LRP4

UniProt ID: O75096
Organism: Homo sapiens
Review Status: COMPLETE
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Gene Description

LRP4 is a 1,905-residue type-I cell-surface receptor with a large extracellular LDL-receptor-family repeat region, one transmembrane helix, and a cytoplasmic tail. At the neuromuscular junction, its ectodomain binds neuronal AGRIN and MuSK and assembles them into a signaling complex that activates MuSK and promotes acetylcholine-receptor clustering and synapse formation. In osteoblast-lineage cells, LRP4 directly binds the secreted inhibitor sclerostin and facilitates its suppression of canonical Wnt signaling and bone formation. Human variants that perturb different surfaces of the third beta-propeller selectively disrupt these neuromuscular or bone-regulatory functions, causing congenital myasthenic syndrome, Cenani-Lenz syndactyly syndrome, or sclerosteosis-like bone overgrowth.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005886 plasma membrane
IBA
GO_REF:0000033
ACCEPT
Summary: IBA inference associates LRP4 with plasma membrane; exact ordered source provenance and any live annotation extensions are retained.
Reason: Plasma-membrane localization is core to LRP4's type-I membrane coreceptor architecture and is independently consistent with the reviewed human record and human agrin-LRP4-MuSK structure.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
FB:FBgn0030706 SUPPORTS TRANSFER
FB:FBgn0051092 SUPPORTS TRANSFER
MGI:MGI:1340044 SUPPORTS TRANSFER
MGI:MGI:1860083 SUPPORTS TRANSFER
MGI:MGI:2138856 SUPPORTS TRANSFER
MGI:MGI:2442252 SUPPORTS TRANSFER
MGI:MGI:96765 SUPPORTS TRANSFER
MGI:MGI:96828 SUPPORTS TRANSFER
PANTHER:PTN002557696 SUPPORTS TRANSFER
RGD:2998 SUPPORTS TRANSFER
RGD:619731 SUPPORTS TRANSFER
RGD:727887 SUPPORTS TRANSFER
UniProtKB:E1BGJ0 SUPPORTS TRANSFER
UniProtKB:P01130 SUPPORTS TRANSFER
UniProtKB:P01131 SUPPORTS TRANSFER
UniProtKB:P98155 SUPPORTS TRANSFER
UniProtKB:Q07954 SUPPORTS TRANSFER
UniProtKB:Q14114 SUPPORTS TRANSFER
UniProtKB:Q86YD5 SUPPORTS TRANSFER
UniProtKB:Q9NPF0 SUPPORTS TRANSFER
UniProtKB:Q9Y561 SUPPORTS TRANSFER
UniProtKB:Q9Y5Q5 SUPPORTS TRANSFER
WB:WBGene00003071 SUPPORTS TRANSFER
WB:WBGene00015083 SUPPORTS TRANSFER
WB:WBGene00019811 SUPPORTS TRANSFER
GO:0048699 generation of neurons
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: IBA inference associates LRP4 with generation of neurons; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
PANTHER:PTN000056094 SUPPORTS TRANSFER
RGD:619731 SUPPORTS TRANSFER
GO:0050808 synapse organization
IBA
GO_REF:0000033
ACCEPT
Summary: IBA inference associates LRP4 with synapse organization; exact ordered source provenance and any live annotation extensions are retained.
Reason: Synapse organization is a defining conserved LRP4 role, especially in neuromuscular-junction assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
MGI:MGI:2442252 SUPPORTS TRANSFER
PANTHER:PTN000056094 SUPPORTS TRANSFER
RGD:619731 SUPPORTS TRANSFER
GO:0090090 negative regulation of canonical Wnt signaling pathway
IBA
GO_REF:0000033
ACCEPT
Summary: IBA inference associates LRP4 with negative regulation of canonical Wnt signaling pathway; exact ordered source provenance and any live annotation extensions are retained.
Reason: Negative regulation of canonical Wnt signaling is supported by direct human genetics and SOST-facilitator experiments as well as the curated phylogeny.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
MGI:MGI:2442252 SUPPORTS TRANSFER
PANTHER:PTN000056094 SUPPORTS TRANSFER
UniProtKB:O75096 SUPPORTS TRANSFER
Target self-WITH/FROM is expected IBA provenance and reflects target experimental grounding; it is not circular.
GO:0005509 calcium ion binding
IEA
GO_REF:0000002
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with calcium ion binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: The molecular property is compatible with LRP4 but is structural, generic, or ligand-contextual rather than the receptor's defining coreceptor activity.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
InterPro:IPR001881 SUPPORTS TRANSFER
InterPro:IPR018097 SUPPORTS TRANSFER
GO:0005886 plasma membrane
IEA
GO_REF:0000120
ACCEPT
Summary: IEA inference associates LRP4 with plasma membrane; exact ordered source provenance and any live annotation extensions are retained.
Reason: Plasma-membrane localization is core to LRP4's single-pass receptor architecture.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
UniProtKB-SubCell:SL-0039 SUPPORTS TRANSFER
GO:0046718 symbiont entry into host cell
IEA
GO_REF:0000108
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with symbiont entry into host cell; exact ordered source provenance and any live annotation extensions are retained.
Reason: Yellow-fever-virus entry is experimentally supported for human LRP4, but pathogen exploitation is not the receptor’s core endogenous function.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
GO:0001618 SOURCE WEAK OR INFERRED
This logical-inference source is an ontology term, not a gene-level experimental donor.
GO:0048731 system development
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with system development; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
ARBA:ARBA00027702 SOURCE WEAK OR INFERRED
The ARBA rule is electronic provenance rather than direct experimental evidence.
GO:0160108 animal gross anatomical part developmental process
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with animal gross anatomical part developmental process; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
ARBA:ARBA00098139 SOURCE WEAK OR INFERRED
The ARBA rule is electronic provenance rather than direct experimental evidence.
GO:0005515 protein binding
IPI
PMID:12421765
Protein-protein interactions between large proteins: two-hyb...
UNDECIDED
Summary: PMID:12421765 provides curator-assessed IPI evidence relating LRP4 to protein binding.
Reason: The cached abstract describes a large-protein yeast-two-hybrid screen but does not identify the LRP4-WHRN pair or establish which WHRN domain binds. Under curator-deference rules the generic IPI annotation remains UNDECIDED.
GO:0005515 protein binding
IPI
PMID:21471202
Bone overgrowth-associated mutations in the LRP4 gene impair...
MODIFY
Summary: PMID:21471202 provides curator-assessed IPI evidence relating LRP4 to protein binding.
Reason: The direct LRP4-SOST interaction is informative, but generic protein binding obscures the demonstrated facilitator role. Current GO:0015026 coreceptor activity captures LRP4 cooperation in SOST-dependent inhibition of Wnt signaling.
Proposed replacements: coreceptor activity
Supporting Evidence:
PMID:21471202
Biochemical assays with recombinant proteins confirmed that sclerostin LRP4 interaction is direct. Interestingly, in vitro overexpression and RNAi-mediated knockdown experiments revealed that LRP4 specifically facilitates the previously described inhibitory action of sclerostin on Wnt1/Ξ²-catenin signaling.
GO:0005515 protein binding
IPI
PMID:28514442
Architecture of the human interactome defines protein commun...
KEEP AS NON CORE
Summary: PMID:28514442 provides curator-assessed IPI evidence relating LRP4 to protein binding.
Reason: The curator-made BioPlex IPI is retained as human AP-MS co-association evidence, and targeted PMID:21471202 independently supports LRP4-SOST binding. The screen row itself does not establish coreceptor activity, so generic binding remains non-core rather than being promoted to that functional term.
Supporting Evidence:
PMID:21471202
Biochemical assays with recombinant proteins confirmed that sclerostin LRP4 interaction is direct. Interestingly, in vitro overexpression and RNAi-mediated knockdown experiments revealed that LRP4 specifically facilitates the previously described inhibitory action of sclerostin on Wnt1/Ξ²-catenin signaling.
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
KEEP AS NON CORE
Summary: PMID:33961781 provides curator-assessed IPI evidence relating LRP4 to protein binding.
Reason: The curator-made BioPlex IPI is retained as human AP-MS co-association evidence, and targeted PMID:21471202 independently supports LRP4-SOST binding. The screen row itself does not establish coreceptor activity, so generic binding remains non-core rather than being promoted to that functional term.
Supporting Evidence:
PMID:21471202
Biochemical assays with recombinant proteins confirmed that sclerostin LRP4 interaction is direct. Interestingly, in vitro overexpression and RNAi-mediated knockdown experiments revealed that LRP4 specifically facilitates the previously described inhibitory action of sclerostin on Wnt1/Ξ²-catenin signaling.
GO:0005515 protein binding
IPI
PMID:36115835
Quantitative fragmentomics allow affinity mapping of interac...
MODIFY
Summary: PMID:36115835 provides curator-assessed IPI evidence relating LRP4 to protein binding.
Reason: This quantitative fragmentomics study measured interactions between the LRP4 C-terminal PDZ-binding motif and an ordered panel of PDZ-domain proteins. Current GO:0030165 PDZ domain binding captures the common mechanism more specifically than generic protein binding.
Proposed replacements: PDZ domain binding
GO:0007167 enzyme-linked receptor protein signaling pathway
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with enzyme-linked receptor protein signaling pathway; exact ordered source provenance and any live annotation extensions are retained.
Reason: LRP4 is an extracellular coreceptor for the MUSK receptor tyrosine kinase; this broad pathway term captures a core signaling role without assigning kinase activity to LRP4.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0009986 cell surface
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with cell surface; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0014069 postsynaptic density
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with postsynaptic density; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0015026 coreceptor activity
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with coreceptor activity; exact ordered source provenance and any live annotation extensions are retained.
Reason: Coreceptor activity is a defining molecular role of LRP4 in the AGRIN-LRP4-MUSK signaling complex and in SOST-dependent Wnt inhibition.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0030425 dendrite
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with dendrite; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0030971 receptor tyrosine kinase binding
IEA
GO_REF:0000120
ACCEPT
Summary: IEA inference associates LRP4 with receptor tyrosine kinase binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: Binding the receptor tyrosine kinase MUSK is a defining molecular interaction in LRP4-dependent neuromuscular-junction assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0031594 neuromuscular junction
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with neuromuscular junction; exact ordered source provenance and any live annotation extensions are retained.
Reason: The neuromuscular junction is a defining site of LRP4 action.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0034185 apolipoprotein binding
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA inference associates LRP4 with apolipoprotein binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: This electronic transfer traces only to inferred rat Lrp4 records and no primary rat or human LRP4 apolipoprotein-binding evidence was identified. The LDL-receptor-family fold does not establish the multiligand lipoprotein-binding repertoire of LRP1 or LRP2 in LRP4.
Propagation Review
Root cause: PROPAGATION BAD
Failure modes: SOURCE EVIDENCE WEAK
Sources checked:
UniProtKB:Q9QYP1 SOURCE WEAK OR INFERRED
ensembl:ENSRNOP00000021353 SOURCE WEAK OR INFERRED
GO:0035022 positive regulation of Rac protein signal transduction
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with positive regulation of Rac protein signal transduction; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0042803 protein homodimerization activity
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: IEA inference associates LRP4 with protein homodimerization activity; exact ordered source provenance and any live annotation extensions are retained.
Reason: The reviewed human record only transfers a nonspecific homooligomer statement by similarity, and the electronic rat sources do not establish a dimer stoichiometry or a dimer-specific molecular activity for human LRP4. The term is therefore more specific than the accessible evidence.
Propagation Review
Root cause: TERM SCOPING PROBLEM
Failure modes: GRANULARITY MISMATCH
Sources checked:
UniProtKB:Q8VI56 SOURCE WEAK OR INFERRED
ensembl:ENSMUSP00000028689 SOURCE WEAK OR INFERRED
GO:0043025 neuronal cell body
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with neuronal cell body; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0044853 plasma membrane raft
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with plasma membrane raft; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0045202 synapse
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with synapse; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0048813 dendrite morphogenesis
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with dendrite morphogenesis; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0050771 negative regulation of axonogenesis
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with negative regulation of axonogenesis; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0050808 synapse organization
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with synapse organization; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0051124 synaptic assembly at neuromuscular junction
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with synaptic assembly at neuromuscular junction; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0071340 skeletal muscle acetylcholine-gated channel clustering
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with skeletal muscle acetylcholine-gated channel clustering; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0097060 synaptic membrane
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with synaptic membrane; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0097104 postsynaptic membrane assembly
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with postsynaptic membrane assembly; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0097105 presynaptic membrane assembly
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with presynaptic membrane assembly; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0097110 scaffold protein binding
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: IEA inference associates LRP4 with scaffold protein binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: Scaffold-protein binding is plausible in neuronal LRP4 biology but is secondary to its core extracellular coreceptor activity.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
ensembl:ENSRNOP00000021353 SUPPORTS TRANSFER
GO:0150052 regulation of postsynapse assembly
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with regulation of postsynapse assembly; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:1901631 positive regulation of presynaptic membrane organization
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with positive regulation of presynaptic membrane organization; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:1904395 positive regulation of skeletal muscle acetylcholine-gated channel clustering
IEA
GO_REF:0000107
ACCEPT
Summary: IEA inference associates LRP4 with positive regulation of skeletal muscle acetylcholine-gated channel clustering; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
ensembl:ENSMUSP00000028689 SUPPORTS TRANSFER
GO:0001618 virus receptor activity
IMP
PMID:41162706
Multiple LDLRΒ family members act as entry receptors for yell...
KEEP AS NON CORE
Summary: PMID:41162706 provides curator-assessed IMP evidence relating LRP4 to virus receptor activity.
Reason: The 2026 study directly establishes LRP4-dependent yellow-fever-virus entry, but this is pathogen exploitation rather than an endogenous core function.
GO:0005886 plasma membrane
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with plasma membrane; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0150094 amyloid-beta clearance by cellular catabolic process
IMP
PMID:18289866
LRP promotes endocytosis and degradation, but not transcytos...
REMOVE
Summary: PMID:18289866 provides curator-assessed IMP evidence relating LRP4 to amyloid-beta clearance by cellular catabolic process.
Reason: The paper explicitly defines β€œmLRP4” as an LRP mini-receptor in an LRP amyloid-beta experiment; it does not study the LRP4 gene product O75096. This is a verified homonym/construct misattribution.
Supporting Evidence:
PMID:18289866
MDCK cells were stably transfected with Pgp or mLRP4, an LRP mini-receptor.
GO:0044853 plasma membrane raft
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with plasma membrane raft; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0009986 cell surface
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with cell surface; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0014069 postsynaptic density
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with postsynaptic density; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0030971 receptor tyrosine kinase binding
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with receptor tyrosine kinase binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0031594 neuromuscular junction
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with neuromuscular junction; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0051124 synaptic assembly at neuromuscular junction
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with synaptic assembly at neuromuscular junction; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0097110 scaffold protein binding
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with scaffold protein binding; exact ordered source provenance and any live annotation extensions are retained.
Reason: The molecular property is compatible with LRP4 but is structural, generic, or ligand-contextual rather than the receptor's defining coreceptor activity.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0030425 dendrite
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with dendrite; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0043025 neuronal cell body
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with neuronal cell body; exact ordered source provenance and any live annotation extensions are retained.
Reason: This localization is biologically compatible but is a secondary neuronal subcompartment or broad contextual location rather than the defining plasma-membrane/neuromuscular-junction site.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0097060 synaptic membrane
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with synaptic membrane; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0050771 negative regulation of axonogenesis
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with negative regulation of axonogenesis; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0048813 dendrite morphogenesis
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ISS inference associates LRP4 with dendrite morphogenesis; exact ordered source provenance and any live annotation extensions are retained.
Reason: This is a broad developmental, neuronal, cargo-specific, or downstream context rather than LRP4's core molecular role.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0050808 synapse organization
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with synapse organization; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q9QYP1 SUPPORTS TRANSFER
GO:0071340 skeletal muscle acetylcholine-gated channel clustering
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with skeletal muscle acetylcholine-gated channel clustering; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0097104 postsynaptic membrane assembly
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with postsynaptic membrane assembly; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0097105 presynaptic membrane assembly
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with presynaptic membrane assembly; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0001822 kidney development
IDA
PMID:20381006
LRP4 mutations alter Wnt/beta-catenin signaling and cause li...
KEEP AS NON CORE
Summary: PMID:20381006 provides curator-assessed IDA evidence relating LRP4 to kidney development.
Reason: Human recessive LRP4 variants establish a developmental requirement associated with kidney malformations; this organismal phenotype is important but non-core relative to the receptor’s molecular role.
GO:0009986 cell surface
IDA
PMID:20381006
LRP4 mutations alter Wnt/beta-catenin signaling and cause li...
ACCEPT
Summary: PMID:20381006 provides curator-assessed IDA evidence relating LRP4 to cell surface.
Reason: Cell-surface localization is consistent with LRP4's experimentally established type-I membrane topology and receptor functions; curator evidence is accepted despite the abstract not detailing the localization assay.
GO:0060173 limb development
IDA
PMID:20381006
LRP4 mutations alter Wnt/beta-catenin signaling and cause li...
KEEP AS NON CORE
Summary: PMID:20381006 provides curator-assessed IDA evidence relating LRP4 to limb development.
Reason: Human LRP4 variants establish an essential role in limb development, but this pleiotropic developmental outcome is non-core.
GO:0090090 negative regulation of canonical Wnt signaling pathway
IMP
PMID:20381006
LRP4 mutations alter Wnt/beta-catenin signaling and cause li...
ACCEPT
Summary: PMID:20381006 provides curator-assessed IMP evidence relating LRP4 to negative regulation of canonical Wnt signaling pathway.
Reason: The human genetics study explicitly states that LRP4 antagonizes LRP6-mediated canonical Wnt activation and that patient mutations lose this function.
GO:1901631 positive regulation of presynaptic membrane organization
ISS
GO_REF:0000024
ACCEPT
Summary: ISS inference associates LRP4 with positive regulation of presynaptic membrane organization; exact ordered source provenance and any live annotation extensions are retained.
Reason: The annotation is consistent with LRP4's conserved core role as a cell-surface organizer/coreceptor for AGRIN-MUSK signaling and synapse assembly.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q8VI56 SUPPORTS TRANSFER
GO:0030279 negative regulation of ossification
IMP
PMID:21471202
Bone overgrowth-associated mutations in the LRP4 gene impair...
KEEP AS NON CORE
Summary: PMID:21471202 provides curator-assessed IMP evidence relating LRP4 to negative regulation of ossification.
Reason: The SOST-facilitator study supports inhibition of bone mineralization/ossification, but this tissue-level output is non-core.
GO:0090090 negative regulation of canonical Wnt signaling pathway
IMP
PMID:21471202
Bone overgrowth-associated mutations in the LRP4 gene impair...
ACCEPT
Summary: PMID:21471202 provides curator-assessed IMP evidence relating LRP4 to negative regulation of canonical Wnt signaling pathway.
Reason: Direct interaction, perturbation, and disease-variant evidence support LRP4-mediated negative regulation of canonical Wnt signaling.
GO:0090090 negative regulation of canonical Wnt signaling pathway
IDA
PMID:20093106
The low-density lipoprotein receptor-related protein 10 is a...
ACCEPT
Summary: PMID:20093106 provides curator-assessed IDA evidence relating LRP4 to negative regulation of canonical Wnt signaling pathway.
Reason: The cached abstract explicitly centers the distinct LRP10 protein and does not expose the LRP4 experiment seen by the curator. Nevertheless, direct human genetic and perturbation evidence in PMID:20381006 and PMID:21471202 independently establishes this exact LRP4 function, so the term is accepted while retaining the citation caveat rather than overruling the curator.
Supporting Evidence:
PMID:20093106
Here, we found that LRP10, a new member of the LDLR gene family, inhibits the canonical Wnt/beta-catenin signaling pathway.
PMID:20381006
By sequencing candidate genes, we identified recessive LRP4 mutations in 12 families with CLS. LRP4 belongs to the low-density lipoprotein (LDL) receptor-related proteins (LRPs), which are essential for various developmental processes. LRP4 is known to antagonize LRP6-mediated activation of canonical Wnt signaling, a function that is lost by the identified mutations.

Core Functions

At the skeletal-muscle neuromuscular junction, LRP4 functions as the extracellular coreceptor that binds neuronal AGRIN and MuSK and brings them into a ternary signaling complex. This enables AGRIN-dependent MuSK activation, acetylcholine-receptor clustering, and coordinated pre- and postsynaptic assembly. Mammalian cell and muscle experiments establish the mechanism, while a human extracellular-fragment cryo-EM structure directly resolves the 1:1:1 complex and human congenital-myasthenia variants impair AGRIN and MuSK binding. The evidence does not make LRP4 a kinase or establish that its cytoplasmic tail supplies the catalytic signal.

Supporting Evidence:
  • PMID:18957220
    report that LRP4, a low-density lipoprotein receptor (LDLR)-related protein, is expressed specifically in myotubes and binds to neuronal agrin. Its expression enables agrin binding and MuSK signaling in cells that otherwise do not respond to agrin. Suppression of LRP4 expression in muscle cells attenuates agrin binding, agrin-induced MuSK tyrosine phosphorylation, and AChR clustering. LRP4 also forms a complex with MuSK in a manner that is stimulated by agrin.
  • PMID:37252960
    Here, we report the cryo-EM structure of the extracellular ternary complex of agrin/LRP4/MuSK in a stoichiometry of 1:1:1. This structure reveals that arc-shaped LRP4 simultaneously recruits both agrin and MuSK to its central cavity, thereby promoting a direct interaction between agrin and MuSK.
  • PMID:24234652
    The identified mutations in LRP4 are located at the edge of its 3rd beta-propeller domain and decrease binding affinity of LRP4 for both MuSK and agrin. Mutations in the LRP4 3rd beta-propeller domain were previously reported to impair Wnt signaling and cause bone diseases including Cenani-Lenz syndactyly syndrome and sclerosteosis-2. By analyzing naturally occurring and artificially introduced mutations in the LRP4 3rd beta-propeller domain, we show that the edge of the domain regulates the MuSK signaling whereas its central cavity governs Wnt signaling.

In osteoblast-lineage cells, plasma-membrane LRP4 binds sclerostin and acts as a facilitator/coreceptor for its inhibition of canonical Wnt signaling. Direct recombinant-protein binding, LRP4 overexpression and knockdown, and human bone-overgrowth variants support this pathway. Mouse Lrp4-DKK1 and bone phenotypes provide additional in-vivo context, but the direct human evidence is strongest for sclerostin and should not be generalized to every Wnt antagonist or LDLR-family ligand.

Supporting Evidence:
  • PMID:21471202
    Biochemical assays with recombinant proteins confirmed that sclerostin LRP4 interaction is direct. Interestingly, in vitro overexpression and RNAi-mediated knockdown experiments revealed that LRP4 specifically facilitates the previously described inhibitory action of sclerostin on Wnt1/Ξ²-catenin signaling.
  • PMID:21471202
    Notably, we identified two mutations in LRP4 (R1170W and W1186S) in patients suffering from bone overgrowth. We found that these mutations impair LRP4 interaction with sclerostin and its concomitant sclerostin facilitator effect.
  • PMID:20381006
    By sequencing candidate genes, we identified recessive LRP4 mutations in 12 families with CLS. LRP4 belongs to the low-density lipoprotein (LDL) receptor-related proteins (LRPs), which are essential for various developmental processes. LRP4 is known to antagonize LRP6-mediated activation of canonical Wnt signaling, a function that is lost by the identified mutations.

References

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Suggested Questions for Experts

Q: How does full-length membrane-embedded human LRP4 couple the extracellular AGRIN-LRP4-MuSK assembly to MuSK activation and synaptic organization?

Q: How do distinct surfaces of LRP4 beta-propeller 3 select AGRIN/MuSK versus sclerostin/Wnt pathway partners in different tissues?

Q: Which PDZ-domain proteins bind the full-length LRP4 cytoplasmic tail endogenously, and do any regulate neuromuscular or bone signaling?

Q: Does human LRP4 directly facilitate DKK1 signaling in osteoblasts, or is its established human Wnt-inhibitory role specific to sclerostin?

Q: Does LRP4 undergo regulated cleavage, endocytosis, or recycling that changes the balance between its AGRIN/MuSK and sclerostin/Wnt functions?

Suggested Experiments

Experiment: Endogenously tag LRP4 and MuSK in human myotubes and combine ligand-timed single-molecule imaging, phosphoproteomics, and AChR-clustering assays with rescue by interface-specific LRP4 variants.

Hypothesis: AGRIN stabilizes a full-length LRP4-MuSK assembly whose geometry, rather than an intrinsic LRP4 catalytic activity, drives MuSK activation and AChR clustering.

Type: endogenous neuromuscular receptor-complex analysis

Experiment: Determine a membrane-context structure of full-length human LRP4 with AGRIN and MuSK by cryo-electron tomography or single-particle cryo-EM, including the transmembrane segments and cytoplasmic tails.

Hypothesis: Membrane anchoring and unresolved LRP4 regions impose additional geometry on the AGRIN-LRP4-MuSK complex that is absent from the soluble 21-1350 ectodomain structure.

Type: full-length membrane-protein structural analysis

Experiment: Introduce pathway-selective third-propeller variants into isogenic human osteoblasts and myotubes, then compare sclerostin/Wnt inhibition, AGRIN/MuSK activation, and receptor surface abundance.

Hypothesis: Spatially distinct third-propeller surfaces independently encode LRP4's bone and neuromuscular functions rather than causing a shared folding defect.

Type: separation-of-function variant analysis

Experiment: Validate candidate PDZ-tail partners by endogenous co-immunoprecipitation and proximity labeling in human myotubes and osteoblasts, followed by partner knockout and rescue assays in each signaling context.

Hypothesis: Only a subset of peptide-binding PDZ proteins forms physiological LRP4 complexes, with tissue-specific effects on receptor localization or signaling.

Type: endogenous cytoplasmic-tail interactome validation

Knowledge Gaps

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

Gap: The architecture and signaling behavior of full-length membrane-embedded human LRP4 in the AGRIN-LRP4-MuSK complex remain unresolved.

OPEN BIOLOGY MF_DARK

What is known: The human cryo-EM study used LRP4 residues 21-1350, and substantial extracellular regions, the transmembrane helix, and the entire cytoplasmic tail are absent or unresolved. It establishes extracellular partner recruitment, not the complete receptor activation mechanism.

Significance: A full-length structure is needed to connect extracellular coreceptor assembly to membrane organization and intracellular MuSK activation.

What would resolve it: Resolve a membrane-context full-length human AGRIN-LRP4-MuSK complex and test structural predictions with interface-selective rescue assays.

Provenance (the field's own admissions):

Gap: Direct endogenous human-neuromuscular evidence for LRP4 complex dynamics and signaling is limited.

OPEN BIOLOGY BP_DARK

What is known: Foundational functional experiments use mammalian cell lines and mouse muscle, while human evidence comes from a soluble-fragment structure and disease variants. These sources strongly support conserved mechanism but do not directly visualize full-length LRP4 signaling at a human neuromuscular junction.

Significance: Human endogenous measurements are needed to define complex lifetime, pathway stoichiometry, and variant effects in the native synaptic context.

What would resolve it: Use patient-derived or engineered human neuromuscular cocultures for endogenous complex imaging, MuSK phosphosignaling, and AChR-clustering rescue.

Gap: The molecular basis for pathway-selective effects of LRP4 beta-propeller variants is incomplete.

OPEN BIOLOGY RESIDUAL_SUBGAP

What is known: Human variants and mutagenesis distinguish a MuSK-regulating edge from a Wnt-regulating central cavity, but folding, trafficking, partner occupancy, and tissue-specific effects have not been compared in one system.

Significance: Resolving this distinction is essential for interpreting disease variants without assuming that every allele disrupts both core pathways.

What would resolve it: Compare surface expression, structure, AGRIN/MuSK binding, sclerostin binding, and downstream signaling for a systematic variant panel in matched human cell models.

Provenance (the field's own admissions):

Gap: The physiological LRP4 cytoplasmic-tail interactome and its contribution to either core pathway are unknown.

OPEN BIOLOGY MF_DARK

What is known: Fragmentomics supports PDZ-domain binding by an LRP4 tail peptide, but the assay does not establish that every partner binds full-length LRP4 in cells or regulates neuromuscular or bone signaling.

Significance: Partner validation is required before assigning specific scaffold complexes or downstream tail-mediated processes to LRP4.

What would resolve it: Test endogenous full-length LRP4 complexes and partner-specific functional effects in human myotubes and osteoblasts.

πŸ“š Additional Documentation

Notes

(LRP4-notes.md)

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