LRG1

UniProt ID: P02750
Organism: Homo sapiens
Review Status: COMPLETE
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Gene Description

LRG1 is a secreted, extensively glycosylated leucine-rich-repeat protein produced as a 347-amino-acid precursor with a cleaved signal peptide (residues 1-35) and an eight-LRR mature chain (residues 36-347). It circulates in plasma and is produced prominently by hepatocytes and neutrophils. LRG1 has no known catalytic activity; it functions extracellularly as a context-dependent receptor-binding and signaling modulator. In endothelial cells exposed to TGF-beta, LRG1 binds endoglin and type I and type II TGF-beta receptor components, favors the pro-angiogenic ALK1-SMAD1/5 branch, and promotes endothelial proliferation and angiogenesis. In mouse fibroblasts it can instead enhance TGF-beta-SMAD2 signaling and profibrotic responses without requiring endoglin, illustrating that its signaling output depends on cellular receptor context. LRG1 glycosylation also gates a distinct interaction with the adhesion G protein-coupled receptor LPHN2: removal of the N325 glycan increases receptor binding and LPHN2-dependent angiogenic and neurotrophic activity in experimental systems. LRG1 additionally contributes to inflammatory-cell, re-epithelialization, vascular, and NETosis responses during wound repair, with opposing net effects in normal and diabetic mouse wounds. No alternative protein isoforms are reported.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005576 extracellular region
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference places active LRG1 in the extracellular region.
Reason: LRG1 is a signal-peptide-containing secreted glycoprotein, so this broad active compartment is correct and central to its receptor-modulating function.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0005114 type II transforming growth factor beta receptor binding
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference assigns binding to type II TGF-beta receptors.
Reason: Direct human experiments independently show LRG1 interaction with TGFBR2, making the conserved inference biologically sound and core.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0030511 positive regulation of transforming growth factor beta receptor signaling pathway
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference assigns positive regulation of TGF-beta receptor signaling.
Reason: Human LRG1 reorganizes the endothelial TGF-beta receptor complex toward pro-angiogenic SMAD1/5 signaling; this is a central mechanistic process.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0034713 type I transforming growth factor beta receptor binding
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference assigns binding to type I TGF-beta receptors.
Reason: Direct human experiments show LRG1 association with ALK1 and ALK5, independently supporting this conserved molecular function.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0035313 wound healing, spreading of epidermal cells
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Phylogenetic inference assigns a role in epidermal spreading during wound healing.
Reason: LRG1-dependent keratinocyte migration and re-epithelialization support the term, but it is a tissue-specific downstream outcome of extracellular signaling.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0045766 positive regulation of angiogenesis
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference assigns positive regulation of angiogenesis.
Reason: Angiogenesis is a well-supported major output of LRG1-mediated TGF-beta receptor-complex modulation in endothelial cells.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
PANTHER:PTN008456974
UniProtKB:P02750
GO:0005576 extracellular region
IEA
GO_REF:0000120
ACCEPT
Summary: Automated orthology and UniProt subcellular-location mappings place LRG1 extracellularly.
Reason: The mouse ortholog transfer and secreted-location mapping agree with the human signal peptide, plasma protein identity, and curated secretion statement.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q91XL1
ensembl:ENSMUSP00000038048
UniProtKB-SubCell:SL-0243
GO:0005515 protein binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
MODIFY
Summary: Interaction experiments detect LRG1 binding to the TGF-beta accessory receptor endoglin.
Reason: The interaction is well supported, but generic protein binding is uninformative; type III transforming growth factor beta receptor binding precisely captures interaction with endoglin.
Supporting Evidence:
PMID:23868260
In primary brain ECs LRG1 was present in immunoprecipitates of TΞ²RII, ALK1, ALK5 and the auxiliary receptor ENG (Fig 4a).
GO:0005515 protein binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
MODIFY
Summary: Interaction experiments detect LRG1 binding to the type I TGF-beta receptor ALK5/TGFBR1.
Reason: The interaction is well supported, but the partner supports the more informative type I transforming growth factor beta receptor binding term.
Supporting Evidence:
PMID:23868260
In primary brain ECs LRG1 was present in immunoprecipitates of TΞ²RII, ALK1, ALK5 and the auxiliary receptor ENG (Fig 4a).
GO:0005515 protein binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
MODIFY
Summary: Interaction experiments detect LRG1 association with the type I TGF-beta receptor ALK1/ACVRL1.
Reason: Generic protein binding loses the biologically informative receptor identity; type I transforming growth factor beta receptor binding is the appropriate specific term.
Supporting Evidence:
PMID:23868260
In primary brain ECs LRG1 was present in immunoprecipitates of TΞ²RII, ALK1, ALK5 and the auxiliary receptor ENG (Fig 4a).
GO:0005515 protein binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
MODIFY
Summary: Interaction experiments detect LRG1 binding to TGFBR2.
Reason: Generic protein binding should be replaced by the directly supported type II transforming growth factor beta receptor binding term.
Supporting Evidence:
PMID:23868260
Immunoprecipitation of the receptor ectodomain revealed coimmunoprecipitation of LRG1 with ALK5, TΞ²RII and ENG indicating a direct interaction of LRG1 with these individual receptors (Fig 4b).
GO:0005515 protein binding
IPI
PMID:28514442
Architecture of the human interactome defines protein commun...
MARK AS OVER ANNOTATED
Summary: A proteome-scale interaction screen reports LRG1 association with cytochrome c.
Reason: Protein binding is uninformative. Targeted surface-plasmon-resonance assays independently show human LRG binding to horse and snake cytochrome c, but not yeast cytochrome c; human CYCS was not tested, and extracellular binding or a physiological role has not been demonstrated in vivo. The high-throughput interaction is therefore retained without promoting it to a core LRG1 function.
Supporting Evidence:
PMID:20442399
Human LRG bound horse and snake Cyt c with dissociation constants of 1.58 x 10(-13) M and 1.65 x 10(-10) M, respectively, but did not bind yeast Cyt c
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
MARK AS OVER ANNOTATED
Summary: A cell-specific proteome-scale interaction map reports LRG1 association with cytochrome c.
Reason: Protein binding is uninformative. Targeted surface-plasmon-resonance assays independently show human LRG binding to horse and snake cytochrome c, but not yeast cytochrome c; human CYCS was not tested, and extracellular binding or a physiological role has not been demonstrated in vivo. The high-throughput interaction is therefore retained without promoting it to a core LRG1 function.
Supporting Evidence:
PMID:20442399
Human LRG bound horse and snake Cyt c with dissociation constants of 1.58 x 10(-13) M and 1.65 x 10(-10) M, respectively, but did not bind yeast Cyt c
GO:0005160 transforming growth factor beta receptor binding
IEA
GO_REF:0000107
ACCEPT
Summary: Ensembl orthology transfer assigns TGF-beta receptor binding from mouse Lrg1.
Reason: The orthology transfer is independently corroborated by direct human binding to TGFBR1, TGFBR2, ACVRL1, and endoglin.
Propagation Review
Root cause: NO FAILURE CORE
Sources checked:
UniProtKB:Q91XL1
ensembl:ENSMUSP00000038048
GO:0140149 non-collagenous component of interstitial matrix
TAS
PMID:36399478
MatrisomeDB 2.0: 2023 updates to the ECM-protein knowledge d...
MARK AS OVER ANNOTATED
Summary: MatrisomeDB curator inference places LRG1 among non-collagenous interstitial-matrix components.
Reason: MatrisomeDB supports LRG1 as a secreted matrisome-associated protein, but the available evidence does not establish it as a structural non-collagenous component of interstitial matrix. This cellular-component term therefore overstates the evidence.
Supporting Evidence:
PMID:36399478
We further divided this in-silico predicted matrisome into two main divisions: (i) structural components of the ECM or the β€˜core matrisome’ including ECM glycoproteins, collagens, and proteoglycans, which are highly insoluble in nature and (ii) matrisome-associated proteins including ECM-affiliated proteins, ECM regulators, and secreted factors known or expected to bind to structural components of the ECM.
GO:0010718 positive regulation of epithelial to mesenchymal transition
IDA
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: Recombinant and loss-of-function experiments support LRG1 promotion of epithelial-to-mesenchymal transition.
Reason: The effect is experimentally supported, but it is a context-dependent cellular response downstream of LRG1 signaling rather than the defining molecular function.
Supporting Evidence:
PMID:32887674
Our study discovered that rhLRG1 induces the expression of EMT markers, which is consistent with the promoting effect of LRG1 in EMT and colorectal cancer metastasis (16).
GO:0001938 positive regulation of endothelial cell proliferation
IMP
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
KEEP AS NON CORE
Summary: LRG1 and TGF-beta1 together stimulate proliferation of mouse brain endothelial cells from WT and Lrg1-null mice.
Reason: The cited Fig. 5b experiment directly supports a proliferative output in mouse brain endothelial cells. The occurs_in human umbilical-vein endothelial-cell extension is retained unchanged from the current QuickGO source annotation, but this mouse-cell experiment does not independently establish that cell-type context; proliferation remains a non-core downstream process.
Supporting Evidence:
PMID:23868260
The addition of LRG1 on its own had no effect, but TGFΞ²1 and LRG1 in combination increased proliferation significantly in both WT and Lrg1 null ECs (Fig 5b).
GO:0005114 type II transforming growth factor beta receptor binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
ACCEPT
Summary: A cell-free recombinant receptor-ectodomain assay shows LRG1 binding to TGFBR2.
Reason: Co-immunoprecipitation of separately expressed receptor ectodomains in HEK293 conditioned media directly supports this specific molecular function. The occurs_in human umbilical-vein endothelial-cell extension is retained unchanged from the current QuickGO source annotation, but is not established by this cell-free binding assay.
Supporting Evidence:
PMID:23868260
Immunoprecipitation of the receptor ectodomain revealed coimmunoprecipitation of LRG1 with ALK5, TΞ²RII and ENG indicating a direct interaction of LRG1 with these individual receptors (Fig 4b).
GO:0005515 protein binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
MODIFY
Summary: A cell-free recombinant receptor-ectodomain assay shows LRG1 binding to endoglin.
Reason: The cell-free receptor-ectodomain interaction is sound, but generic protein binding obscures the biologically relevant interaction with endoglin, a type III TGF-beta receptor. The occurs_in human umbilical-vein endothelial-cell extension is retained unchanged from the current QuickGO source annotation, but is not established by this binding assay.
Supporting Evidence:
PMID:23868260
LRG1 binds directly to the TGF-Ξ² accessory receptor endoglin, which, in the presence of TGF-Ξ²1, results in promotion of the pro-angiogenic Smad1/5/8 signalling pathway.
GO:0010838 positive regulation of keratinocyte proliferation
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: Lrg1 loss and gain perturb keratinocyte proliferation in wounded skin and cultured keratinocytes.
Reason: The proliferative effect is supported, but it is a wound-context downstream response rather than LRG1's defining receptor-binding activity.
Supporting Evidence:
PMID:32887674
Here, we showed reduced number of proliferating keratinocytes as indicated by Ki67 staining at the wound edge of Lrg1-deficient mice.
GO:0030511 positive regulation of transforming growth factor beta receptor signaling pathway
IDA
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
ACCEPT
Summary: LRG1 shifts endothelial TGF-beta signaling toward pro-angiogenic SMAD1/5 activity upstream of endothelial proliferation.
Reason: Receptor-complex binding, signaling assays, knockdown, and rescue experiments establish positive modulation of the pathway as a central LRG1 mechanism.
Supporting Evidence:
PMID:23868260
LRG1 antibody blockade inhibits this switch and attenuates angiogenesis. These studies reveal a new regulator of angiogenesis that mediates its effect by modulating TGF-Ξ² signalling.
GO:0030511 positive regulation of transforming growth factor beta receptor signaling pathway
IDA
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
ACCEPT
Summary: Wound-healing experiments support LRG1-dependent TGF-beta receptor signaling in skin.
Reason: Although assayed in a wound context, the result independently supports the same central receptor-signaling mechanism established in endothelial cells.
Supporting Evidence:
PMID:32887674
Furthermore, we demonstrated that LRG1-mediated NET formation is dependent on activation of the Akt pathway through TGFΞ² type I receptor ALK5, which is in agreement with LRG1-mediated TGFΞ² signaling in ECs (11), fibroblasts (34), glioma cells (46), and T-helper 17 cells (47).
GO:0034713 type I transforming growth factor beta receptor binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
ACCEPT
Summary: A cell-free recombinant receptor-ectodomain assay shows LRG1 binding to ALK5/TGFBR1.
Reason: Co-immunoprecipitation of separately expressed receptor ectodomains in HEK293 conditioned media supports this specific and central receptor-binding activity. The occurs_in human umbilical-vein endothelial-cell extension is retained unchanged from the current QuickGO source annotation, but is not established by this cell-free binding assay.
Supporting Evidence:
PMID:23868260
Immunoprecipitation of the receptor ectodomain revealed coimmunoprecipitation of LRG1 with ALK5, TΞ²RII and ENG indicating a direct interaction of LRG1 with these individual receptors (Fig 4b).
GO:0034713 type I transforming growth factor beta receptor binding
IPI
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
ACCEPT
Summary: A cell-free recombinant receptor-ectodomain assay shows endoglin-dependent LRG1 association with ALK1/ACVRL1.
Reason: The assay supports an endoglin-dependent association with ALK1 rather than unconditional binary binding. The occurs_in human umbilical-vein endothelial-cell extension is retained unchanged from the current QuickGO source annotation, but is not established by this cell-free ectodomain assay.
Supporting Evidence:
PMID:23868260
These studies revealed that LRG1 only associated with ALK1 in the presence of ENG, to which it bound, and this was enhanced by the addition of TGFΞ²1 (Fig 4d).
GO:0035313 wound healing, spreading of epidermal cells
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: Lrg1 deficiency delays epidermal spreading and re-epithelialization during cutaneous wound repair.
Reason: The wound phenotype is directly supported but is a tissue-specific downstream role rather than the core receptor-modulating molecular function.
Supporting Evidence:
PMID:32887674
Lrg1 deficiency leads to a significant delay in normal wound healing as a consequence of impaired inflammation, reepithelialization, and angiogenesis.
GO:0045766 positive regulation of angiogenesis
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
ACCEPT
Summary: LRG1 promotes vascular growth in the wound bed and in dermal endothelial-cell assays.
Reason: This tissue-qualified study independently supports angiogenesis as a major biological output of LRG1 signaling.
Supporting Evidence:
PMID:32887674
We further showed that LRG1 promotes angiogenesis by mediating HDMEC proliferation, migration, and the ability to form tube-like structures.
GO:0045766 positive regulation of angiogenesis
IMP
PMID:23868260
LRG1 promotes angiogenesis by modulating endothelial TGF-Ξ² s...
ACCEPT
Summary: Genetic loss, antibody blockade, and endothelial assays show that LRG1 promotes angiogenesis.
Reason: The study establishes angiogenesis as a major LRG1 output through endothelial TGF-beta receptor-complex modulation.
Supporting Evidence:
PMID:23868260
We show that in the presence of transforming growth factor-Ξ²1 (TGF-Ξ²1), LRG1 is mitogenic to endothelial cells and promotes angiogenesis.
GO:0051546 keratinocyte migration
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: LRG1 gain and loss alter keratinocyte migration during re-epithelialization.
Reason: Keratinocyte migration is supported as a wound-context response downstream of extracellular LRG1 signaling, but is not a core molecular function.
Supporting Evidence:
PMID:32887674
Consistent with the previous report (21), we showed that LRG1 overexpression and knockdown affect keratinocyte migration.
GO:0060054 positive regulation of epithelial cell proliferation involved in wound healing
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: Lrg1 deficiency reduces proliferating keratinocytes and delays wound re-epithelialization.
Reason: This is a well-supported but tissue-specific downstream wound-healing process.
Supporting Evidence:
PMID:32887674
Here, we showed reduced number of proliferating keratinocytes as indicated by Ki67 staining at the wound edge of Lrg1-deficient mice.
GO:0061756 leukocyte adhesion to vascular endothelial cell
IMP
PMID:32887674
A Multifunctional Role of Leucine-Rich Ξ±-2-Glycoprotein 1 in...
KEEP AS NON CORE
Summary: LRG1 promotes neutrophil adhesion and inflammatory-cell recruitment during wound healing.
Reason: The IMP evidence supports the leukocyte-adhesion phenotype, but it is a specialized inflammatory outcome rather than the core receptor-binding mechanism.
Supporting Evidence:
PMID:32887674
Our study showed that LRG1 promotes neutrophil adhesion, likely by inducing the expression of neutrophil adhesion molecule, L-selectin.
GO:0005576 extracellular region
HDA
PMID:27068509
Extracellular matrix remodelling in response to venous hyper...
ACCEPT
Summary: ECM-enriched proteomics detects LRG1 in human saphenous-vein tissue.
Reason: The curated tissue-qualified proteomic localization agrees with LRG1 secretion; extracellular region is broad but correct.
Supporting Evidence:
PMID:27068509
Varicose saphenous veins removed during phlebectomy and normal saphenous veins obtained during coronary artery bypass surgery were collected for proteomics analysis. Extracellular matrix proteins were enriched from venous tissues.
GO:0005576 extracellular region
TAS
Reactome:R-HSA-6798745
ACCEPT
Summary: Reactome places LRG1 extracellularly after exocytosis from tertiary neutrophil granules.
Reason: The pathway-specific source is consistent with LRG1 being a secreted plasma glycoprotein; the broad extracellular destination is correct.
GO:0005576 extracellular region
TAS
Reactome:R-HSA-6798749
ACCEPT
Summary: Reactome places LRG1 extracellularly after exocytosis from specific neutrophil granules.
Reason: Granule exocytosis supplies a biologically plausible route to the independently established extracellular compartment.
GO:0005576 extracellular region
TAS
Reactome:R-HSA-6800434
ACCEPT
Summary: Reactome places LRG1 extracellularly after exocytosis from ficolin-1-rich neutrophil granules.
Reason: The pathway-specific route is consistent with secretion and supports the broad extracellular localization.
GO:0035580 specific granule lumen
TAS
Reactome:R-HSA-6798749
KEEP AS NON CORE
Summary: Reactome assigns LRG1 to the lumen of specific neutrophil granules before exocytosis.
Reason: This is a plausible specialized storage compartment linked to neutrophil expression, but it is contextual rather than the general site of LRG1 action.
GO:1904724 tertiary granule lumen
TAS
Reactome:R-HSA-6798745
KEEP AS NON CORE
Summary: Reactome assigns LRG1 to the lumen of tertiary neutrophil granules before exocytosis.
Reason: The granule localization is compatible with neutrophil production and regulated release, but is a cell-specific storage site.
GO:1904813 ficolin-1-rich granule lumen
TAS
Reactome:R-HSA-6800434
KEEP AS NON CORE
Summary: Reactome assigns LRG1 to the lumen of ficolin-1-rich neutrophil granules before exocytosis.
Reason: This is a specialized neutrophil storage compartment and not the general extracellular site where released LRG1 acts.
GO:0070062 extracellular exosome
HDA
PMID:23533145
In-depth proteomic analyses of exosomes isolated from expres...
KEEP AS NON CORE
Summary: Shotgun proteomics detects LRG1 in extracellular exosome preparations from expressed prostatic secretions.
Reason: The high-throughput detection is retained with curator deference, but exosome association is sample-specific and not necessary to define LRG1's secreted signaling role.
Supporting Evidence:
PMID:23533145
Using expressed prostatic secretions in urine (EPS-urine), exosome preparations were characterized by a shotgun proteomics procedure.
GO:0005576 extracellular region
HDA
PMID:16502470
Human colostrum: identification of minor proteins in the aqu...
ACCEPT
Summary: Proteomics detects LRG1 in the aqueous phase of human colostrum.
Reason: Detection in an extracellular body fluid is consistent with LRG1 secretion; the colostrum extension appropriately preserves the sampling context.
Supporting Evidence:
PMID:16502470
We have investigated the low abundance proteins in the aqueous phase of human colostrum, after depletion of the major proteins secretory IgA, lactoferrin, alpha-lactalbumin and HSA by immunoabsorption, using 2-D LC and gel-based proteomic methods.
GO:0005576 extracellular region
IDA
PMID:20363744
Prolyl 3-hydroxylase 1 null mice display abnormalities in fi...
ACCEPT
Summary: A curator-assigned experimental annotation places LRG1 in the extracellular region.
Reason: The cached record is abstract-only and does not expose the LRG1 assay, so the experimental curator call is retained; extracellular localization is independently clear from LRG1's signal peptide, secretion record, and plasma occurrence.
GO:0005576 extracellular region
NAS
PMID:12223515
Molecular characterization and expression analysis of leucin...
ACCEPT
Summary: A sequence and expression study supports LRG1 as an extracellular glycoprotein associated with granulocytic differentiation.
Reason: Although the cached abstract emphasizes expression rather than localization, the curator statement agrees with the signal peptide and the established identity of mature LRG1 as a circulating secreted protein.
GO:0005576 extracellular region
NAS
PMID:14718574
The human plasma proteome: a nonredundant list developed by ...
ACCEPT
Summary: Human plasma-proteome evidence places LRG1 in an extracellular body fluid.
Reason: Plasma detection directly supports the broad extracellular-region annotation and is concordant with the reviewed UniProt secreted location.
Supporting Evidence:
PMID:14718574
The resulting nonredundant list confirms the presence of a number of interesting candidate marker proteins in plasma and serum.
GO:0016020 membrane
NAS
PMID:3856868
Periodicity of leucine and tandem repetition of a 24-amino a...
REMOVE
Summary: An early sequence analysis proposed that LRG1 is membrane-derived or membrane-associated.
Reason: The NAS annotation reflects a historical sequence-based hypothesis rather than observed membrane localization. It is superseded by the experimentally supported cleaved signal peptide (residues 1-35), soluble secreted mature plasma protein (residues 36-347), and absence of a transmembrane segment, so the membrane annotation should be removed.
Supporting Evidence:
PMID:3856868
These structural characteristics and the homology to mitochondrial proteins and apolipoproteins suggest that LRG is a membrane-derived or membrane-associated protein containing a series of domains capable of bipolar surface orientation.
GO:0061761 alpha-latrotoxin receptor binding
IPI
PMID:37121976
Crystal structure of LRG1 and the functional significance of...
NEW
Summary: Biochemical and cell-based experiments show glycan-state-dependent binding of human LRG1 to the alpha-latrotoxin receptor family member LPHN2.
Reason: GO:0061761 specifically captures binding to an alpha-latrotoxin receptor and is more informative than generic G protein-coupled receptor or protein binding. High-affinity LPHN2 binding is strongest after deglycosylation, particularly removal of the N325 glycan; downstream angiogenic and neurotrophic assays were performed in mouse explants, so this row asserts the directly tested binding activity without generalizing those physiological outputs to humans.
Supporting Evidence:
PMID:37121976
In addition, our biochemical and cell-biological analyses found that the deglycosylation of LRG1, particularly the removal of glycans on N325, is critical for the high-affinity binding of LRG1 to LPHN2 and thus promotes LRG1/LPHN2-mediated angiogenic and neurotrophic processes in mouse tissue explants, even under normal glucose conditions.
PMID:37121976
Recombinant human LRG1 protein and human LPHN2 variant proteins (Lec, Olf, Lec/Olf domain, or ecto-full domain) were expressed and purified as described previously20.

Core Functions

Extracellular LRG1 binds components of the TGF-beta receptor system and modulates pathway output according to ligand, receptor, and cell context. Direct one-to-one ectodomain binding is established for TGFBR1/ALK5, TGFBR2, and the type III receptor endoglin; association with ACVRL1/ALK1 is conditional on endoglin and is enhanced by TGF-beta1. In endothelial cells, LRG1 alone is not an agonist, but in the presence of TGF-beta1 it favors the pro-angiogenic ALK1-SMAD1/5 branch and promotes angiogenesis. Enhancement of TGF-beta-SMAD2 signaling in mouse fibroblasts is endoglin-independent, showing that downstream output is cell-context-dependent rather than a universal ALK1 mechanism.

Supporting Evidence:
  • PMID:23868260
    Immunoprecipitation of the receptor ectodomain revealed coimmunoprecipitation of LRG1 with ALK5, TΞ²RII and ENG indicating a direct interaction of LRG1 with these individual receptors (Fig 4b).
  • PMID:23868260
    These studies revealed that LRG1 only associated with ALK1 in the presence of ENG, to which it bound, and this was enhanced by the addition of TGFΞ²1 (Fig 4d).
  • PMID:23868260
    The addition of LRG1 alone did not activate Smad2/3 or Smad1/5, but in combination with TGFβ1 there was a dramatic induction of Smad1/5 phosphorylation showing that LRG1 requires the presence of TGFβ1 to stimulate the pro-angiogenic TßRII-ALK1-Smad1/5/8 pathway (Fig 5a).
  • PMID:29279415
    Although endoglin, an accessory TGF-Ξ² receptor, is essential for LRG to promote TGF-Ξ² signaling in endothelial cells during angiogenesis, we found that endoglin did not contribute to the ability of LRG to enhance Smad2 phosphorylation in fibroblasts.

Human LRG1 binds the adhesion G protein-coupled receptor LPHN2 through a glycan-state-dependent activity. Removal of LRG1 glycans, especially the N325 glycan, increases high-affinity receptor binding and LPHN2-dependent signaling. Angiogenic and neurotrophic outputs have been demonstrated in mouse tissue explants and diabetic mouse models, whereas the abundance and physiological generation of the relevant deglycosylated human LRG1 species remain unknown; this is therefore a conditional receptor-binding activity, not a constitutive property assigned to all circulating LRG1.

Supporting Evidence:
  • PMID:37121976
    In addition, our biochemical and cell-biological analyses found that the deglycosylation of LRG1, particularly the removal of glycans on N325, is critical for the high-affinity binding of LRG1 to LPHN2 and thus promotes LRG1/LPHN2-mediated angiogenic and neurotrophic processes in mouse tissue explants, even under normal glucose conditions.
  • PMID:37121976
    Collectively, these data indicate a novel role of LRG1 glycans as molecular switches that can tune the range of LRG1's cellular functions, particularly the LRG1/LPHN2 signaling axis.

References

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Suggested Questions for Experts

Q: Which endogenous human LRG1 glycoforms bind and activate LPHN2, where are they generated, and under which physiological or disease conditions?

Q: What receptor stoichiometry and cell-state variables determine whether LRG1 enhances ALK1-SMAD1/5 or ALK5-SMAD2 signaling?

Q: Does native human LRG1 bind human extracellular cytochrome c and neutralize a defined cytochrome-c-dependent activity in vivo?

Q: Which cellular sources of LRG1 account for its opposing net effects in normal and diabetic wound healing?

Suggested Experiments

Experiment: Quantify human LRG1 binding to purified TGFBR1, TGFBR2, endoglin, ACVRL1, and TGF-beta in defined combinations by orthogonal biophysical methods; determine complex stoichiometry and structure, then test matched receptor perturbations and SMAD outputs in primary human endothelial cells and fibroblasts.

Hypothesis: Endoglin and TGF-beta jointly stabilize an LRG1-containing receptor assembly that favors endothelial ALK1-SMAD1/5 signaling, whereas a distinct assembly supports fibroblast ALK5-SMAD2 signaling.

Type: Structural biochemistry and pathway reconstitution

Experiment: Isolate LRG1 from human plasma, neutrophils, and relevant tissue fluids; map site-specific glycoforms by quantitative glycoproteomics, fractionate them, and compare LPHN1-3 binding and LPHN2-dependent signaling with recombinant native, deglycosylated, and N325-mutant controls.

Hypothesis: A discrete endogenous human LRG1 glycoform lacking or remodeling the N325 glycan is the physiologically active LPHN2 ligand.

Type: Glycoproteomics, receptor binding, and signaling assays

Experiment: Measure binding kinetics between native human LRG1 glycoforms and human cytochrome c, map the interaction interface, and test LRG1 dose-response and loss-of-function effects in human-cell assays exposed to extracellular cytochrome c before pursuing an in-vivo model.

Hypothesis: Human LRG1 recognizes human extracellular cytochrome c and changes its extracellular bioactivity or clearance.

Type: Species-matched biophysics and functional perturbation

Experiment: Combine lineage-restricted Lrg1 deletion or rescue with normal and diabetic mouse wound models, time-resolved inflammation, re-epithelialization, angiogenesis, and NETosis measurements, and validate conserved mechanisms in organotypic human skin cultures with source-specific LRG1 perturbation.

Hypothesis: Hepatic, neutrophil-derived, and local tissue LRG1 make separable contributions to normal repair and diabetes-associated NETosis and delayed wound closure.

Type: Cell-source-resolved wound model and human tissue validation

Knowledge Gaps

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

Gap: The structure, stoichiometry, and cell-context determinants of the LRG1-TGF-beta receptor assembly are unresolved, including how endoglin and ligand exposure govern conditional ALK1 association and selection between ALK1-SMAD1/5 and ALK5-SMAD2 outputs.

OPEN BIOLOGY RESIDUAL_SUBGAP

What is known: LRG1 directly binds isolated ALK5, TGFBR2, and endoglin ectodomains, while ALK1 association requires endoglin and is enhanced by TGF-beta1; endothelial and fibroblast assays establish distinct downstream outputs.

Significance: Defining the receptor assembly is necessary to predict when LRG1 will drive angiogenic rather than profibrotic TGF-beta signaling and to target the mechanism selectively.

What would resolve it: Determine structures and binding stoichiometries of human LRG1 with defined TGFBR1, TGFBR2, endoglin, ACVRL1, and TGF-beta combinations, then reconstitute branch-specific signaling in primary human endothelial cells and fibroblasts.

Provenance (the field's own admissions):

Gap: The physiological source, tissue distribution, and abundance of LPHN2-activating deglycosylated LRG1 in humans are unknown, as is the precise structural basis for LPHN2 selectivity over LPHN1 and LPHN3.

OPEN BIOLOGY RESIDUAL_SUBGAP

What is known: Removing LRG1 glycans, especially at N325, increases LPHN2 binding, and the resulting angiogenic and neurotrophic outputs have been shown in mouse tissue explants and diabetic mouse models.

Significance: Resolving this gap will establish whether glycan-regulated LPHN2 signaling is a physiological human mechanism or primarily an experimentally induced state.

What would resolve it: Use site-resolved glycoproteomics and LPHN2-binding assays on endogenous human plasma, neutrophil, and tissue LRG1, coupled to structural analysis of a human LRG1-LPHN2 complex and receptor-paralog selectivity tests.

Provenance (the field's own admissions):

Gap: It is unknown whether human LRG1 binds human extracellular cytochrome c with physiologically relevant affinity and neutralizes cytochrome-c activity in vivo.

OPEN BIOLOGY RESIDUAL_SUBGAP

What is known: Human LRG1 bound horse and snake cytochrome c, but not yeast cytochrome c, in surface-plasmon-resonance assays; independent proteome-scale screens report a human LRG1-CYCS interaction without locally accessible pair-specific assay detail.

Significance: This distinguishes a physiological extracellular damage-response function from a plausible but incompletely tested interaction.

What would resolve it: Measure binding of native human LRG1 glycoforms to human cytochrome c, map the interface, and test whether LRG1 alters extracellular cytochrome-c activity or clearance in human-cell and in-vivo models.

Provenance (the field's own admissions):

πŸ“š Additional Documentation

Notes

(LRG1-notes.md)

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