STIP1

UniProt ID: P31948
Organism: Homo sapiens
Review Status: COMPLETE
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Gene Description

STIP1 (stress-induced phosphoprotein 1), better known as HOP (Hsp70-Hsp90 organizing protein) or p60, is a cytoplasmic (and partly nuclear) TPR-domain adaptor co-chaperone. It contains three TPR (tetratricopeptide repeat) domains. The TPR1 domain binds the C-terminal EEVD motif of HSP70 (HSPA8/HSC70), while the TPR2A and TPR2B domains bind the C-terminal MEEVD motif of HSP90. By simultaneously engaging both chaperones, HOP physically bridges the HSP70 and HSP90 systems and coordinates the transfer of client proteins from HSP70 to HSP90 during the chaperone cycle. HOP has no catalytic activity; it functions as a scaffold/adaptor that modulates HSP90 conformation and client maturation. It is a defining component of the HSP70-HSP90 multichaperone machine and participates in the maturation of diverse clients including protein kinases and steroid hormone receptors.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0051879 Hsp90 protein binding
IBA
GO_REF:0000033
ACCEPT
Summary: HOP binds HSP90 directly via its TPR2A/TPR2B domains (HSP90 C-terminal MEEVD). This is a core molecular function.
Reason: Directly supported by UniProt FUNCTION/DOMAIN and by experimental HSP90 interaction data; HSP90 binding is central to HOP's adaptor role.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
The TPR 4, 5 and 6 repeats (also called TPR2A domain) and TPR 7, 8 and 9 repeats (also called TPR2B domain) interact with HSP90.
GO:0005634 nucleus
IEA
GO_REF:0000120
KEEP AS NON CORE
Summary: HOP is detected in the nucleus in addition to the cytoplasm; nuclear shuttling is documented.
Reason: UniProt lists Nucleus as a subcellular location; nuclear pool is real but HOP's principal chaperone-organizing function is cytoplasmic.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005737 cytoplasm
IEA
GO_REF:0000044
ACCEPT
Summary: HOP is predominantly cytoplasmic, where it organizes the HSP70-HSP90 machine.
Reason: Matches UniProt subcellular location; cytoplasm is the genuine principal compartment of HOP.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0120293 dynein axonemal particle
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: By-similarity localization to the dynein axonemal particle, reflecting a role of HOP/co-chaperones in cytoplasmic preassembly of axonemal dynein in ciliated cells.
Reason: Specialized, by-similarity localization in ciliated-cell contexts; peripheral to HOP's general cytoplasmic chaperone-organizing function.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Dynein axonemal particle
GO:0005515 protein binding
IPI
PMID:20029029
Regulation of epidermal growth factor receptor trafficking b...
KEEP AS NON CORE
Summary: Interaction with EGFR (P00533), an HSP90 client kinase. Bare protein binding is uninformative; the interaction is consistent with HOP's role in client maturation but is a generic binding term.
Reason: Records a real interaction with an HSP90 client (EGFR), but bare protein binding is uninformative and not elevated to core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:21044950
Genome-wide YFP fluorescence complementation screen identifi...
KEEP AS NON CORE
Summary: High-throughput interaction (partner Q96AP0). Bare protein binding is uninformative and the partner is not part of HOP's core chaperone function.
Reason: Records a real interaction but bare protein binding is uninformative; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:21170051
Mixed Hsp90-cochaperone complexes are important for the prog...
MODIFY
Summary: Interaction with HSP90AB1 (P08238). Bare protein binding is uninformative; this HSP90 interaction is better captured as Hsp90 protein binding.
Reason: The WITH partner is HSP90AB1 (P08238); the interaction is precisely captured as Hsp90 protein binding, a core HOP function.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:21360678
Label-free quantitative proteomics and SAINT analysis enable...
KEEP AS NON CORE
Summary: Interaction with PPP5C (P53041), a TPR-domain HSP90 co-chaperone phosphatase. Bare protein binding is uninformative; the partner is chaperone-machinery-related.
Reason: Real interaction with a co-chaperone (PPP5C) within the HSP90 machine, but bare protein binding is uninformative; not elevated to core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:25036637
A quantitative chaperone interaction network reveals the arc...
MODIFY
Summary: Quantitative chaperone interaction network capturing HOP with HSP90AB1 (P08238) and CDC37L1 (Q7L3B6) among others. Bare protein binding is uninformative; the meaningful partner is HSP90.
Reason: WITH partners include HSP90AB1 (P08238); the chaperone-network interaction is precisely captured as Hsp90 protein binding.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:28330616
Systematic Analysis of Human Protein Phosphatase Interaction...
KEEP AS NON CORE
Summary: Interaction with PPP5C (P53041), an HSP90 co-chaperone. Bare protein binding is uninformative.
Reason: Real interaction with a chaperone-machinery component but uninformative protein binding term; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:28514442
Architecture of the human interactome defines protein commun...
MODIFY
Summary: Binary interactome capturing HOP with HSP90AB1 (P08238) and another partner. Bare protein binding is uninformative; the meaningful partner is HSP90.
Reason: WITH partner HSP90AB1 (P08238); precisely captured as Hsp90 protein binding.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:31980649
Extensive rewiring of the EGFR network in colorectal cancer ...
KEEP AS NON CORE
Summary: Interaction with EGFR (P00533) and ERBB2 (P04626), HSP90 client kinases. Bare protein binding is uninformative; consistent with HOP's client-maturation role.
Reason: Records real interactions with HSP90 client kinases, but bare protein binding is uninformative and not elevated to core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:32814053
Interactome Mapping Provides a Network of Neurodegenerative ...
KEEP AS NON CORE
Summary: Large neurodegeneration interactome capturing many partners (e.g. MYC, p53), none being core chaperones. Bare protein binding is uninformative.
Reason: High-throughput interactome partners unrelated to HOP's chaperone-organizing function; uninformative protein binding.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
KEEP AS NON CORE
Summary: BioPlex interactome capturing HOP interactions (partners O15484, Q8IWD4). Bare protein binding is uninformative and these are not core chaperone partners.
Reason: Records real high-throughput interactions but uninformative protein binding; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:35140242
Human transcription factor protein interaction networks.
KEEP AS NON CORE
Summary: Interaction with p53 (P04637). Bare protein binding is uninformative; p53 is an HSP90 client but this is a generic binding annotation.
Reason: Real interaction with an HSP90 client (p53), but bare protein binding is uninformative; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:35271311
OpenCell: Endogenous tagging for the cartography of human ce...
MODIFY
Summary: Chaperome interaction-landscape study capturing HOP with HSP90AB1 (P08238). Bare protein binding is uninformative; the meaningful partner is HSP90.
Reason: WITH partner HSP90AB1 (P08238); precisely captured as Hsp90 protein binding.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:40205054
Multimodal cell maps as a foundation for structural and func...
KEEP AS NON CORE
Summary: Multimodal cell-maps interactome (partner Q8IWD4). Bare protein binding is uninformative.
Reason: Records a real high-throughput interaction but uninformative protein binding; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005634 nucleus
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Sequence-similarity-based nuclear localization, consistent with HOP's documented nuclear pool.
Reason: Consistent with UniProt Nucleus location; nuclear pool is real but non-core relative to the cytoplasmic chaperone-organizing role.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005737 cytoplasm
ISS
GO_REF:0000024
ACCEPT
Summary: Sequence-similarity-based cytoplasm localization, consistent with HOP's principal compartment.
Reason: Matches UniProt subcellular location; cytoplasm is the genuine principal compartment.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0120293 dynein axonemal particle
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Sequence-similarity-based localization to the dynein axonemal particle (ciliated-cell dynein preassembly context).
Reason: Specialized by-similarity localization; peripheral to HOP's general chaperone-organizing function.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Dynein axonemal particle
GO:0051879 Hsp90 protein binding
IPI
PMID:29127155
Tumor suppressor Tsc1 is a new Hsp90 co-chaperone that facil...
ACCEPT
Summary: Experimental evidence for HOP binding HSP90 within the HSP70-HSP90 multichaperone complex. Core molecular function.
Reason: Directly supported; HOP is a defining HSP90-binding component of the multichaperone machine described in this study and in UniProt SUBUNIT.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Probably forms a complex composed of chaperones HSP90 and HSP70, co-chaperones STIP1/HOP, CDC37, PPP5C, PTGES3/p23, TSC1 and client protein TSC2
GO:0101031 protein folding chaperone complex
IDA
PMID:29127155
Tumor suppressor Tsc1 is a new Hsp90 co-chaperone that facil...
ACCEPT
Summary: HOP is part of the HSP70-HSP90 multichaperone complex, a folding chaperone complex. This is an accurate and core complex annotation.
Reason: Directly supported by UniProt SUBUNIT (HSP90/HSP70/STIP1/CDC37/PPP5C/p23/TSC1 complex); HOP is a defining organizing subunit.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Probably forms a complex composed of chaperones HSP90 and HSP70, co-chaperones STIP1/HOP, CDC37, PPP5C, PTGES3/p23, TSC1 and client protein TSC2
GO:0005515 protein binding
IPI
PMID:23349634
A newly uncovered group of distantly related lysine methyltr...
KEEP AS NON CORE
Summary: Interaction with the lysine methyltransferase EEF1AKMT3 (Q96AZ1). Bare protein binding is uninformative.
Reason: Records a real interaction but bare protein binding is uninformative; not core.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Interacts with EEF1AKMT3
GO:0032991 protein-containing complex
IDA
PMID:23349634
A newly uncovered group of distantly related lysine methyltr...
KEEP AS NON CORE
Summary: Generic protein-containing complex annotation from a methyltransferase study. Less informative than the specific HSP70-HSP90 multichaperone complex term.
Reason: Generic complex term; the informative complex annotation is GO:0101031 (HSP70-HSP90 multichaperone complex).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Interacts with EEF1AKMT3
GO:0005515 protein binding
IPI
PMID:24880080
SMYD2-dependent HSP90 methylation promotes cancer cell proli...
MODIFY
Summary: Interaction with HSP90AB1 (P08238), modulated by SMYD2-dependent HSP90AB1 methylation. Bare protein binding is uninformative; the meaningful partner is HSP90.
Reason: WITH partner HSP90AB1 (P08238); precisely captured as Hsp90 protein binding.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Interacts with HSP90AB1; upon SMYD2-dependent HSP90AB1 methylation
GO:0005515 protein binding
IPI
PMID:23431407
Distinct roles of molecular chaperones HSP90Ξ± and HSP90Ξ² in ...
KEEP AS NON CORE
Summary: Interaction with partner P56696. Bare protein binding is uninformative and not part of HOP's core chaperone function.
Reason: Records a real interaction but bare protein binding is uninformative; not core.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0005515 protein binding
GO:0005515 protein binding
IPI
PMID:27353360
The FNIP co-chaperones decelerate the Hsp90 chaperone cycle ...
MODIFY
Summary: Interaction with HSP90AA1 (P07900) and the FLCN/FNIP co-chaperone module. Bare protein binding is uninformative; the meaningful partner is HSP90.
Reason: WITH partner HSP90AA1 (P07900); precisely captured as Hsp90 protein binding, a core HOP function.
Proposed replacements: Hsp90 protein binding
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
Acts as a co-chaperone for HSP90AA1
GO:0005829 cytosol
TAS
Reactome:R-HSA-3371503
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-5618085
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-5618098
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-5618105
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-5618107
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-5618110
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005829 cytosol
TAS
Reactome:R-HSA-9696271
ACCEPT
Summary: Curated (Reactome) cytosolic localization, consistent with HOP's cytoplasmic chaperone role.
Reason: Cytosol is the genuine principal compartment of HOP; consistent with UniProt (one of several redundant Reactome cytosol annotations).
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0003723 RNA binding
HDA
PMID:22658674
Insights into RNA biology from an atlas of mammalian mRNA-bi...
MARK AS OVER ANNOTATED
Summary: mRNA-interactome-capture detection of HOP as an RNA-binder. No characterized RNA-dependent function for HOP exists.
Reason: High-throughput RNA-interactome capture without a validated functional role; not part of HOP's chaperone-organizing function.
Supporting Evidence:
file:human/STIP1/STIP1-goa.tsv
GO:0003723 RNA binding
GO:0005634 nucleus
TAS
PMID:16130169
Proteomics of human umbilical vein endothelial cells applied...
KEEP AS NON CORE
Summary: Curated nuclear localization of HOP, consistent with its documented nuclear pool.
Reason: Nuclear pool is real (UniProt lists Nucleus) but non-core relative to the cytoplasmic chaperone-organizing role.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005634 nucleus
TAS
PMID:1569099
Molecular cloning and expression of a transformation-sensiti...
KEEP AS NON CORE
Summary: Early curated nuclear localization of HOP/p60.
Reason: Consistent with the documented nuclear pool; non-core relative to the cytoplasmic function.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm
GO:0005794 Golgi apparatus
TAS
PMID:1569099
Molecular cloning and expression of a transformation-sensiti...
KEEP AS NON CORE
Summary: Early curated Golgi localization of HOP/p60. Not corroborated by the current UniProt subcellular-location record (cytoplasm/nucleus/dynein axonemal particle).
Reason: Historical localization not supported as a principal compartment in current UniProt; peripheral to HOP's cytoplasmic chaperone function.
Supporting Evidence:
file:human/STIP1/STIP1-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm

Core Functions

Hsp70-Hsp90 organizing protein (HOP); a TPR-domain adaptor co-chaperone that binds HSP90 (via TPR2A/TPR2B) and bridges it to HSP70/HSC70 (via TPR1), coordinating transfer of client proteins from HSP70 to HSP90.

Molecular Function:
Hsp90 protein binding
Cellular Locations:
Supporting Evidence:
  • file:human/STIP1/STIP1-uniprot.txt
    Mediates the association of the molecular chaperones HSPA8/HSC70 and HSP90
  • file:human/STIP1/STIP1-uniprot.txt
    The TPR 4, 5 and 6 repeats (also called TPR2A domain) and TPR 7, 8 and 9 repeats (also called TPR2B domain) interact with HSP90.

Component of the HSP70-HSP90 multichaperone machine that organizes client maturation; HOP acts as a scaffold without catalytic activity, holding HSP90 in a client-loading-competent state.

Molecular Function:
Hsp90 protein binding
Cellular Locations:
Supporting Evidence:
  • file:human/STIP1/STIP1-uniprot.txt
    Probably forms a complex composed of chaperones HSP90 and HSP70, co-chaperones STIP1/HOP, CDC37, PPP5C, PTGES3/p23, TSC1 and client protein TSC2

References

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Suggested Questions for Experts

Q: How is the HOP-mediated HSP70-to-HSP90 client handoff regulated in human cells, and is HOP strictly required or can clients transfer via HOP-independent routes (as suggested in some organisms)?

Q: What determines the cytoplasmic versus nuclear partitioning of HOP, and does the nuclear pool have a chaperone-independent function?

Q: Do the alternative HOP isoforms (P31948-2, P31948-3) differ in TPR-domain composition and thus in HSP70/HSP90 binding or client specificity?

Suggested Experiments

Experiment: Reconstitute the HSP70-HOP-HSP90 client-transfer reaction in vitro with a model client (e.g. a kinase or steroid receptor) and TPR-domain point mutants to dissect the HSP70-binding (TPR1) versus HSP90-binding (TPR2A/2B) contributions to handoff.

Experiment: HOP knockout/knockdown followed by client stability profiling (kinases, steroid receptors) to quantify HOP dependence of the HSP90 clientele in human cells.

Experiment: Crosslinking-MS or cryo-EM of the human HSP70-HOP-HSP90 intermediate to define the architecture of the client-loading complex.

πŸ“š Additional Documentation

Notes

(STIP1-notes.md)

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Pn Notes

(STIP1-pn-notes.md)

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πŸ“„ View Raw YAML

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