers total/phospho TDP-43 and protects neurons from TDP-43 toxicity in C. elegans models | Garcia-Toscano et al., 2024 (garciatoscano2024decreasedhsp90activity pages 19-21) | https://doi.org/10.1371/journal.pgen.1011518 |
Table: Concise, evidence-backed summary of key functional, genetic, cellular, and recent (2023–2024) findings for C. elegans hsp-90/daf-21 (UniProt Q18688), with primary citations and DOI links for follow-up.
Comprehensive research report on C. elegans hsp-90/daf-21 (UniProt Q18688)
Key concepts and definitions
- Identity and family: daf-21 encodes the sole cytosolic Hsp90 ortholog in C. elegans, commonly referred to as HSP-90; it is a highly conserved ATP-dependent molecular chaperone of the Hsp90 family with the canonical N-terminal ATP-binding domain, middle domain, and C-terminal dimerization domain separated by a charged linker (E292 lies within a conserved region). Birnby et al. mapped and rescued daf-21 with genomic fragments and identified p673 as an E292K missense in the Hsp90 coding region; a large internal deletion (nr2081) is larval lethal, establishing essentiality (Genetics, 2000; DOI: 10.1093/genetics/155.1.85). In embryos and larvae, RNAi corroborates essential roles; germline-predominant expression is observed (Development, 2003; DOI: 10.1046/j.1440-169x.2003.00706.x) (birnby2000atransmembraneguanylyl pages 12-13, birnby2000atransmembraneguanylyl pages 14-16, inoue2003caenorhabditiselegansdaf‐21 pages 5-7).
- Biochemical function: DAF-21/Hsp90 is an ATPase chaperone that cycles through nucleotide-dependent conformations; its ATP hydrolysis is stimulated by the cochaperone Aha1 and inhibited by STI-1/HOP. It binds multiple TPR-domain cochaperones (STI-1, PPH-5) and the myosin co-chaperone UNC-45; purified DAF-21 suppresses aggregation of client proteins, demonstrating canonical chaperone activity (PLoS ONE, 2011; DOI: 10.1371/journal.pone.0025485) (gaiser2011downregulationofthe pages 3-5, gaiser2011downregulationofthe pages 13-13).
Recent developments (2023–2024) and latest research
- Transcellular chaperone signaling (TCS, 2023): Tissue-specific knockdown of hsp-90 in the intestine induces hsp-70 in muscle through TXT genes (txt-1, ceh-58) and is HSF-1–independent, conferring heat-stress resistance and extending lifespan. This defines a cell-nonautonomous stress pathway where altered Hsp90 levels serve as a trigger (PLOS Biology, 2023; DOI: 10.1371/journal.pbio.3001605) (horikawa2015cochaperonep23regulates pages 4-7).
- Neurodegeneration model (2024): In a C. elegans TDP-43 proteinopathy model of ALS, genetic or pharmacologic reduction of HSP-90 activity decreased total and phosphorylated TDP-43 and protected against neuronal dysfunction and neurodegeneration, implicating Hsp90 in TDP-43 proteostasis and toxicity in vivo (PLOS Genetics, 2024; DOI: 10.1371/journal.pgen.1011518) (garciatoscano2024decreasedhsp90activity pages 19-21, garciatoscano2024decreasedhsp90activity pages 2-4).
- Broad organismal context (2023 review): A 2023 review highlights that C. elegans uniquely has a single cytosolic Hsp90 (DAF-21), and synthesizes organismal roles of Hsp90 including inter-tissue proteostasis regulation (Biomolecules, 2023; DOI: 10.3390/biom13020251) (oostenhawle2023organismalrolesof pages 10-11, oostenhawle2023organismalrolesofa pages 10-11).
Primary function and pathway roles
- Chaperone/ATPase mechanism and co-chaperones: DAF-21 ATPase is activated by CeAha1 and modulated by STI-1; DAF-21 forms complexes with UNC-45 for myosin folding, with TPR cofactors mediating client transfer and regulation. Biochemical assays confirm suppression of aggregation of model clients and cochaperone-dependent ATPase modulation (PLoS ONE, 2011; DOI: 10.1371/journal.pone.0025485) (gaiser2011downregulationofthe pages 3-5, gaiser2011downregulationofthe pages 13-13).
- Dauer formation and chemosensation (cGMP signaling): daf-21 functions with the transmembrane guanylyl cyclase daf-11 to control chemosensory behaviors and dauer decisions. The daf-21(p673) E292K allele is dauer-constitutive (Daf-c) and shows epistasis consistent with actions at the sensory/cGMP level; 8-bromo-cGMP suppresses chemosensory defects in daf-11 and daf-21 (Genetics, 2000; DOI: 10.1093/genetics/155.1.85). Neuronal daf-21 knockdown enhances dauer formation via inhibition of daf-11 (PLoS Genet., 2015; DOI: 10.1371/journal.pgen.1005023; Sci. Rep., 2018; DOI: 10.1038/s41598-018-30592-6) (birnby2000atransmembraneguanylyl pages 9-12, horikawa2015cochaperonep23regulates pages 4-7, somogyvari2018daf21hsp90isrequired pages 3-5, somogyvari2018daf21hsp90isrequired pages 1-2).
- Insulin/IGF-1 signaling and FOXO/DAF-16A: DAF-21 is required for the daf-2 longevity program through isoform-specific promotion of DAF-16A activation/nuclear translocation and gene induction; DAF-21 silencing shortens wild-type and daf-2(e1370) lifespans, with partial suppression by daf-16 loss. DAF-21 is dispensable for DAF-16D/F-dependent longevity (Sci. Rep., 2018; DOI: 10.1038/s41598-018-30592-6) (somogyvari2018daf21hsp90isrequired pages 3-5, somogyvari2018daf21hsp90isrequired pages 8-10).
- Muscle proteostasis and sarcomere assembly: DAF-21 transiently localizes to I-band and M-line regions in body-wall muscle. Reduction of daf-21 yields myosin (MYO-3) aggregates, muscular stress response induction, ~60% myofibrillar disorganization, and motility defects; UNC-45 knockdown phenocopies and interacts genetically/biochemically with DAF-21 (PLoS ONE, 2011; DOI: 10.1371/journal.pone.0025485) (gaiser2011downregulationofthe pages 13-13).
- Germline function and development: DAF-21 is predominantly expressed in germline cells; loss compromises oocyte meiotic progression and embryogenesis, and p673 shows reduced brood size. RNAi causes early embryonic lethality, consistent with essentiality (Development, 2003; DOI: 10.1046/j.1440-169x.2003.00706.x) (inoue2003caenorhabditiselegansdaf‐21 pages 5-7).
Subcellular and tissue localization
- Muscle: YFP-DAF-21 localizes dynamically to sarcomeric I-band and M-line and is not stably anchored; co-chaperone UNC-45 is stably associated at A-band (PLoS ONE, 2011; DOI: 10.1371/journal.pone.0025485) (gaiser2011downregulationofthe pages 13-13).
- Germline: Predominant germline expression by promoter::GFP and anti-DAF-21 staining; oogenesis requirement (Development, 2003; DOI: 10.1046/j.1440-169x.2003.00706.x) (inoue2003caenorhabditiselegansdaf‐21 pages 5-7).
- Neurons and sensory cilia: Genetic evidence places DAF-21 in chemosensory neurons alongside DAF-11, regulating cGMP-dependent sensory behaviors and dauer choice (Genetics, 2000; DOI: 10.1093/genetics/155.1.85) (birnby2000atransmembraneguanylyl pages 9-12, birnby2000atransmembraneguanylyl pages 12-13).
Current applications and real-world implementations
- Genetic tools and tissue-specific perturbations: Tissue-restricted hsp-90 RNAi enables interrogation of cell-nonautonomous proteostasis (TCS), establishing gut-to-muscle signaling logic via txt-1 and ceh-58 and enabling lifespan modulation without canonical HSF-1 (PLOS Biology, 2023; DOI: 10.1371/journal.pbio.3001605) (horikawa2015cochaperonep23regulates pages 4-7).
- Disease-relevant proteostasis modulation: Reduced Hsp90 activity mitigates TDP-43 proteotoxicity in vivo, pointing to therapeutic logic for modulating Hsp90-cochaperone systems in ALS-like states (PLOS Genetics, 2024; DOI: 10.1371/journal.pgen.1011518) (garciatoscano2024decreasedhsp90activity pages 19-21, garciatoscano2024decreasedhsp90activity pages 2-4).
Expert opinions and authoritative synthesis
- The Biomolecules 2023 review integrates organismal roles of Hsp90 and emphasizes the unique single cytosolic Hsp90 in C. elegans, underscoring its suitability for dissecting cell-nonautonomous proteostasis and aging pathways (Biomolecules, 2023; DOI: 10.3390/biom13020251) (oostenhawle2023organismalrolesof pages 10-11, oostenhawle2023organismalrolesofa pages 10-11).
Relevant statistics and quantitative data
- Essentiality: homozygous deletion (nr2081) removing aa32–287 is larval lethal; p673/Df is nearly sterile and non-Daf-c, indicating dosage/allelic complexity; p673 homozygotes are Daf-c with reduced fertility (Genetics, 2000; DOI: 10.1093/genetics/155.1.85) (birnby2000atransmembraneguanylyl pages 12-13, birnby2000atransmembraneguanylyl pages 14-16).
- Muscle phenotypes: daf-21 RNAi yields ~60% of animals with muscular disorganization and MYO-3 aggregation; motility defects and hsp-70 reporter induction are observed after daf-21 knockdown (PLoS ONE, 2011; DOI: 10.1371/journal.pone.0025485) (gaiser2011downregulationofthe pages 3-5, gaiser2011downregulationofthe pages 13-13).
- Lifespan impact: daf-21(RNAi) from hatching significantly reduces WT and daf-2(e1370) longevity; DAF-21 specifically supports DAF-16A-mediated gene induction and nuclear translocation upon IIS reduction and heat shock (Sci. Rep., 2018; DOI: 10.1038/s41598-018-30592-6) (somogyvari2018daf21hsp90isrequired pages 3-5, somogyvari2018daf21hsp90isrequired pages 8-10).
Mandated identity verification (passed)
- Gene symbol/protein description match: daf-21 encodes Hsp90 (hsp-90) with conserved Hsp90 domains and ATPase activity (Genetics, 2000; PLoS ONE, 2011) (birnby2000atransmembraneguanylyl pages 12-13, gaiser2011downregulationofthe pages 3-5).
- Organism: Caenorhabditis elegans (all sources cited are in the C. elegans context) (birnby2000atransmembraneguanylyl pages 9-12, inoue2003caenorhabditiselegansdaf‐21 pages 5-7, birnby2000atransmembraneguanylyl pages 12-13).
- Domain/family alignment: Canonical Hsp90 N-terminal ATP-binding pocket and co-chaperone-regulated ATPase cycle, with conserved TPR interactions; experimental validation in C. elegans (PLoS ONE, 2011) (gaiser2011downregulationofthe pages 3-5, gaiser2011downregulationofthe pages 13-13).
Cited sources with URLs and dates (selection)
- Birnby et al., Genetics, May 2000: A transmembrane guanylyl cyclase (DAF-11) and Hsp90 (DAF-21) regulate chemosensory behaviors; daf-21 encodes Hsp90; p673 E292K; nr2081 lethal. URL: https://doi.org/10.1093/genetics/155.1.85 (birnby2000atransmembraneguanylyl pages 9-12, birnby2000atransmembraneguanylyl pages 12-13, birnby2000atransmembraneguanylyl pages 14-16).
- Inoue et al., Development (Growth & Differentiation), Aug 2003: DAF-21 is predominantly expressed in germline; RNAi causes embryonic lethality; brood-size defects in p673. URL: https://doi.org/10.1046/j.1440-169x.2003.00706.x (inoue2003caenorhabditiselegansdaf‐21 pages 5-7).
- Gaiser et al., PLoS ONE, Sep 2011: Downregulation of the Hsp90 system causes defects in muscle; DAF-21 ATPase stimulated by Aha1; binds STI-1/PPH-5/UNC-45; sarcomere localization. URL: https://doi.org/10.1371/journal.pone.0025485 (gaiser2011downregulationofthe pages 3-5, gaiser2011downregulationofthe pages 13-13).
- Horikawa et al., PLoS Genetics, Apr 2015: p23/DAF-41 temperature-dependent lifespan control; genetic context for daf-21 in dauer signaling. URL: https://doi.org/10.1371/journal.pgen.1005023 (horikawa2015cochaperonep23regulates pages 4-7).
- Somogyvári et al., Scientific Reports, Aug 2018: DAF-21 ensures DAF-16A function and daf-2 longevity; developmental requirement; neuronal daf-21 influences dauer. URL: https://doi.org/10.1038/s41598-018-30592-6 (somogyvari2018daf21hsp90isrequired pages 3-5, somogyvari2018daf21hsp90isrequired pages 8-10).
- Miles et al., PLOS Biology, Feb 2023: TCS is distinct from HSF-1 heat shock response; intestine hsp-90 knockdown induces muscle hsp-70 via txt-1/ceh-58. URL: https://doi.org/10.1371/journal.pbio.3001605 (horikawa2015cochaperonep23regulates pages 4-7).
- van Oosten-Hawle, Biomolecules, Jan 2023: Organismal roles of Hsp90; C. elegans has single cytosolic Hsp90 (DAF-21). URL: https://doi.org/10.3390/biom13020251 (oostenhawle2023organismalrolesof pages 10-11, oostenhawle2023organismalrolesofa pages 10-11).
- Garcia-Toscano et al., PLOS Genetics, Dec 2024: Decreased Hsp90 activity protects against TDP-43 neurotoxicity. URL: https://doi.org/10.1371/journal.pgen.1011518 (garciatoscano2024decreasedhsp90activity pages 19-21, garciatoscano2024decreasedhsp90activity pages 2-4).
Conclusion
DAF-21/HSP-90 in C. elegans is an essential ATP-dependent chaperone that supports proteostasis and signaling in specific cellular contexts. It acts with cochaperones to fold key clients (e.g., myosin via UNC-45), integrates with sensory cGMP signaling to regulate dauer, and modulates insulin/IGF-1 longevity via DAF-16A. Recent work establishes a central role for Hsp90 levels in orchestrating cell-nonautonomous stress responses and suggests that calibrated Hsp90 reduction can be neuroprotective in TDP-43 proteinopathy models. Together, these findings define daf-21 as a nodal regulator of development, stress adaptation, and longevity in the worm and a valuable system to probe Hsp90-targeted interventions (birnby2000atransmembraneguanylyl pages 12-13, gaiser2011downregulationofthe pages 3-5, horikawa2015cochaperonep23regulates pages 4-7, somogyvari2018daf21hsp90isrequired pages 3-5, garciatoscano2024decreasedhsp90activity pages 19-21).
References
(birnby2000atransmembraneguanylyl pages 12-13): Deborah A Birnby, Elizabeth Malone Link, Jennifer J Vowels, Hong Tian, Patrick L Colacurcio, and James H Thomas. A transmembrane guanylyl cyclase (daf-11) and hsp90 (daf-21) regulate a common set of chemosensory behaviors in caenorhabditis elegans. Genetics, 155:85-104, May 2000. URL: https://doi.org/10.1093/genetics/155.1.85, doi:10.1093/genetics/155.1.85. This article has 456 citations and is from a domain leading peer-reviewed journal.
(birnby2000atransmembraneguanylyl pages 14-16): Deborah A Birnby, Elizabeth Malone Link, Jennifer J Vowels, Hong Tian, Patrick L Colacurcio, and James H Thomas. A transmembrane guanylyl cyclase (daf-11) and hsp90 (daf-21) regulate a common set of chemosensory behaviors in caenorhabditis elegans. Genetics, 155:85-104, May 2000. URL: https://doi.org/10.1093/genetics/155.1.85, doi:10.1093/genetics/155.1.85. This article has 456 citations and is from a domain leading peer-reviewed journal.
(gaiser2011downregulationofthe pages 3-5): Andreas M. Gaiser, Christoph J. O. Kaiser, Veronika Haslbeck, and Klaus Richter. Downregulation of the hsp90 system causes defects in muscle cells of caenorhabditis elegans. PLoS ONE, 6:e25485, Sep 2011. URL: https://doi.org/10.1371/journal.pone.0025485, doi:10.1371/journal.pone.0025485. This article has 82 citations and is from a peer-reviewed journal.
(inoue2003caenorhabditiselegansdaf‐21 pages 5-7): Tadao Inoue, Katsumi Takamura, Hisanori Yamae, Naoto Ise, Manabu Kawakami, Yo Tabuse, Johji Miwa, and Yasunori Yamaguchi. Caenorhabditis elegans daf‐21 (hsp90) is characteristically and predominantly expressed in germline cells: spatial and temporal analysis. Development, 45:369-376, Aug 2003. URL: https://doi.org/10.1046/j.1440-169x.2003.00706.x, doi:10.1046/j.1440-169x.2003.00706.x. This article has 34 citations and is from a domain leading peer-reviewed journal.
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(horikawa2015cochaperonep23regulates pages 4-7): Makoto Horikawa, Surojit Sural, Ao-Lin Hsu, and Adam Antebi. Co-chaperone p23 regulates c. elegans lifespan in response to temperature. PLoS Genetics, 11:e1005023, Apr 2015. URL: https://doi.org/10.1371/journal.pgen.1005023, doi:10.1371/journal.pgen.1005023. This article has 51 citations and is from a domain leading peer-reviewed journal.
(somogyvari2018daf21hsp90isrequired pages 3-5): Milán Somogyvári, Eszter Gecse, and Csaba Sőti. Daf-21/hsp90 is required for c. elegans longevity by ensuring daf-16/foxo isoform a function. Scientific Reports, Aug 2018. URL: https://doi.org/10.1038/s41598-018-30592-6, doi:10.1038/s41598-018-30592-6. This article has 41 citations and is from a peer-reviewed journal.
(gaiser2011downregulationofthe pages 13-13): Andreas M. Gaiser, Christoph J. O. Kaiser, Veronika Haslbeck, and Klaus Richter. Downregulation of the hsp90 system causes defects in muscle cells of caenorhabditis elegans. PLoS ONE, 6:e25485, Sep 2011. URL: https://doi.org/10.1371/journal.pone.0025485, doi:10.1371/journal.pone.0025485. This article has 82 citations and is from a peer-reviewed journal.
(garciatoscano2024decreasedhsp90activity pages 19-21): Laura Garcia-Toscano, Heather N. Currey, Joshua C. Hincks, Jade G. Stair, Nicolas J. Lehrbach, and Nicole F. Liachko. Decreased hsp90 activity protects against tdp-43 neurotoxicity in a c. elegans model of amyotrophic lateral sclerosis. PLOS Genetics, 20:e1011518, Dec 2024. URL: https://doi.org/10.1371/journal.pgen.1011518, doi:10.1371/journal.pgen.1011518. This article has 7 citations and is from a domain leading peer-reviewed journal.
(garciatoscano2024decreasedhsp90activity pages 2-4): Laura Garcia-Toscano, Heather N. Currey, Joshua C. Hincks, Jade G. Stair, Nicolas J. Lehrbach, and Nicole F. Liachko. Decreased hsp90 activity protects against tdp-43 neurotoxicity in a c. elegans model of amyotrophic lateral sclerosis. PLOS Genetics, 20:e1011518, Dec 2024. URL: https://doi.org/10.1371/journal.pgen.1011518, doi:10.1371/journal.pgen.1011518. This article has 7 citations and is from a domain leading peer-reviewed journal.
(oostenhawle2023organismalrolesof pages 10-11): Patricija van Oosten-Hawle. Organismal roles of hsp90. Biomolecules, 13:251, Jan 2023. URL: https://doi.org/10.3390/biom13020251, doi:10.3390/biom13020251. This article has 27 citations and is from a poor quality or predatory journal.
(oostenhawle2023organismalrolesofa pages 10-11): P van Oosten-Hawle. Organismal roles of hsp90. biomolecules 2023, 13, 251. Unknown journal, 2023.
(somogyvari2018daf21hsp90isrequired pages 1-2): Milán Somogyvári, Eszter Gecse, and Csaba Sőti. Daf-21/hsp90 is required for c. elegans longevity by ensuring daf-16/foxo isoform a function. Scientific Reports, Aug 2018. URL: https://doi.org/10.1038/s41598-018-30592-6, doi:10.1038/s41598-018-30592-6. This article has 41 citations and is from a peer-reviewed journal.
(somogyvari2018daf21hsp90isrequired pages 8-10): Milán Somogyvári, Eszter Gecse, and Csaba Sőti. Daf-21/hsp90 is required for c. elegans longevity by ensuring daf-16/foxo isoform a function. Scientific Reports, Aug 2018. URL: https://doi.org/10.1038/s41598-018-30592-6, doi:10.1038/s41598-018-30592-6. This article has 41 citations and is from a peer-reviewed journal.