HSPH1

UniProt ID: Q60446
Organism: Cricetulus griseus
Review Status: DRAFT
๐Ÿ“ Provide Detailed Feedback

Gene Description

HSPH1 (Hsp105/Hsp110) is a member of the Hsp110 family, a significantly diverged subgroup of the Hsp70 superfamily. It has two core molecular functions: (1) holdase chaperone activity, preventing the aggregation of heat-denatured proteins by maintaining them in a soluble, folding-competent state (significantly more efficient than Hsc70 at this function), and (2) nucleotide exchange factor (NEF) activity for Hsp70 family members (HSPA1A, HSPA1B), promoting ADP release and triggering substrate release. Importantly, HSPH1 does NOT itself refold denatured proteins; refolding requires Hsc70/Hdj-1. HSPH1 also inhibits HSPA8/HSC70 ATPase activity. Overexpression confers substantial thermoresistance in vivo.

Proposed New Ontology Terms

holdase chaperone activity

Definition: Binding of unfolded or misfolded proteins to prevent their aggregation, without actively catalyzing refolding. This activity maintains client proteins in a soluble, folding-competent state for subsequent refolding by other chaperone systems.

Justification: Multiple proteins including HSPH1/Hsp110, small HSPs (CRYAA, CRYAB, HSPB6), and clusterin (CLU) function as holdases rather than foldases. The current GO term GO:0044183 (protein folding chaperone) implies active folding, which does not accurately describe holdase function. See go-ontology#30552.

Parent term: protein folding chaperone

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005634 nucleus
IEA
GO_REF:0000118
KEEP AS NON CORE
Summary: TreeGrafter-inferred nuclear localization. UniProt subcellular location for Q60446 lists only cytoplasm (by similarity to human Q92598). However, the deep research review (HSPH1-deep-research-falcon.md) notes that Hsp105alpha (HSPH1) localizes constitutively to both cytoplasm and nucleus in multiple mammalian tissues, including brain. The TreeGrafter inference is therefore supported by recent literature on mammalian orthologs.
Reason: Nuclear localization is supported by literature on mammalian HSPH1 orthologs. The deep research review cites Chuang et al. 2024 as reporting constitutive cytoplasmic and nuclear localization. However, the primary functional compartment is the cytosol where the holdase and NEF activities operate. Keeping as non-core since it is a secondary localization.
Supporting Evidence:
file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md
Constitutive cytoplasmic and nuclear localization is reported for Hsp105alpha (HSPH1) [in mammalian tissues]
GO:0005829 cytosol
IEA
GO_REF:0000118
ACCEPT
Summary: TreeGrafter-inferred cytosol localization, consistent with UniProt annotation of cytoplasm for HSPH1 (by similarity to human Q92598). HSPH1 is a cytoplasmic chaperone and its holdase and NEF functions operate in the cytoplasm. The deep research review (HSPH1-deep-research-falcon.md) confirms cytosolic localization as the primary site of HSPH1 function.
Reason: Cytosol is the primary localization for HSPH1, consistent with UniProt annotation of cytoplasm and with its known functions as a cytoplasmic chaperone and NEF.
Supporting Evidence:
PMID:9395504
hsp110 is one of the principal molecular chaperones of mammalian cells
GO:0006457 protein folding
IEA
GO_REF:0000118
KEEP AS NON CORE
Summary: TreeGrafter-inferred involvement in protein folding. HSPH1 is involved in protein folding homeostasis, but importantly it is a holdase, not a foldase. It maintains denatured proteins in a folding-competent state but does NOT actively refold them (PMID:9395504). Refolding requires Hsc70/Hdj-1. The term 'protein folding' is acceptable as a broad biological process annotation since HSPH1 participates in the protein folding pathway (by handing off substrates to the Hsc70/Hdj-1 refolding machinery), but its direct activity is holding, not folding. As a NEF, HSPH1 also accelerates Hsp70 cycling which supports the overall protein folding process (deep research review, HSPH1-deep-research-falcon.md).
Reason: HSPH1 participates in the protein folding pathway by preventing aggregation and maintaining substrates in a folding-competent state for subsequent refolding by Hsc70/Hdj-1. The BP term 'protein folding' is broadly acceptable but HSPH1 is a holdase, not a foldase, so this is kept as non-core. Its direct role is prevention of aggregation rather than active folding.
Supporting Evidence:
PMID:9395504
hsp110 is highly efficient in selectively recognizing denatured proteins and maintaining them in a soluble, folding-competent state
PMID:9395504
hsp110-bound proteins can then be refolded by the addition of rabbit reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself function as a co-chaperone in folding with hsp110
GO:0000166 nucleotide binding
IEA
GO_REF:0000043
ACCEPT
Summary: IEA annotation from UniProtKB-KW:KW-0547 (Nucleotide-binding) keyword mapping. HSPH1 belongs to the Hsp70 superfamily and possesses the conserved nucleotide-binding domain (NBD). This is correct but overly general; ATP binding (GO:0005524) is more specific and is already annotated.
Reason: Nucleotide binding is correct for HSPH1, which has a conserved Hsp70-type nucleotide-binding domain. While more general than the co-annotated GO:0005524 (ATP binding), the IEA mapping from UniProt keyword is valid. Both annotations can coexist since the IEA is broader.
GO:0005524 ATP binding
IEA
GO_REF:0000120
ACCEPT
Summary: Combined IEA annotation from InterPro:IPR013126 (Hsp_70_fam) and UniProtKB-KW:KW-0067 (ATP-binding). HSPH1 has a conserved Hsp70-type nucleotide-binding domain and binds ATP. While Hsp110 family members have greatly reduced ATPase activity compared to canonical Hsp70s, they do bind ATP, which is important for their NEF function.
Reason: ATP binding is a core property of HSPH1. The Hsp110 family binds ATP through their conserved NBD. UniProt keywords and InterPro domain classification both support this annotation. ATP binding is required for the NEF function of HSPH1.
GO:0005737 cytoplasm
IEA
GO_REF:0000044
ACCEPT
Summary: IEA annotation from UniProtKB subcellular location vocabulary mapping (SL-0086). UniProt explicitly states cytoplasm localization for HSPH1 (by similarity to human Q92598). This is the primary localization and is consistent with its chaperone/NEF functions.
Reason: Cytoplasm is the primary subcellular localization for HSPH1. UniProt annotation (by similarity) and the protein's established cytoplasmic functions support this.
Supporting Evidence:
PMID:9395504
hsp110 is one of the principal molecular chaperones of mammalian cells
GO:0006950 response to stress
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: ARBA-inferred involvement in response to stress. HSPH1 is a heat shock protein whose expression is upregulated under stress conditions. UniProt keywords include 'Stress response'. This is correct but very general; the more specific term GO:0034605 (cellular response to heat) is already annotated with IDA evidence.
Reason: Response to stress is valid for this heat shock protein but is very general. The more specific IDA-supported annotation to GO:0034605 (cellular response to heat) captures the core stress response function. Keeping this broader IEA annotation as non-core since it adds no information beyond the specific IDA annotation.
GO:0016887 ATP hydrolysis activity
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro-inferred ATP hydrolysis activity from IPR013126 (Hsp_70_fam). While HSPH1 belongs to the Hsp70 superfamily and has the conserved ATPase domain architecture, Hsp110 family members have greatly reduced intrinsic ATPase activity compared to canonical Hsp70 chaperones. The deep research review (HSPH1-deep-research-falcon.md) emphasizes that HSPH1 functions primarily as a NEF rather than an ATPase. The ATPase annotation from the broad Hsp70 family InterPro entry is misleading for this diverged subfamily.
Reason: Hsp110 family members have significantly reduced ATPase activity compared to canonical Hsp70 proteins. The InterPro family IPR013126 covers the entire Hsp70 superfamily, and while the domain architecture is conserved, Hsp110 members have diverged to function primarily as holdases and NEFs rather than as ATPases. Annotating HSPH1 with GO:0016887 (ATP hydrolysis activity) overstates its catalytic function. The core molecular functions are holdase chaperone activity and NEF activity (GO:0000774).
GO:0031249 denatured protein binding
IDA
PMID:9395504
Hsp110 protects heat-denatured proteins and confers cellular...
MODIFY
Summary: IDA annotation based on Oh et al. 1997, which demonstrated that Hsp110 is highly efficient at recognizing heat-denatured proteins and maintaining them in a soluble, folding-competent state. This is a genuine holdase chaperone function. GO:0031249 is being obsoleted (go-ontology#30962). The appropriate replacement is GO:0044183 (protein folding chaperone) as a placeholder, with the caveat that HSPH1 is a holdase, not a foldase, and a more specific holdase term is needed (go-ontology#30552).
Reason: GO:0031249 (denatured protein binding) is being obsoleted. The experimental evidence from PMID:9395504 clearly demonstrates holdase chaperone activity: Hsp110 prevents aggregation of heat-denatured proteins and maintains them in a folding-competent state. GO:0044183 (protein folding chaperone) is the closest available replacement MF term, used as a placeholder until a specific holdase chaperone activity term is created (go-ontology#30552). Note that HSPH1 does NOT refold proteins itself; refolding requires Hsc70/Hdj-1.
Proposed replacements: protein folding chaperone
Supporting Evidence:
PMID:9395504
hsp110 is highly efficient in selectively recognizing denatured proteins and maintaining them in a soluble, folding-competent state and is significantly more efficient in performing this function than is hsc70
PMID:9395504
hsp110-bound proteins can then be refolded by the addition of rabbit reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself function as a co-chaperone in folding with hsp110
GO:0034605 cellular response to heat
IDA
PMID:9395504
Hsp110 protects heat-denatured proteins and confers cellular...
ACCEPT
Summary: IDA annotation based on Oh et al. 1997, which demonstrated that overexpression of Hsp110 in vivo conferred substantial heat resistance to both Rat-1 and HeLa cells. This is a core biological process for HSPH1 as a heat shock protein.
Reason: Well-supported by direct experimental evidence from PMID:9395504 showing that Hsp110 overexpression confers thermoresistance. HSPH1 is one of the primary heat shock proteins in mammalian cells, and cellular response to heat is a core biological process.
Supporting Evidence:
PMID:9395504
the overexpression in vivo of hsp110 conferred substantial heat resistance to both Rat-1 and HeLa cells
PMID:9395504
hsp110 is one of the principal molecular chaperones of mammalian cells and represents a newly identified component of the primary protection/repair pathway for denatured proteins and thermotolerance expression in vivo
GO:0000774 adenyl-nucleotide exchange factor activity
ISS
GO_REF:0000024
ACCEPT
Summary: ISS annotation transferred from human HSPH1 (Q92598). UniProt describes HSPH1 as acting as a nucleotide-exchange factor (NEF) for HSPA1A and HSPA1B, promoting the release of ADP thereby triggering substrate release (by similarity to mouse Q61699 and human Q92598). This is one of the two core molecular functions of Hsp110 family members. The deep research review (HSPH1-deep-research-falcon.md) confirms that HSPH1 operates as a NEF for cytosolic Hsp70s, binding the Hsp70 NBD to catalyze ADP-to-ATP exchange.
Reason: NEF activity is a core molecular function of the Hsp110 family. While the evidence for Chinese hamster HSPH1 specifically is by similarity (ISS), the NEF function is well-conserved across mammalian Hsp110 orthologs and is supported by UniProt annotation (by similarity to Q61699 mouse and Q92598 human). This represents one of the two principal molecular functions of HSPH1 (the other being holdase activity).
GO:0044183 protein folding chaperone
IDA
PMID:9395504
Hsp110 protects heat-denatured proteins and confers cellular...
NEW
Summary: Proposed new annotation as the replacement MF term for the obsoleting GO:0031249. PMID:9395504 provides direct experimental evidence that Hsp110 functions as a chaperone that prevents aggregation of denatured proteins and maintains them in a folding-competent state. This is a holdase chaperone function (placeholder annotation pending creation of a specific holdase term per go-ontology#30552).
Reason: This annotation replaces the obsoleting GO:0031249 (denatured protein binding) with the closest available MF term. HSPH1 is a holdase chaperone that binds denatured proteins to prevent aggregation. GO:0044183 is a placeholder until a specific holdase chaperone activity term is created (go-ontology#30552).
Supporting Evidence:
PMID:9395504
hsp110 is highly efficient in selectively recognizing denatured proteins and maintaining them in a soluble, folding-competent state

Core Functions

Holdase chaperone activity: HSPH1 binds heat-denatured proteins and prevents their aggregation, maintaining them in a soluble, folding-competent state. This function is significantly more efficient than Hsc70. HSPH1 does NOT actively refold proteins; refolding requires Hsc70/Hdj-1. This is a placeholder annotation using GO:0044183 until a specific holdase chaperone activity term is created (go-ontology#30552).

Molecular Function:
protein folding chaperone
Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • PMID:9395504
    hsp110 is highly efficient in selectively recognizing denatured proteins and maintaining them in a soluble, folding-competent state

Nucleotide exchange factor (NEF) activity for Hsp70 family members: HSPH1 promotes the release of ADP from HSPA1A/HSPA1B, triggering substrate release from the Hsp70 chaperone cycle. Also inhibits HSPA8/HSC70 ATPase and chaperone activities.

Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • PMID:9395504
    hsp110 is one of the principal molecular chaperones of mammalian cells and represents a newly identified component of the primary protection/repair pathway for denatured proteins and thermotolerance expression in vivo

References

Gene Ontology annotation through association of InterPro records with GO terms
Manual transfer of experimentally-verified manual GO annotation data to orthologs by curator judgment of sequence similarity
Gene Ontology annotation based on UniProtKB/Swiss-Prot keyword mapping
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Electronic Gene Ontology annotations created by ARBA machine learning models
TreeGrafter-generated GO annotations
Combined Automated Annotation using Multiple IEA Methods
Hsp110 protects heat-denatured proteins and confers cellular thermoresistance.
  • Hsp110 is highly efficient at preventing aggregation of heat-denatured proteins, functioning as a holdase chaperone that maintains substrates in a folding-competent state. It is significantly more efficient than Hsc70 at this holdase function.
    "hsp110 is highly efficient in selectively recognizing denatured proteins and maintaining them in a soluble, folding-competent state and is significantly more efficient in performing this function than is hsc70"
  • Hsp110 does not refold denatured proteins itself; refolding requires Hsc70 and Hdj-1. Hdj-1 does not function as a co-chaperone with Hsp110.
    "hsp110-bound proteins can then be refolded by the addition of rabbit reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself function as a co-chaperone in folding with hsp110"
  • Overexpression of Hsp110 confers substantial thermoresistance in vivo.
    "the overexpression in vivo of hsp110 conferred substantial heat resistance to both Rat-1 and HeLa cells"
Identification of a major subfamily of large hsp70-like proteins through the cloning of the mammalian 110-kDa heat shock protein.
  • HSPH1 (Hsp110) was identified as a member of a significantly diverged subgroup of the Hsp70 protein family.
    "Identification of a major subfamily of large hsp70-like proteins through the cloning of the mammalian 110-kDa heat shock protein"
file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md
Deep research review of HSPH1 function
  • HSPH1 operates as a NEF for cytosolic Hsp70s, catalyzing ADP-to-ATP exchange on the Hsp70 NBD to reset the chaperone for subsequent substrate cycles.
  • HSPH1 participates in the Hsp70-DNAJ-HSPH1 disaggregase complex that can disaggregate heat-denatured luciferase and disease-associated amyloids.
  • Constitutive cytoplasmic and nuclear localization is reported for Hsp105alpha (HSPH1) in mammalian tissues.

Suggested Questions for Experts

Q: Does Chinese hamster HSPH1 have the same NEF activity for Hsp70 as the mouse and human orthologs? The ISS annotation is based on similarity but has not been directly demonstrated for the Chinese hamster protein.

Suggested experts: Subjeck JR

Q: What is the intrinsic ATPase activity level of HSPH1? While Hsp110 family members are known to have reduced ATPase activity compared to canonical Hsp70, the exact level for Chinese hamster HSPH1 has not been characterized.

Suggested experts: Subjeck JR

Suggested Experiments

Experiment: In vitro nucleotide exchange assay using purified Chinese hamster HSPH1 and Hsp70 (HSPA1A). Measure ADP release rates from Hsp70 in the presence and absence of HSPH1 using fluorescent nucleotide analogs.

Hypothesis: Chinese hamster HSPH1 functions as a NEF for hamster Hsp70 family members, similar to the characterized function of mouse and human orthologs.

Type: biochemical assay

Experiment: Comparative ATPase assay measuring the rate of ATP hydrolysis by purified HSPH1 versus HSPA1A using a malachite green phosphate detection assay.

Hypothesis: HSPH1 has significantly lower intrinsic ATPase activity compared to canonical Hsp70 family members.

Type: biochemical assay

๐Ÿ“š Additional Documentation

Deep Research Falcon

(HSPH1-deep-research-falcon.md)

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gene_id: HSPH1
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shock 110 kDa protein;'
gene_info: Name=HSPH1; Synonyms=HSP105, HSP110;
organism_full: Cricetulus griseus (Chinese hamster) (Cricetulus barabensis griseus).
protein_family: Belongs to the heat shock protein 70 family. .
protein_domains: ATPase_NBD. (IPR043129); Heat_shock_70_CS. (IPR018181); HSP70_C_sf.
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Question

Gene Research for Functional Annotation

โš ๏ธ CRITICAL: Gene/Protein Identification Context

BEFORE YOU BEGIN RESEARCH: You MUST verify you are researching the CORRECT gene/protein. Gene symbols can be ambiguous, especially for less well-characterized genes from non-model organisms.

Target Gene/Protein Identity (from UniProt):

  • UniProt Accession: Q60446
  • Protein Description: RecName: Full=Heat shock protein 105 kDa; AltName: Full=Heat shock 110 kDa protein;
  • Gene Information: Name=HSPH1; Synonyms=HSP105, HSP110;
  • Organism (full): Cricetulus griseus (Chinese hamster) (Cricetulus barabensis griseus).
  • Protein Family: Belongs to the heat shock protein 70 family. .
  • Key Domains: ATPase_NBD. (IPR043129); Heat_shock_70_CS. (IPR018181); HSP70_C_sf. (IPR029048); HSP70_peptide-bd_sf. (IPR029047); Hsp_70_fam. (IPR013126)

MANDATORY VERIFICATION STEPS:

  1. Check if the gene symbol "HSPH1" matches the protein description above
  2. Verify the organism is correct: Cricetulus griseus (Chinese hamster) (Cricetulus barabensis griseus).
  3. Check if protein family/domains align with what you find in literature
  4. If you find literature for a DIFFERENT gene with the same or similar symbol, STOP

If Gene Symbol is Ambiguous or You Cannot Find Relevant Literature:

DO NOT PROCEED WITH RESEARCH ON A DIFFERENT GENE. Instead:
- State clearly: "The gene symbol 'HSPH1' is ambiguous or literature is limited for this specific protein"
- Explain what you found (e.g., "Found extensive literature on a different gene with the same symbol in a different organism")
- Describe the protein based ONLY on the UniProt information provided above
- Suggest that the protein function can be inferred from domain/family information

Research Target:

Please provide a comprehensive research report on the gene HSPH1 (gene ID: HSPH1, UniProt: Q60446) in CRIGR.

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.

Output

Question: You are an expert researcher providing comprehensive, well-cited information.

Provide detailed information focusing on:
1. Key concepts and definitions with current understanding
2. Recent developments and latest research (prioritize 2023-2024 sources)
3. Current applications and real-world implementations
4. Expert opinions and analysis from authoritative sources
5. Relevant statistics and data from recent studies

Format as a comprehensive research report with proper citations. Include URLs and publication dates where available.
Always prioritize recent, authoritative sources and provide specific citations for all major claims.

Gene Research for Functional Annotation

โš ๏ธ CRITICAL: Gene/Protein Identification Context

BEFORE YOU BEGIN RESEARCH: You MUST verify you are researching the CORRECT gene/protein. Gene symbols can be ambiguous, especially for less well-characterized genes from non-model organisms.

Target Gene/Protein Identity (from UniProt):

  • UniProt Accession: Q60446
  • Protein Description: RecName: Full=Heat shock protein 105 kDa; AltName: Full=Heat shock 110 kDa protein;
  • Gene Information: Name=HSPH1; Synonyms=HSP105, HSP110;
  • Organism (full): Cricetulus griseus (Chinese hamster) (Cricetulus barabensis griseus).
  • Protein Family: Belongs to the heat shock protein 70 family. .
  • Key Domains: ATPase_NBD. (IPR043129); Heat_shock_70_CS. (IPR018181); HSP70_C_sf. (IPR029048); HSP70_peptide-bd_sf. (IPR029047); Hsp_70_fam. (IPR013126)

MANDATORY VERIFICATION STEPS:

  1. Check if the gene symbol "HSPH1" matches the protein description above
  2. Verify the organism is correct: Cricetulus griseus (Chinese hamster) (Cricetulus barabensis griseus).
  3. Check if protein family/domains align with what you find in literature
  4. If you find literature for a DIFFERENT gene with the same or similar symbol, STOP

If Gene Symbol is Ambiguous or You Cannot Find Relevant Literature:

DO NOT PROCEED WITH RESEARCH ON A DIFFERENT GENE. Instead:
- State clearly: "The gene symbol 'HSPH1' is ambiguous or literature is limited for this specific protein"
- Explain what you found (e.g., "Found extensive literature on a different gene with the same symbol in a different organism")
- Describe the protein based ONLY on the UniProt information provided above
- Suggest that the protein function can be inferred from domain/family information

Research Target:

Please provide a comprehensive research report on the gene HSPH1 (gene ID: HSPH1, UniProt: Q60446) in CRIGR.

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.

Plan and verification
- Identity: The target is HSPH1 (aliases HSP110/HSP105), a large Hsp70-like chaperone; UniProt Q60446 corresponds to HSPH1 from Cricetulus griseus (Chinese hamster). Literature on mammalian HSPH1 describes the same protein class and domain architecture, supporting cross-species functional inference (Hsp70-like NBD and SBD) (boopathy2024characterizingtheregulationb pages 34-39, chuang2024definingasmallmolecule pages 5-8).

Comprehensive research report on HSPH1 (HSP110/HSP105)

1) Key concepts and definitions with current understanding
- Definition and family: HSPH1 encodes a large heat shock protein of the Hsp110 subfamily within the Hsp70 superfamily. It contains an Hsp70-like N-terminal nucleotide-binding domain (NBD/ATPase) and a substrate-binding domain (SBD) (conceptual alignment with Hsp70 architecture described in 2024 sources) (boopathy2024characterizingtheregulationb pages 34-39). Functionally, HSPH1 operates as a nucleotide-exchange factor (NEF) for Hsp70s, catalyzing ADP-to-ATP exchange on the Hsp70 NBD to reset the chaperone for subsequent substrate cycles (boopathy2024characterizingtheregulationb pages 34-39). URL/date: 2024 review summary (no DOI provided in excerpt) (boopathy2024characterizingtheregulationb pages 34-39).
- Mechanistic role in the Hsp70 system: As a NEF, HSPH1 binding to Hsp70โ€™s NBD accelerates nucleotide exchange; ATP rebinding opens Hsp70 and promotes client release. HSPH1 thereby increases Hsp70 folding throughput and participates in chaperone-mediated disaggregation with Hsp70 and J-domain proteins (DNAJs) (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 22-25). URL/date: Chuang et al., bioRxiv, Jan 2024, https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 22-25).
- Holdase-like properties: Under severe stress, HSPH1 has been described to help keep heat-denatured clients soluble and refolding-competent, complementing Hsp70 action (tandon2024athreedimensionalvalveonchip pages 12-14). URL/date: Tandon et al., Acta Biomaterialia, Sep 2024, https://doi.org/10.1016/j.actbio.2024.07.036 (tandon2024athreedimensionalvalveonchip pages 12-14).

2) Recent developments and latest research (2023โ€“2024 priority)
- Small-molecule modulation of the Hsp70โ€“HSPH1 disaggregase: A 2024 study defined dihydropyrimidine scaffolds that stimulate human Hsp70โ€“DNAJโ€“HSPH1 disaggregase activity. The reference compound 115-7c increased disaggregation approximately two-fold in vitro, and an optimized analog (โ€œcompound 18โ€) enhanced activity across a range of Hsp70/Hsp40/Hsp110 stoichiometries with selectivity for DnaJB1/DnaJB4-containing sets, without detectable cytotoxicity in cell assays (bioRxiv, Jan 2024) (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 25-29). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 25-29).
- Disaggregation mechanism and stoichiometry: The same 2024 work showed both human HSPH1 (Hsp105) and HSPH2 (Apg2) stimulate Hsp70-mediated disaggregation and reactivation of luciferase and can promote dissolution of disease-relevant amyloids (e.g., ฮฑ-synuclein), with outcomes depending on Hsp70/Hsp40/Hsp110 ratios; inappropriate stoichiometry can impair activity (bioRxiv, Jan 2024) (chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 22-25). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 22-25).
- System-level induction in neuronal heat stress: In human neuronal models, heat exposure induced coordinated upregulation of HSPH1 alongside DNAJB1 and BAG3 within the Hsp70 network, consistent with HSF1-driven heat shock response dynamics (Biology, Mar 2023) (alharbi2023profilingthehsp70 pages 17-18). URL/date: https://doi.org/10.3390/biology12030416 (alharbi2023profilingthehsp70 pages 17-18).
- Disease model observation (microphysiological system): In a 3D valve-on-chip model of early calcific aortic valve disease, HSPH1 abundance increased in diseased tissues, consistent with roles in proteostasis under stress and potential links to survival signaling pathways (Acta Biomaterialia, Sep 2024) (tandon2024athreedimensionalvalveonchip pages 12-14). URL/date: https://doi.org/10.1016/j.actbio.2024.07.036 (tandon2024athreedimensionalvalveonchip pages 12-14).

3) Primary function, substrates/clients, and localization
- Primary biochemical function: HSPH1 is a NEF for cytosolic Hsp70s (e.g., HSPA1A/HSPA8), binding the Hsp70 NBD to catalyze ADPโ†’ATP exchange; this resets Hsp70 to the open, low-affinity state, releasing bound substrate and enabling additional folding cycles (boopathy2024characterizingtheregulationb pages 34-39). This enhances Hsp70 folding activity and supports disaggregation (boopathy2024characterizingtheregulationb pages 34-39). URL/date: 2024 review summary (no DOI provided in excerpt) (boopathy2024characterizingtheregulationb pages 34-39).
- Disaggregation substrates: In reconstituted human systems, Hsp70โ€“DNAJโ€“HSPH1/2 can disaggregate heat-denatured luciferase and disease-associated amyloids including ฮฑ-synuclein; related studies cite activity on tau and polyQ-expanded proteins in Hsp70 disaggregase paradigms (bioRxiv, 2024; literature compilation) (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8, ansari2025investigatingtherole pages 139-140). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8); summary citing multi-article evidence (ansari2025investigatingtherole pages 139-140).
- Cellular localization: HSPH1 (Hsp105ฮฑ) localizes constitutively to cytoplasm and nucleus in multiple mammalian tissues, including brain; a stress-inducible Hsp105ฮฒ splice variant exists (not always tested in vitro) (chuang2024definingasmallmolecule pages 5-8). URL/date: bioRxiv, Jan 2024, https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 5-8).

4) Pathways, protein quality control, and organelle/stress compartment dynamics
- HSF1-driven heat shock response: HSPH1 is induced as part of the HSF1-regulated chaperone network during heat shock, with reported rapid transcript and delayed protein accumulation kinetics in neuronal heat-stress models (peak gene induction ~1 h; protein upregulated over the subsequent hours) (Biology, Mar 2023) (alharbi2023profilingthehsp70 pages 17-18). URL/date: https://doi.org/10.3390/biology12030416 (alharbi2023profilingthehsp70 pages 17-18).
- Hsp70 co-chaperone circuitry and fate decisions: BAG3 biases Hsp70 clients toward autophagy (aggrephagy), while CHIP (a U-box E3) promotes ubiquitination leading to proteasomal degradation; HSPH1, by accelerating Hsp70 cycling, functionally couples into these fate pathways and contributes to disaggregation upstream of BAG3- or CHIP-directed clearance (boopathy2024characterizingtheregulationb pages 34-39). URL/date: 2024 review summary (no DOI provided in excerpt) (boopathy2024characterizingtheregulationb pages 34-39).
- Stress granules and translation recovery: Hsp70 systems participate in stress granule disassembly and translation restart; HSPH1, as the principal cytosolic Hsp70 NEF in metazoans, supports this process by enabling efficient Hsp70 cycles under stress (boopathy2024characterizingtheregulationb pages 34-39). URL/date: 2024 review summary (no DOI provided in excerpt) (boopathy2024characterizingtheregulationb pages 34-39).
- Aggrephagy and proteasome/autophagy crosstalk: Mechanistic compendia indicate that Hsp70 disaggregation machinery (with HSPH1/NEF activity) intersects with proteolysis/autophagy pathways for aggregate processing. Summaries emphasize that Hsp110 shapes early steps of disaggregation by Hsp70 and that disaggregation can feed into proteolytic routing depending on co-chaperone context (compilation citing 2024โ€“2025 primary studies) (ansari2025investigatingtherole pages 139-140, ansari2025investigatingtherolea pages 139-140). URL/date: 2025 review citing 2024 mechanistic work (no DOI in excerpt) (ansari2025investigatingtherole pages 139-140, ansari2025investigatingtherolea pages 139-140).

5) Oncologic and disease correlations; expert analyses
- Cancer biology and signaling: Reviews and summaries link HSPH1 to survival signaling (e.g., NF-ฮบB) and proliferation under stress; HSPH1 can modulate pathways such as ฮฒ-catenin via PP2A recruitment and is reported to be phosphorylated by CK2 (site Ser509) (MDPI IJMS review, Jan 2024) (zhang2024heatshockproteins pages 19-19). URL/date: https://doi.org/10.3390/ijms25020876 (zhang2024heatshockproteins pages 19-19).
- Biomarker and therapeutic context: In the 3D valve-on-chip study, HSPH1 abundance increased in diseased samples, positioning it among proteostasis-linked changes associated with early disease processes (Acta Biomaterialia, Sep 2024) (tandon2024athreedimensionalvalveonchip pages 12-14). URL/date: https://doi.org/10.1016/j.actbio.2024.07.036 (tandon2024athreedimensionalvalveonchip pages 12-14). Additional oncology-focused compendia highlight emergent HSP110 inhibitors and roles in tumor stress adaptation and drug resistance (summary of recent studies) (ansari2025investigatingtherolea pages 139-140). URL/date: 2025 review citing 2020โ€“2024 primary works (no DOI in excerpt) (ansari2025investigatingtherolea pages 139-140).
- Expert network perspective: The 2024 small-molecule work emphasizes that chaperone stoichiometry is a critical determinant of disaggregase output in human cells and neurons, suggesting tissue-specific vulnerabilities if Hsp70/Hsp40/HSPH1 ratios drift with stress or aging (bioRxiv, Jan 2024) (chuang2024definingasmallmolecule pages 25-29). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 25-29).

6) Subcellular dynamics and stress-induced relocalization
- Cytosol and nucleus: Constitutive cytoplasmic and nuclear localization is reported for Hsp105ฮฑ (HSPH1) (bioRxiv, Jan 2024) (chuang2024definingasmallmolecule pages 5-8). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 5-8).
- Stress granules: Hsp70 systems contribute to stress granule disassembly and translational recovery after stress; by serving as the NEF for Hsp70, HSPH1 supports efficient chaperone cycling in these compartments (boopathy2024characterizingtheregulationb pages 34-39). URL/date: 2024 review summary (no DOI provided in excerpt) (boopathy2024characterizingtheregulationb pages 34-39).

7) Chinese hamster/CHO evidence and orthology-based inference
- Direct Cricetulus griseus/CHO data: We did not retrieve specific CHO/Hamster primary studies for HSPH1 within 2020โ€“2026 searches above. Given the strong conservation of Hsp110/Hsp70 systems and domain architecture, the mammalian evidence for HSPH1โ€™s NEF role, cytosolic/nuclear localization, and participation in disaggregation is highly likely to translate to Chinese hamster HSPH1 (UniProt Q60446) (boopathy2024characterizingtheregulationb pages 34-39, chuang2024definingasmallmolecule pages 5-8). Where species-specific findings are absent, conclusions are inferred by orthology and conserved domain function (boopathy2024characterizingtheregulationb pages 34-39, chuang2024definingasmallmolecule pages 5-8).

8) Current applications and real-world implementations; relevant statistics
- Therapeutic modulation: In vitro, the Hsp70โ€“DNAJโ€“HSPH1 disaggregase can be pharmacologically enhanced; the small molecule 115-7c produced about a two-fold increase in disaggregation activity against model substrates, and โ€œcompound 18โ€ further optimized activity across multiple chaperone combinations while remaining non-toxic in tested paradigms (bioRxiv, Jan 2024) (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 25-29). URL/date: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 25-29).
- Disease-model implementation: A 3D valve-on-chip platform modeling early calcific aortic valve disease reported increased HSPH1 abundance in diseased tissues, aligning with proteostasis stress signatures in pathogenesis (Acta Biomaterialia, Sep 2024) (tandon2024athreedimensionalvalveonchip pages 12-14). URL/date: https://doi.org/10.1016/j.actbio.2024.07.036 (tandon2024athreedimensionalvalveonchip pages 12-14).
- Cancer-focused expert analysis: Summaries from 2024โ€“2025 compilations describe HSPH1 as part of adaptive survival networks in tumors and discuss emerging HSP110 inhibitors in preclinical studies; while specific hazard ratios are not provided in the excerpts here, these sources position HSPH1 as a potential biomarker/target in oncology, warranting context- and tumor-type-specific validation (zhang2024heatshockproteins pages 19-19, ansari2025investigatingtherolea pages 139-140). URL/date: https://doi.org/10.3390/ijms25020876 (2024) (zhang2024heatshockproteins pages 19-19); 2025 review (no DOI in excerpt) (ansari2025investigatingtherolea pages 139-140).

Mechanistic summary and functional annotation (for HSPH1, Cricetulus griseus, UniProt Q60446)
- Molecular function: Hsp70 NEF activity that accelerates ADPโ†’ATP exchange on Hsp70 NBD to promote client release and increase folding/disaggregation throughput; large Hsp70-like chaperone with NBD and SBD (boopathy2024characterizingtheregulationb pages 34-39, chuang2024definingasmallmolecule pages 1-5).
- Biological processes: Protein folding; protein disaggregation with Hsp70 and J-domain proteins; maintenance of proteostasis under heat and proteotoxic stress; interface with client routing to autophagy (via BAG3) or UPS (via CHIP) (chuang2024definingasmallmolecule pages 1-5, boopathy2024characterizingtheregulationb pages 34-39).
- Cellular localization: Cytosol and nucleus under basal conditions; participates in stress responses influencing stress-granule dynamics and translation recovery (chuang2024definingasmallmolecule pages 5-8, boopathy2024characterizingtheregulationb pages 34-39).
- Pathways: Heat shock response (HSF1-regulated induction); Hsp70 chaperone cycle; proteostasis/quality control interconnections with autophagy and proteasome systems (alharbi2023profilingthehsp70 pages 17-18, boopathy2024characterizingtheregulationb pages 34-39, ansari2025investigatingtherole pages 139-140).
- Applications: Pharmacologic stimulation of Hsp70โ€“HSPH1 disaggregase; disease-model upregulation in microphysiological systems; oncology biomarker/target potential (chuang2024definingasmallmolecule pages 1-5, tandon2024athreedimensionalvalveonchip pages 12-14, zhang2024heatshockproteins pages 19-19, ansari2025investigatingtherolea pages 139-140).

Notes on evidence quality and recency
- 2024 primary preprint provides detailed mechanistic and pharmacologic insights (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 25-29). 2024 Acta Biomaterialia offers peer-reviewed disease-model evidence (tandon2024athreedimensionalvalveonchip pages 12-14). 2023 neuronal study corroborates network induction and timing under heat stress (alharbi2023profilingthehsp70 pages 17-18). Reviews/summaries integrate additional mechanistic connections and cancer relevance (boopathy2024characterizingtheregulationb pages 34-39, zhang2024heatshockproteins pages 19-19, ansari2025investigatingtherolea pages 139-140).

References (URLs and publication dates)
- Chuang et al., Defining a small-molecule stimulator of the human Hsp70-disaggregase system with selectivity for DnaJB proteins. bioRxiv, Jan 2024. URL: https://doi.org/10.1101/2024.01.11.575109 (chuang2024definingasmallmolecule pages 1-5, chuang2024definingasmallmolecule pages 5-8, chuang2024definingasmallmolecule pages 25-29, chuang2024definingasmallmolecule pages 22-25).
- Tandon et al., A three-dimensional valve-on-chip microphysiological system implicates cell cycle progression, cholesterol metabolism and protein homeostasis in early calcific aortic valve disease progression. Acta Biomaterialia, 186:167โ€“184, Sep 2024. URL: https://doi.org/10.1016/j.actbio.2024.07.036 (tandon2024athreedimensionalvalveonchip pages 12-14).
- Alharbi et al., Profiling the Hsp70 chaperone network in heat-induced proteotoxic stress models of human neurons. Biology, 12:416, Mar 2023. URL: https://doi.org/10.3390/biology12030416 (alharbi2023profilingthehsp70 pages 17-18).
- Boopathy, Characterizing the regulation of the inducible HSP70 mRNA in yeasts and mammalian cells subjected to heat shock. 2024 (mechanism summary; NEF/disaggregation; stress granules and BAG3/CHIP routing noted in excerpts) (boopathy2024characterizingtheregulationb pages 34-39).
- Zhang and Bi, Heat Shock Proteins and Breast Cancer. International Journal of Molecular Sciences, 25:876, Jan 2024. URL: https://doi.org/10.3390/ijms25020876 (zhang2024heatshockproteins pages 19-19).
- Ansari, Investigating the Role of Hsp110 in Prion Propagation and Fragmentation. 2025 (cites 2024 mechanistic studies on Hsp110 shaping early disaggregation steps and links to proteolytic pathways; used here as a compendium) (ansari2025investigatingtherole pages 139-140, ansari2025investigatingtherolea pages 139-140).

Species alignment statement
- No contradictions were found regarding HSPH1 naming or protein family/domain features. Where Cricetulus/CHO-specific data were absent, function/localization were inferred from conserved mammalian HSPH1 ortholog evidence and the shared Hsp70-like NBD/SBD domain architecture (boopathy2024characterizingtheregulationb pages 34-39, chuang2024definingasmallmolecule pages 5-8).

References

  1. (boopathy2024characterizingtheregulationb pages 34-39): LR Alagar Boopathy. Characterizing the regulation of the inducible hsp70 mrna in yeasts and mammalian cells subjected to heat shock. Unknown journal, 2024.

  2. (chuang2024definingasmallmolecule pages 5-8): Edward Chuang, Ryan R. Cupo, Jeffrey R. Bevan, Mikhaila L. Rice, Shuli Mao, Erica L. Gorenberg, Korrie L. Mack, Donna M. Huryn, Peter Wipf, Jeffrey L. Brodsky, and James Shorter. Defining a small-molecule stimulator of the human hsp70-disaggregase system with selectivity for dnajb proteins. bioRxiv, Jan 2024. URL: https://doi.org/10.1101/2024.01.11.575109, doi:10.1101/2024.01.11.575109. This article has 2 citations and is from a poor quality or predatory journal.

  3. (chuang2024definingasmallmolecule pages 1-5): Edward Chuang, Ryan R. Cupo, Jeffrey R. Bevan, Mikhaila L. Rice, Shuli Mao, Erica L. Gorenberg, Korrie L. Mack, Donna M. Huryn, Peter Wipf, Jeffrey L. Brodsky, and James Shorter. Defining a small-molecule stimulator of the human hsp70-disaggregase system with selectivity for dnajb proteins. bioRxiv, Jan 2024. URL: https://doi.org/10.1101/2024.01.11.575109, doi:10.1101/2024.01.11.575109. This article has 2 citations and is from a poor quality or predatory journal.

  4. (chuang2024definingasmallmolecule pages 22-25): Edward Chuang, Ryan R. Cupo, Jeffrey R. Bevan, Mikhaila L. Rice, Shuli Mao, Erica L. Gorenberg, Korrie L. Mack, Donna M. Huryn, Peter Wipf, Jeffrey L. Brodsky, and James Shorter. Defining a small-molecule stimulator of the human hsp70-disaggregase system with selectivity for dnajb proteins. bioRxiv, Jan 2024. URL: https://doi.org/10.1101/2024.01.11.575109, doi:10.1101/2024.01.11.575109. This article has 2 citations and is from a poor quality or predatory journal.

  5. (tandon2024athreedimensionalvalveonchip pages 12-14): Ishita Tandon, Alan E. Woessner, Laฮฏs A. Ferreira, Christine Shamblin, Gustavo Vaca-Diez, Amanda Walls, Patrick Kuczwara, Alexis Applequist, Denise F. Nascimento, Swastika Tandon, Jin-Woo Kim, Manuel Rausch, Tomasz Timek, Muralidhar Padala, Michael T. Kinter, Dennis Province, Stephanie D. Byrum, Kyle P. Quinn, and Kartik Balachandran. A three-dimensional valve-on-chip microphysiological system implicates cell cycle progression, cholesterol metabolism and protein homeostasis in early calcific aortic valve disease progression. Acta Biomaterialia, 186:167-184, Sep 2024. URL: https://doi.org/10.1016/j.actbio.2024.07.036, doi:10.1016/j.actbio.2024.07.036. This article has 6 citations and is from a domain leading peer-reviewed journal.

  6. (chuang2024definingasmallmolecule pages 25-29): Edward Chuang, Ryan R. Cupo, Jeffrey R. Bevan, Mikhaila L. Rice, Shuli Mao, Erica L. Gorenberg, Korrie L. Mack, Donna M. Huryn, Peter Wipf, Jeffrey L. Brodsky, and James Shorter. Defining a small-molecule stimulator of the human hsp70-disaggregase system with selectivity for dnajb proteins. bioRxiv, Jan 2024. URL: https://doi.org/10.1101/2024.01.11.575109, doi:10.1101/2024.01.11.575109. This article has 2 citations and is from a poor quality or predatory journal.

  7. (alharbi2023profilingthehsp70 pages 17-18): Bothina Mohammed Alharbi, Tahani H. Albinhassan, Razan Ali Alzahrani, Abderrezak Bouchama, Sameer Mohammad, Awatif Abdulaziz Alomari, May Nasser Bin-Jumah, Entissar S. AlSuhaibani, and Shuja Shafi Malik. Profiling the hsp70 chaperone network in heat-induced proteotoxic stress models of human neurons. Biology, 12:416, Mar 2023. URL: https://doi.org/10.3390/biology12030416, doi:10.3390/biology12030416. This article has 7 citations and is from a poor quality or predatory journal.

  8. (ansari2025investigatingtherole pages 139-140): M Ansari. Investigating the role of hsp110 in prion propagation and fragmentation. Unknown journal, 2025.

  9. (ansari2025investigatingtherolea pages 139-140): M Ansari. Investigating the role of hsp110 in prion propagation and fragmentation. Unknown journal, 2025.

  10. (zhang2024heatshockproteins pages 19-19): Miao Zhang and Xiaowen Bi. Heat shock proteins and breast cancer. International Journal of Molecular Sciences, 25:876, Jan 2024. URL: https://doi.org/10.3390/ijms25020876, doi:10.3390/ijms25020876. This article has 38 citations and is from a poor quality or predatory journal.

Citations

  1. boopathy2024characterizingtheregulationb pages 34-39
  2. tandon2024athreedimensionalvalveonchip pages 12-14
  3. ansari2025investigatingtherole pages 139-140
  4. chuang2024definingasmallmolecule pages 5-8
  5. zhang2024heatshockproteins pages 19-19
  6. ansari2025investigatingtherolea pages 139-140
  7. chuang2024definingasmallmolecule pages 25-29
  8. chuang2024definingasmallmolecule pages 1-5
  9. chuang2024definingasmallmolecule pages 22-25
  10. https://doi.org/10.1101/2024.01.11.575109
  11. https://doi.org/10.1016/j.actbio.2024.07.036
  12. https://doi.org/10.3390/biology12030416
  13. https://doi.org/10.3390/ijms25020876
  14. https://doi.org/10.1101/2024.01.11.575109,
  15. https://doi.org/10.1016/j.actbio.2024.07.036,
  16. https://doi.org/10.3390/biology12030416,
  17. https://doi.org/10.3390/ijms25020876,

๐Ÿ“„ View Raw YAML

id: Q60446
gene_symbol: HSPH1
product_type: PROTEIN
status: DRAFT
taxon:
  id: NCBITaxon:10029
  label: Cricetulus griseus
description: >-
  HSPH1 (Hsp105/Hsp110) is a member of the Hsp110 family, a significantly diverged subgroup
  of the Hsp70 superfamily. It has two core molecular functions: (1) holdase chaperone activity,
  preventing the aggregation of heat-denatured proteins by maintaining them in a soluble,
  folding-competent state (significantly more efficient than Hsc70 at this function), and
  (2) nucleotide exchange factor (NEF) activity for Hsp70 family members (HSPA1A, HSPA1B),
  promoting ADP release and triggering substrate release. Importantly, HSPH1 does NOT itself
  refold denatured proteins; refolding requires Hsc70/Hdj-1. HSPH1 also inhibits HSPA8/HSC70
  ATPase activity. Overexpression confers substantial thermoresistance in vivo.
existing_annotations:
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IEA
  original_reference_id: GO_REF:0000118
  review:
    summary: >-
      TreeGrafter-inferred nuclear localization. UniProt subcellular location for Q60446
      lists only cytoplasm (by similarity to human Q92598). However, the deep research
      review (HSPH1-deep-research-falcon.md) notes that Hsp105alpha (HSPH1) localizes
      constitutively to both cytoplasm and nucleus in multiple mammalian tissues,
      including brain. The TreeGrafter inference is therefore supported by recent
      literature on mammalian orthologs.
    action: KEEP_AS_NON_CORE
    reason: >-
      Nuclear localization is supported by literature on mammalian HSPH1 orthologs.
      The deep research review cites Chuang et al. 2024 as reporting constitutive
      cytoplasmic and nuclear localization. However, the primary functional
      compartment is the cytosol where the holdase and NEF activities operate.
      Keeping as non-core since it is a secondary localization.
    additional_reference_ids:
      - file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md
    supported_by:
      - reference_id: file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md
        supporting_text: >-
          Constitutive cytoplasmic and nuclear localization is reported for Hsp105alpha
          (HSPH1) [in mammalian tissues]

- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IEA
  original_reference_id: GO_REF:0000118
  review:
    summary: >-
      TreeGrafter-inferred cytosol localization, consistent with UniProt annotation of
      cytoplasm for HSPH1 (by similarity to human Q92598). HSPH1 is a cytoplasmic chaperone
      and its holdase and NEF functions operate in the cytoplasm. The deep research review
      (HSPH1-deep-research-falcon.md) confirms cytosolic localization as the primary site
      of HSPH1 function.
    action: ACCEPT
    reason: >-
      Cytosol is the primary localization for HSPH1, consistent with UniProt annotation
      of cytoplasm and with its known functions as a cytoplasmic chaperone and NEF.
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is one of the principal molecular chaperones of mammalian cells

- term:
    id: GO:0006457
    label: protein folding
  evidence_type: IEA
  original_reference_id: GO_REF:0000118
  review:
    summary: >-
      TreeGrafter-inferred involvement in protein folding. HSPH1 is involved in protein
      folding homeostasis, but importantly it is a holdase, not a foldase. It maintains
      denatured proteins in a folding-competent state but does NOT actively refold them
      (PMID:9395504). Refolding requires Hsc70/Hdj-1. The term 'protein folding' is
      acceptable as a broad biological process annotation since HSPH1 participates in
      the protein folding pathway (by handing off substrates to the Hsc70/Hdj-1 refolding
      machinery), but its direct activity is holding, not folding. As a NEF, HSPH1 also
      accelerates Hsp70 cycling which supports the overall protein folding process (deep
      research review, HSPH1-deep-research-falcon.md).
    action: KEEP_AS_NON_CORE
    reason: >-
      HSPH1 participates in the protein folding pathway by preventing aggregation and
      maintaining substrates in a folding-competent state for subsequent refolding by
      Hsc70/Hdj-1. The BP term 'protein folding' is broadly acceptable but HSPH1 is a
      holdase, not a foldase, so this is kept as non-core. Its direct role is prevention
      of aggregation rather than active folding.
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is highly efficient in selectively recognizing denatured proteins and
          maintaining them in a soluble, folding-competent state
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110-bound proteins can then be refolded by the addition of rabbit
          reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself
          function as a co-chaperone in folding with hsp110

- term:
    id: GO:0000166
    label: nucleotide binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000043
  review:
    summary: >-
      IEA annotation from UniProtKB-KW:KW-0547 (Nucleotide-binding) keyword mapping.
      HSPH1 belongs to the Hsp70 superfamily and possesses the conserved nucleotide-binding
      domain (NBD). This is correct but overly general; ATP binding (GO:0005524) is more
      specific and is already annotated.
    action: ACCEPT
    reason: >-
      Nucleotide binding is correct for HSPH1, which has a conserved Hsp70-type
      nucleotide-binding domain. While more general than the co-annotated GO:0005524
      (ATP binding), the IEA mapping from UniProt keyword is valid. Both annotations
      can coexist since the IEA is broader.

- term:
    id: GO:0005524
    label: ATP binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: >-
      Combined IEA annotation from InterPro:IPR013126 (Hsp_70_fam) and UniProtKB-KW:KW-0067
      (ATP-binding). HSPH1 has a conserved Hsp70-type nucleotide-binding domain and binds ATP.
      While Hsp110 family members have greatly reduced ATPase activity compared to canonical
      Hsp70s, they do bind ATP, which is important for their NEF function.
    action: ACCEPT
    reason: >-
      ATP binding is a core property of HSPH1. The Hsp110 family binds ATP through
      their conserved NBD. UniProt keywords and InterPro domain classification both
      support this annotation. ATP binding is required for the NEF function of HSPH1.

- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  review:
    summary: >-
      IEA annotation from UniProtKB subcellular location vocabulary mapping (SL-0086).
      UniProt explicitly states cytoplasm localization for HSPH1 (by similarity to human
      Q92598). This is the primary localization and is consistent with its chaperone/NEF
      functions.
    action: ACCEPT
    reason: >-
      Cytoplasm is the primary subcellular localization for HSPH1. UniProt annotation
      (by similarity) and the protein's established cytoplasmic functions support this.
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is one of the principal molecular chaperones of mammalian cells

- term:
    id: GO:0006950
    label: response to stress
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  review:
    summary: >-
      ARBA-inferred involvement in response to stress. HSPH1 is a heat shock protein
      whose expression is upregulated under stress conditions. UniProt keywords include
      'Stress response'. This is correct but very general; the more specific term
      GO:0034605 (cellular response to heat) is already annotated with IDA evidence.
    action: KEEP_AS_NON_CORE
    reason: >-
      Response to stress is valid for this heat shock protein but is very general.
      The more specific IDA-supported annotation to GO:0034605 (cellular response to
      heat) captures the core stress response function. Keeping this broader IEA
      annotation as non-core since it adds no information beyond the specific IDA
      annotation.

- term:
    id: GO:0016887
    label: ATP hydrolysis activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  review:
    summary: >-
      InterPro-inferred ATP hydrolysis activity from IPR013126 (Hsp_70_fam). While HSPH1
      belongs to the Hsp70 superfamily and has the conserved ATPase domain architecture,
      Hsp110 family members have greatly reduced intrinsic ATPase activity compared to
      canonical Hsp70 chaperones. The deep research review (HSPH1-deep-research-falcon.md)
      emphasizes that HSPH1 functions primarily as a NEF rather than an ATPase. The ATPase
      annotation from the broad Hsp70 family InterPro entry is misleading for this
      diverged subfamily.
    action: MARK_AS_OVER_ANNOTATED
    reason: >-
      Hsp110 family members have significantly reduced ATPase activity compared to
      canonical Hsp70 proteins. The InterPro family IPR013126 covers the entire Hsp70
      superfamily, and while the domain architecture is conserved, Hsp110 members have
      diverged to function primarily as holdases and NEFs rather than as ATPases.
      Annotating HSPH1 with GO:0016887 (ATP hydrolysis activity) overstates its
      catalytic function. The core molecular functions are holdase chaperone activity
      and NEF activity (GO:0000774).

- term:
    id: GO:0031249
    label: denatured protein binding
  evidence_type: IDA
  original_reference_id: PMID:9395504
  review:
    summary: >-
      IDA annotation based on Oh et al. 1997, which demonstrated that Hsp110 is highly
      efficient at recognizing heat-denatured proteins and maintaining them in a soluble,
      folding-competent state. This is a genuine holdase chaperone function. GO:0031249
      is being obsoleted (go-ontology#30962). The appropriate replacement is GO:0044183
      (protein folding chaperone) as a placeholder, with the caveat that HSPH1 is a
      holdase, not a foldase, and a more specific holdase term is needed
      (go-ontology#30552).
    action: MODIFY
    reason: >-
      GO:0031249 (denatured protein binding) is being obsoleted. The experimental
      evidence from PMID:9395504 clearly demonstrates holdase chaperone activity:
      Hsp110 prevents aggregation of heat-denatured proteins and maintains them in
      a folding-competent state. GO:0044183 (protein folding chaperone) is the closest
      available replacement MF term, used as a placeholder until a specific holdase
      chaperone activity term is created (go-ontology#30552). Note that HSPH1 does NOT
      refold proteins itself; refolding requires Hsc70/Hdj-1.
    proposed_replacement_terms:
      - id: GO:0044183
        label: protein folding chaperone
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is highly efficient in selectively recognizing denatured proteins and
          maintaining them in a soluble, folding-competent state and is significantly more
          efficient in performing this function than is hsc70
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110-bound proteins can then be refolded by the addition of rabbit
          reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself
          function as a co-chaperone in folding with hsp110

- term:
    id: GO:0034605
    label: cellular response to heat
  evidence_type: IDA
  original_reference_id: PMID:9395504
  review:
    summary: >-
      IDA annotation based on Oh et al. 1997, which demonstrated that overexpression
      of Hsp110 in vivo conferred substantial heat resistance to both Rat-1 and HeLa
      cells. This is a core biological process for HSPH1 as a heat shock protein.
    action: ACCEPT
    reason: >-
      Well-supported by direct experimental evidence from PMID:9395504 showing that
      Hsp110 overexpression confers thermoresistance. HSPH1 is one of the primary heat
      shock proteins in mammalian cells, and cellular response to heat is a core
      biological process.
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          the overexpression in vivo of hsp110 conferred substantial heat resistance
          to both Rat-1 and HeLa cells
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is one of the principal molecular chaperones of mammalian cells and
          represents a newly identified component of the primary protection/repair
          pathway for denatured proteins and thermotolerance expression in vivo

- term:
    id: GO:0000774
    label: adenyl-nucleotide exchange factor activity
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: >-
      ISS annotation transferred from human HSPH1 (Q92598). UniProt describes HSPH1
      as acting as a nucleotide-exchange factor (NEF) for HSPA1A and HSPA1B, promoting
      the release of ADP thereby triggering substrate release (by similarity to mouse
      Q61699 and human Q92598). This is one of the two core molecular functions of
      Hsp110 family members. The deep research review (HSPH1-deep-research-falcon.md)
      confirms that HSPH1 operates as a NEF for cytosolic Hsp70s, binding the Hsp70
      NBD to catalyze ADP-to-ATP exchange.
    action: ACCEPT
    reason: >-
      NEF activity is a core molecular function of the Hsp110 family. While the evidence
      for Chinese hamster HSPH1 specifically is by similarity (ISS), the NEF function is
      well-conserved across mammalian Hsp110 orthologs and is supported by UniProt
      annotation (by similarity to Q61699 mouse and Q92598 human). This represents one
      of the two principal molecular functions of HSPH1 (the other being holdase activity).
    additional_reference_ids:
      - file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md

# New annotations not in the existing GOA set

- term:
    id: GO:0044183
    label: protein folding chaperone
  evidence_type: IDA
  original_reference_id: PMID:9395504
  review:
    summary: >-
      Proposed new annotation as the replacement MF term for the obsoleting GO:0031249.
      PMID:9395504 provides direct experimental evidence that Hsp110 functions as a
      chaperone that prevents aggregation of denatured proteins and maintains them in
      a folding-competent state. This is a holdase chaperone function (placeholder
      annotation pending creation of a specific holdase term per go-ontology#30552).
    action: NEW
    reason: >-
      This annotation replaces the obsoleting GO:0031249 (denatured protein binding)
      with the closest available MF term. HSPH1 is a holdase chaperone that binds
      denatured proteins to prevent aggregation. GO:0044183 is a placeholder until
      a specific holdase chaperone activity term is created (go-ontology#30552).
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is highly efficient in selectively recognizing denatured proteins and
          maintaining them in a soluble, folding-competent state

references:
- id: GO_REF:0000002
  title: Gene Ontology annotation through association of InterPro records with GO
    terms
  findings: []
- id: GO_REF:0000024
  title: Manual transfer of experimentally-verified manual GO annotation data to orthologs
    by curator judgment of sequence similarity
  findings: []
- id: GO_REF:0000043
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot keyword mapping
  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:0000117
  title: Electronic Gene Ontology annotations created by ARBA machine learning models
  findings: []
- id: GO_REF:0000118
  title: TreeGrafter-generated GO annotations
  findings: []
- id: GO_REF:0000120
  title: Combined Automated Annotation using Multiple IEA Methods
  findings: []
- id: PMID:9395504
  title: Hsp110 protects heat-denatured proteins and confers cellular thermoresistance.
  findings:
    - statement: >-
        Hsp110 is highly efficient at preventing aggregation of heat-denatured proteins,
        functioning as a holdase chaperone that maintains substrates in a folding-competent
        state. It is significantly more efficient than Hsc70 at this holdase function.
      supporting_text: >-
        hsp110 is highly efficient in selectively recognizing denatured proteins and
        maintaining them in a soluble, folding-competent state and is significantly more
        efficient in performing this function than is hsc70
    - statement: >-
        Hsp110 does not refold denatured proteins itself; refolding requires Hsc70 and
        Hdj-1. Hdj-1 does not function as a co-chaperone with Hsp110.
      supporting_text: >-
        hsp110-bound proteins can then be refolded by the addition of rabbit
        reticulocyte lysate or hsc70 and Hdj-1, whereas Hdj-1 does not itself
        function as a co-chaperone in folding with hsp110
    - statement: >-
        Overexpression of Hsp110 confers substantial thermoresistance in vivo.
      supporting_text: >-
        the overexpression in vivo of hsp110 conferred substantial heat resistance
        to both Rat-1 and HeLa cells
- id: PMID:7797574
  title: Identification of a major subfamily of large hsp70-like proteins through
    the cloning of the mammalian 110-kDa heat shock protein.
  findings:
    - statement: >-
        HSPH1 (Hsp110) was identified as a member of a significantly diverged subgroup
        of the Hsp70 protein family.
      supporting_text: >-
        Identification of a major subfamily of large hsp70-like proteins through the
        cloning of the mammalian 110-kDa heat shock protein
      reference_section_type: TITLE
- id: file:CRIGR/HSPH1/HSPH1-deep-research-falcon.md
  title: Deep research review of HSPH1 function
  findings:
    - statement: >-
        HSPH1 operates as a NEF for cytosolic Hsp70s, catalyzing ADP-to-ATP exchange
        on the Hsp70 NBD to reset the chaperone for subsequent substrate cycles.
    - statement: >-
        HSPH1 participates in the Hsp70-DNAJ-HSPH1 disaggregase complex that can
        disaggregate heat-denatured luciferase and disease-associated amyloids.
    - statement: >-
        Constitutive cytoplasmic and nuclear localization is reported for Hsp105alpha
        (HSPH1) in mammalian tissues.
core_functions:
  - description: >-
      Holdase chaperone activity: HSPH1 binds heat-denatured proteins and prevents their
      aggregation, maintaining them in a soluble, folding-competent state. This function
      is significantly more efficient than Hsc70. HSPH1 does NOT actively refold proteins;
      refolding requires Hsc70/Hdj-1. This is a placeholder annotation using GO:0044183
      until a specific holdase chaperone activity term is created (go-ontology#30552).
    molecular_function:
      id: GO:0044183
      label: protein folding chaperone
    directly_involved_in:
      - id: GO:0034605
        label: cellular response to heat
    locations:
      - id: GO:0005829
        label: cytosol
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is highly efficient in selectively recognizing denatured proteins and
          maintaining them in a soluble, folding-competent state
  - description: >-
      Nucleotide exchange factor (NEF) activity for Hsp70 family members: HSPH1 promotes
      the release of ADP from HSPA1A/HSPA1B, triggering substrate release from the Hsp70
      chaperone cycle. Also inhibits HSPA8/HSC70 ATPase and chaperone activities.
    molecular_function:
      id: GO:0000774
      label: adenyl-nucleotide exchange factor activity
    directly_involved_in:
      - id: GO:0034605
        label: cellular response to heat
    locations:
      - id: GO:0005829
        label: cytosol
    supported_by:
      - reference_id: PMID:9395504
        supporting_text: >-
          hsp110 is one of the principal molecular chaperones of mammalian cells and
          represents a newly identified component of the primary protection/repair
          pathway for denatured proteins and thermotolerance expression in vivo
proposed_new_terms:
  - proposed_name: holdase chaperone activity
    proposed_definition: >-
      Binding of unfolded or misfolded proteins to prevent their aggregation, without
      actively catalyzing refolding. This activity maintains client proteins in a soluble,
      folding-competent state for subsequent refolding by other chaperone systems.
    justification: >-
      Multiple proteins including HSPH1/Hsp110, small HSPs (CRYAA, CRYAB, HSPB6),
      and clusterin (CLU) function as holdases rather than foldases. The current
      GO term GO:0044183 (protein folding chaperone) implies active folding, which
      does not accurately describe holdase function. See go-ontology#30552.
    proposed_parent:
      id: GO:0044183
      label: protein folding chaperone
suggested_questions:
  - question: >-
      Does Chinese hamster HSPH1 have the same NEF activity for Hsp70 as the mouse
      and human orthologs? The ISS annotation is based on similarity but has not been
      directly demonstrated for the Chinese hamster protein.
    experts:
      - Subjeck JR
  - question: >-
      What is the intrinsic ATPase activity level of HSPH1? While Hsp110 family members
      are known to have reduced ATPase activity compared to canonical Hsp70, the exact
      level for Chinese hamster HSPH1 has not been characterized.
    experts:
      - Subjeck JR
suggested_experiments:
  - hypothesis: >-
      Chinese hamster HSPH1 functions as a NEF for hamster Hsp70 family members,
      similar to the characterized function of mouse and human orthologs.
    description: >-
      In vitro nucleotide exchange assay using purified Chinese hamster HSPH1 and
      Hsp70 (HSPA1A). Measure ADP release rates from Hsp70 in the presence and
      absence of HSPH1 using fluorescent nucleotide analogs.
    experiment_type: biochemical assay
  - hypothesis: >-
      HSPH1 has significantly lower intrinsic ATPase activity compared to canonical
      Hsp70 family members.
    description: >-
      Comparative ATPase assay measuring the rate of ATP hydrolysis by purified
      HSPH1 versus HSPA1A using a malachite green phosphate detection assay.
    experiment_type: biochemical assay