Affinage · Toxoplasma gondii

GRA3

dense granule protein GRA3 · UniProt A0A125YY45

Annotated
2026-09-05
41 papers in source corpus 19 papers cited in narrative 19 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

GRA3 is a Toxoplasma gondii dense granule protein that is secreted into the parasitophorous vacuole upon host-cell invasion and functions from the parasitophorous vacuole membrane (PVM) as an interface for co-opting host organelles and modulating host immunity (PMID:8195173, PMID:7739670, PMID:19127325). It is a 24 kDa protein carrying an N-terminal secretory signal sequence, a transmembrane domain, and a C-terminal dilysine (KKXX) ER-retrieval motif; the signal peptide alone is necessary and sufficient to target reporter protein to dense granules and the PV (PMID:16149193). Stored as a soluble protein in dense granules, GRA3 is released by constitutive, calcium-independent exocytosis mediated by the NSF/SNAP/SNARE/Rab machinery and inserts directly into the PVM in an oligomeric form through hydrophobic interactions, self-cross-linking without incorporating other parasite or host proteins (PMID:8195173, PMID:9891012). From the PVM, GRA3 engages host membranes and signaling: it binds the host ER protein CAMLG, recruiting ER to the PVM (PMID:19127325); it binds host Golgi membranes, oligomerizes, and drives Golgi tubulation and entry of Golgi material into the PV (PMID:30814066); and it interacts with host STING to promote STING oligomerization and IRF3 nuclear translocation, engaging the cGAS-STING and STAT6 axes to enhance parasite proliferation via ISG56 and STAT6 (PMID:41712610). GRA3 suppresses MHC-I antigen presentation of PV-localized antigens to CD8+ T cells independently of IFN-γ/IRG effectors (PMID:31555296), and contributes to intracellular replication, acute virulence, and chronic cyst persistence in vivo (PMID:17418907, PMID:31266861, PMID:40730529, PMID:37868957), where it is also a component of the cyst wall (PMID:31040239). Expressed in tachyzoites but absent from sporozoites (PMID:8720177), GRA3 (Wh6 allele) can additionally trigger PERK-dependent ER-stress apoptosis in neuronal cells when overexpressed (PMID:35918751).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1993 High

    Established when and where GRA3 is delivered to the host interface, showing dense granule discharge occurs at the apical pole within minutes of invasion and deposits GRA3 at the nascent PVM.

    Evidence IFA, immunoelectron microscopy, and freeze-fracture at defined post-invasion time points in T. gondii

    PMID:8415546

    Open questions at the time
    • Did not define the biochemical mechanism of membrane insertion
    • Did not identify host partners at the PVM
  2. 1994 High

    Answered how a soluble dense granule protein becomes membrane-associated, showing GRA3 inserts directly into the vacuole membrane as a self-associating oligomer via hydrophobic interactions rather than through a stable partner complex.

    Evidence Sucrose flotation, Triton X-114 partitioning, carbonate extraction, and chemical cross-linking in T. gondii

    PMID:8195173

    Open questions at the time
    • Sequence used was later shown to be a chimera
    • Topology and orientation of the inserted protein not resolved
  3. 1994 High

    Defined the molecular architecture by cloning GRA3 cDNA and demonstrating an N-terminal signal sequence sufficient for dense granule/PV targeting, though the original sequence carried an error.

    Evidence cDNA cloning, in vitro translation with dog pancreas microsomes, IFA, Southern/Northern blot in T. gondii

    PMID:7739670

    Open questions at the time
    • Reported sequence and MW later corrected
    • Function beyond localization not addressed
  4. 1995 Medium

    Established stage-specific expression, showing GRA3 is absent from sporozoites and appears at tachyzoite conversion, tying its function to the replicative intracellular stage.

    Evidence IFA and Western blot time course in sporozoite-infected cells in T. gondii

    PMID:8720177

    Open questions at the time
    • Regulatory basis of stage-specific expression not addressed
    • Sporozoite-stage functional consequence not tested
  5. 1999 High

    Defined the secretion pathway, showing GRA3 release is constitutive, calcium-independent, and dependent on NSF/SNAP/SNARE/Rab machinery.

    Evidence Endogenous secretion assay in SLO-permeabilized parasites with recombinant NSF/α-SNAP and inhibitors (NEM, Rab-GDI) in T. gondii

    PMID:9891012

    Open questions at the time
    • Specific SNARE/Rab identities not defined
    • Trigger coupling secretion to invasion not resolved
  6. 2005 High

    Corrected the primary structure, establishing authentic GRA3 as a 24 kDa protein with signal sequence, transmembrane domain, and a C-terminal dilysine ER-retrieval motif consistent with PVM insertion.

    Evidence Sequence analysis, recombinant antibody, GFP targeting, and ESA immunoblot in T. gondii

    PMID:16149193

    Open questions at the time
    • Functional role of the KKXX motif not experimentally tested
    • Host-facing versus lumenal domain topology unresolved
  7. 2007 High

    Delineated GRA3's oligomeric behavior relative to other GRA proteins, showing it co-purifies with soluble GRA2/GRA6/GRA7 complexes but dissociates from GRA2 after membrane insertion.

    Evidence HA-FLAG affinity purification from complemented KO parasites with MS in T. gondii

    PMID:17959262

    Open questions at the time
    • Whether GRA3 forms independent oligomers versus mixed complexes in the membrane unresolved
    • Stoichiometry not defined
  8. 2007 High

    Provided the first genetic test of function, showing GRA3 is dispensable for in vitro growth and bradyzoite formation but contributes to acute virulence in vivo, restorable by complementation.

    Evidence Homologous-recombination KO with complementation and mouse survival in a type II strain of T. gondii

    PMID:17418907

    Open questions at the time
    • Molecular mechanism of virulence contribution not identified
    • Host pathways affected not defined
  9. 2008 Medium

    Identified the first host-cell partner, showing PVM-inserted GRA3 binds the host ER protein CAMLG, providing a molecular basis for ER recruitment to the PVM.

    Evidence Yeast two-hybrid, GST pull-down, co-IP, and co-localization in infected cells for T. gondii

    PMID:19127325

    Open questions at the time
    • Functional consequence of ER recruitment for the parasite not tested
    • Interaction not validated in a knockout background
  10. 2008 Medium

    Placed GRA3 within cytosolic strand-like extensions of the PVM alongside GRA1, GRA7, and rhoptry proteins, extending its localization beyond the vacuole membrane proper.

    Evidence Multi-protein co-labeling IFA at defined time points in T. gondii

    PMID:18809661

    Open questions at the time
    • Functional role of GRA3 in strand structures not defined
    • Fixed-cell co-localization only
  11. 2016 Medium

    Distinguished GRA3 from IRG-modulating effectors by showing it is dispensable for the antitumor immunity induced by uracil-auxotroph vaccines, unlike GRA2/GRA12/GRA24 and several ROPs.

    Evidence Targeted deletion in Δku80 vaccine strain with ovarian tumor model and immune readouts in type I T. gondii

    PMID:27447180

    Open questions at the time
    • Does not exclude a role in other immune contexts
    • Negative result in a specific vaccine setting only
  12. 2019 Medium

    Revealed GRA3's role in host Golgi hijacking, showing it binds Golgi membranes, oligomerizes, and promotes Golgi tubulation and engulfment into the PV while dysregulating anterograde transport.

    Evidence Gene disruption with live host-Golgi imaging and direct Golgi membrane binding assays in T. gondii

    PMID:30814066

    Open questions at the time
    • Direct GRA3 receptor on Golgi not identified
    • Parasite benefit of Golgi engulfment not quantified
  13. 2019 High

    Established a chronic-stage requirement, showing GRA3 deletion in the Pru type II strain severely impairs in vivo cyst development while sparing growth and in vitro differentiation.

    Evidence Targeted deletion with in vivo cyst burden and in vitro bradyzoite assays in Pru type II T. gondii

    PMID:31266861

    Open questions at the time
    • Molecular basis of the cyst defect not defined
    • Whether the defect is intrinsic or immune-driven unresolved
  14. 2019 Medium

    Identified GRA3 as a cyst-wall component, linking its PVM/interface role to the chronic-stage structure.

    Evidence Percoll-enriched cyst wall fragments with CST1 IP and LC-MS/MS in T. gondii

    PMID:31040239

    Open questions at the time
    • Functional contribution to cyst wall integrity not tested
    • Association versus structural role not distinguished
  15. 2019 Medium

    Defined an immune-evasion function, showing GRA3 suppresses MHC-I presentation of PV antigens to CD8+ T cells independently of IFN-γ/IRG effectors.

    Evidence GRA3 deletion with MHC-I antigen presentation assays in macrophages and dendritic cells ±IFN-γ in type II T. gondii

    PMID:31555296

    Open questions at the time
    • Molecular mechanism of antigen presentation suppression not identified
    • Relevant host trafficking step not defined
  16. 2022 Medium

    Uncovered a host-cell-death consequence, showing overexpressed GRA3 (Wh6 allele) triggers PERK-dependent ER-stress apoptosis in neuronal cells.

    Evidence Overexpression in N2a cells with ER-stress/apoptosis marker immunoblotting and PERK/caspase-12 inhibitors in Chinese 1 Wh6 T. gondii

    PMID:35918751

    Open questions at the time
    • Demonstrated by overexpression rather than in infection
    • Allele-specificity versus general GRA3 effect not resolved
  17. 2023 Medium

    Linked GRA3 to CD8+ T-cell resistance during chronic infection, showing GRA3-deficient cysts are more efficiently eliminated by perforin-dependent CD8+ T cells and that persisting cysts are enriched for GRA3 mRNA.

    Evidence GRA3-deficient strain with in vivo cyst burden under CD8+ immunity and qRT-PCR correlation in BALB/c mice with T. gondii

    PMID:37868957

    Open questions at the time
    • Causal versus correlative role of GRA3 mRNA in persistence not resolved
    • Link to MHC-I suppression mechanism not directly tested
  18. 2025 Medium

    Confirmed GRA3's contribution to replication and virulence with CRISPR deletion, showing ME49Δgra3 has reduced plaque formation, smaller PVs, and increased host mortality.

    Evidence CRISPR/Cas9 disruption with Western validation, plaque and PV-size assays, and mouse survival in ME49 T. gondii

    PMID:40730529

    Open questions at the time
    • Increased mortality mechanism not dissected
    • Molecular pathway linking GRA3 to replication not defined here
  19. 2026 Medium

    Identified a signaling mechanism for GRA3-driven proliferation, showing GRA3 binds host STING to promote STING oligomerization, IRF3 nuclear translocation, and STAT6 phosphorylation, with proliferation mediated by ISG56 and STAT6 rather than IFN-β.

    Evidence Co-IP, STING oligomerization and nuclear fractionation assays, KO/overexpression, and ISG56/STAT6 KO/inhibitor proliferation assays in ME49 T. gondii

    PMID:41712610

    Open questions at the time
    • Direct binding interface with STING not mapped
    • How PVM-restricted GRA3 accesses host STING not resolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • How GRA3's single membrane insertion domain simultaneously coordinates ER (CAMLG), Golgi, and STING engagement, and whether these interactions are physically or temporally partitioned, remains unresolved.
  • No structural model of GRA3 at the PVM
  • Topology dictating which domains face the host cytosol undefined
  • Mechanistic link between host-organelle capture, immune evasion, and cyst persistence not integrated

Mechanism profile

Synthesis pass · controlled-vocabulary classification
Partners
CAMLG (HOST ER TYPE II TRANSMEMBRANE PROTEIN)GRA2, GRA6, GRA7 (SOLUBLE COMPLEX, PRE-INSERTION)HOST GOLGI MEMBRANESHOST STINGNSF / Α-SNAP / SNARE / RAB (SECRETION MACHINERY)
Complex memberships
Homo-oligomeric GRA3 assembly in the PVM (self-cross-linking)Soluble GRA2/GRA3/GRA6/GRA7 co-purifying complex prior to membrane insertion

Evidence

Reading pass · 19 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1994 GRA3 is stored as a soluble protein in dense granules and, following secretion into the parasitophorous vacuole, inserts directly into the vacuole membrane in an oligomeric form via hydrophobic interactions; it cannot be removed by sodium carbonate extraction, is not labeled by [1-14C]acetate, and cross-links to itself (forming a ladder on SDS-PAGE) but no other parasite or host proteins co-precipitate with cross-linked GRA3. Sucrose flotation gradients, Triton X-114 phase partitioning, sodium carbonate extraction, chemical cross-linking, SDS-PAGE, [1-14C]acetate labeling The Journal of biological chemistry High 8195173
1994 GRA3 cDNA encodes a 30 kDa protein with an N-terminal signal sequence (sufficient to target GFP to dense granules and PV) and four short hydrophobic regions insufficient to span the membrane; Southern blot indicates a single-copy gene; GRA3 localizes to dense granules, the PVM, and PVM extensions protruding into the host cytoplasm. cDNA cloning, Northern blot, Southern blot, in vitro translation with dog pancreas microsomes, immunofluorescence, immunoblot with affinity-purified antibodies Molecular and biochemical parasitology High 7739670
1993 Dense granule exocytosis (releasing GRA2 and GRA3) occurs in the apical area of the tachyzoite within minutes of invasion; dense granule contents appear first as dense material between parasite and vacuole membranes before the vacuolar network or PVM are immunolabeled; GRA3 antibodies label the PVM and strands extending from PVs into the host-cell cytoplasm. Immunofluorescence assay (IFA), immunoelectron microscopy, serial sectioning, freeze-fracture studies at defined post-invasion time points Parasitology research High 8415546
1999 GRA3 release from dense granules is constitutive and calcium-independent; the NSF/SNAP/SNARE/Rab machinery mediates dense granule exocytosis, as shown by inhibition with N-ethylmaleimide and Rab-GDI, and augmentation with recombinant NSF and α-SNAP in streptolysin O-permeabilized parasites. Endogenous GRA3 secretion assay in intact and SLO-permeabilized parasites, recombinant protein addition, pharmacological inhibitors (NEM, Rab-GDI), GTPγS and cAMP treatments The Journal of biological chemistry High 9891012
2005 The originally published GRA3 sequence is an artificial chimera of two proteins. The authentic GRA3 (24 kDa, recognized by mAb 2H11) contains an N-terminal secretory signal sequence and a transmembrane domain consistent with PVM insertion, plus a C-terminal dilysine (KKXX) ER retrieval motif. The signal peptide alone is necessary and sufficient to target GFP to dense granules and the PV. A Neospora caninum homologue was identified. Sequence analysis, recombinant antibody production, immunoblot of ESA, GFP fusion/targeting assay, bioinformatics identification of KKXX motif Parasitology High 16149193
2007 Within the parasite and the PV soluble fraction, GRA2 and GRA5 form high-molecular-weight oligomeric complexes with multiple GRA proteins; affinity purification of HA-FLAG-tagged GRA2 from ΔeGRA2 complemented parasites co-purified GRA3, GRA6, and GRA7 as the major partners. After membrane insertion, GRA2 associates with GRA5 and GRA7 but not GRA3. Affinity chromatography (HA-FLAG tag), conventional chromatography, complementation of KO parasites with tagged proteins, co-purification/MS Molecular and biochemical parasitology High 17959262
2007 Targeted deletion of GRA3 in a type II strain shows no effect on cell culture growth rate or bradyzoite formation, and does not alter localization of other dense granule proteins or association with host cell organelles; however, at ~4× the LD50, ΔeGRA3 mice survived acute infection, and complementation restored and even enhanced virulence, demonstrating GRA3 contributes to acute virulence in vivo. Homologous recombination gene knockout, plaque assay, bradyzoite differentiation assay, IFA for GRA protein localization, mouse survival experiments, genetic complementation Molecular and biochemical parasitology High 17418907
2008 PVM-inserted GRA3 physically interacts with CAMLG (calcium modulating ligand), a type II transmembrane protein of the host cell ER, as established by yeast two-hybrid screening of a HeLa cDNA library, GST pull-down, and co-immunoprecipitation; GRA3 and CAMLG co-localize at the PVM in infected cells, suggesting GRA3 mediates ER recruitment to the PVM. Yeast two-hybrid (GRA3 ORF vs HeLa cDNA library), GST pull-down assay, co-immunoprecipitation, co-transfection/co-localization by fluorescence microscopy, IFA in infected cells The Korean journal of parasitology Medium 19127325
2008 GRA7 strands in the host cytosol (formed within 10 min of invasion) also contain GRA1 and GRA3, indicating GRA3 is present in cytosolic strand-like structures alongside rhoptry proteins ROP2 and ROP4 in infected host cells. IFA with co-labeling of GRA7, GRA1, GRA3, ROP2, ROP4 at defined post-invasion time points Infection and immunity Medium 18809661
1995 Sporozoites of T. gondii lack GRA3 protein (not detected by IFA or Western blot in sporozoite-infected cells 2–15 h post-inoculation); GRA3 expression first appears 15 h post-inoculation and is present in all intracellular parasites at 24 h, coinciding with expression of the tachyzoite marker SAG1 and onset of replication. Indirect immunofluorescence, Western blot of sporozoite extracts, time-course monitoring of GRA3 and SAG1 expression Molecular and biochemical parasitology Medium 8720177
2019 Deletion of PVM-associated GRA3 in the type II Prugniaud strain induced severe defects in chronic cyst development in vivo without affecting parasite growth rate or in vitro cyst differentiation; acute virulence of PruΔgra3 was reduced but not abolished. Targeted gene deletion (Δgra3) in type II Pru strain, in vivo mouse cyst burden assay, in vitro bradyzoite differentiation, plaque assay mBio High 31266861
2019 TgGRA3 interacts with host Golgi membranes, oligomerizes, and promotes the formation of tubules and entry of host Golgi material into the PV; disruption of the TgGRA3 gene delays engulfment of host Golgi; GRA3 dysregulates anterograde transport in the host cell. Gene disruption (ΔgGRA3), live imaging/fluorescence microscopy of host Golgi, biochemical binding assay of GRA3 to isolated Golgi membranes, oligomerization analysis Biology open Medium 30814066
2022 GRA3 (Wh6 allele) induces neuronal apoptosis via ER stress by activating the PERK pathway; transfection of N2a cells with GRA3Wh6 increased GRP78, p-PERK, cleaved caspase-12, cleaved caspase-3, and CHOP; PERK inhibitor GSK2656157 and caspase-12 inhibitor Z-ATAD-FMK reduced GRA3-induced apoptosis. Plasmid transfection of N2a cells with pEGFP-GRA3Wh6, flow cytometry (apoptosis), immunoblotting for ER stress/apoptosis markers, pharmacological inhibitors (GSK2656157, Z-ATAD-FMK) Parasites & vectors Medium 35918751
2023 GRA3-deficient T. gondii cysts showed significantly enhanced elimination by anti-cyst CD8+ T cells (perforin-mediated) compared to wild-type; cysts that persisted in the presence of CD8+ T cells had significantly greater GRA3 mRNA levels than the total cyst population; GRA3 mRNA levels correlated with successful cyst persistence during active T-cell immunity. GRA3-deficient parasite strain, in vivo cyst burden measurement in BALB/c mice with CD8+ T-cell immunity, qRT-PCR of GRA3 mRNA in persisting vs. total cysts, correlation analysis Frontiers in immunology Medium 37868957
2019 Deletion of GRA3 (PVM-associated) did not affect the antitumor immunity activated by uracil auxotrophic T. gondii mutants, in contrast to deletions of GRA2, GRA12, GRA24, ROP5, ROP17, ROP18, and ROP35, which severely abrogated the antitumor response. Targeted gene deletion in Δku80 vaccine strain, tumor model (established ovarian cancer), flow cytometry, cytokine measurement PLoS genetics Medium 27447180
2026 GRA3 interacts with host STING, promotes STING oligomerization, and nuclear translocation of phosphorylated IRF3, enhancing IFN-β production via the cGAS-STING pathway; GRA3 also promotes STAT6 phosphorylation; the net effect is enhanced parasite proliferation mediated by ISG56 and STAT6 rather than IFN-β itself. Co-immunoprecipitation (GRA3-STING interaction), STING oligomerization assay, nuclear fractionation (p-IRF3 translocation), gene KO/overexpression, IFN-β ELISA, ISG56/STAT6 inhibitor/KO experiments, parasite proliferation assays PLoS neglected tropical diseases Medium 41712610
2025 CRISPR/Cas9-generated ME49Δgra3 parasites show reduced plaque number, a lower proportion of PVs with ≥4 tachyzoites, and significantly increased host mortality in C57BL/6 mice (70% vs 10% at 35 days), demonstrating that GRA3 promotes intracellular replication and virulence in the ME49 type II strain. CRISPR/Cas9 gene disruption, Western blot confirmation of protein absence, crystal violet plaque assay, Giemsa staining of PV sizes, mouse survival experiment Zhongguo xue xi chong bing fang zhi za zhi Medium 40730529
2019 GRA3 (along with GRA2, GRA4, GRA5, GRA7, GRA12) was found in a proteomic characterization of purified cyst wall fragments from in vitro T. gondii cysts, establishing GRA3 as a component of the cyst wall. Percoll gradient enrichment of cyst wall fragments, CST1 immunoprecipitation, LC-MS/MS proteomic analysis mBio Medium 31040239
2019 Deletion of GRA3 increased MHC-I antigen presentation of PV-localized parasite antigens to CD8+ T cells in infected macrophages and/or dendritic cells, indicating that GRA3 suppresses antigen presentation; this suppression operated independently of IFN-γ or IRG effectors (unlike GRA7/ROP5/ROP18). Targeted GRA3 deletion, MHC-I antigen presentation assay with CD8+ T cells, macrophage and dendritic cell infection, ±IFN-γ conditions, comparison with IRG-dependent effectors Frontiers in immunology Medium 31555296

Source papers

Stage 0 corpus · 41 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1993 Kinetics and pattern of organelle exocytosis during Toxoplasma gondii/host-cell interaction. Parasitology research 158 8415546
2003 Serotyping of Toxoplasma gondii infections in humans using synthetic peptides. The Journal of infectious diseases 106 12717631
2008 The Toxoplasma gondii dense granule protein GRA7 is phosphorylated upon invasion and forms an unexpected association with the rhoptry proteins ROP2 and ROP4. Infection and immunity 78 18809661
2019 Toxoplasma gondii Parasitophorous Vacuole Membrane-Associated Dense Granule Proteins Orchestrate Chronic Infection and GRA12 Underpins Resistance to Host Gamma Interferon. mBio 75 31266861
2006 Chimeric antigens of Toxoplasma gondii: toward standardization of toxoplasmosis serodiagnosis using recombinant products. Journal of clinical microbiology 73 16757610
1994 A soluble secretory protein of the intracellular parasite Toxoplasma gondii associates with the parasitophorous vacuole membrane through hydrophobic interactions. The Journal of biological chemistry 68 8195173
2019 Enrichment and Proteomic Characterization of the Cyst Wall from In Vitro Toxoplasma gondii Cysts. mBio 65 31040239
2003 Use of an immunoglobulin G avidity assay based on recombinant antigens for diagnosis of primary Toxoplasma gondii infection during pregnancy. Journal of clinical microbiology 65 14662919
2014 Phylogeny and virulence divergency analyses of Toxoplasma gondii isolates from China. Parasites & vectors 61 24678633
1994 Cloning of a cDNA encoding the dense granule protein GRA3 from Toxoplasma gondii. Molecular and biochemical parasitology 56 7739670
2015 Variation detection based on next-generation sequencing of type Chinese 1 strains of Toxoplasma gondii with different virulence from China. BMC genomics 55 26518334
2003 Molecular dissection of the human B-cell response against Toxoplasma gondii infection by lambda display of cDNA libraries. International journal for parasitology 54 12633654
1999 Constitutive calcium-independent release of Toxoplasma gondii dense granules occurs through the NSF/SNAP/SNARE/Rab machinery. The Journal of biological chemistry 50 9891012
2016 Secretion of Rhoptry and Dense Granule Effector Proteins by Nonreplicating Toxoplasma gondii Uracil Auxotrophs Controls the Development of Antitumor Immunity. PLoS genetics 47 27447180
2019 A systematic review on the role of GRA proteins of Toxoplasma gondii in host immunization. Journal of microbiological methods 38 31442457
2007 Purification of Toxoplasma dense granule proteins reveals that they are in complexes throughout the secretory pathway. Molecular and biochemical parasitology 36 17959262
2009 Induction of partial protection against infection with Toxoplasma gondii genotype II by DNA vaccination with recombinant chimeric tachyzoite antigens. Vaccine 34 19368791
2008 Interaction between parasitophorous vacuolar membrane-associated GRA3 and calcium modulating ligand of host cell endoplasmic reticulum in the parasitism of Toxoplasma gondii. The Korean journal of parasitology 34 19127325
2007 Increased efficiency of homologous recombination in Toxoplasma gondii dense granule protein 3 demonstrates that GRA3 is not necessary in cell culture but does contribute to virulence. Molecular and biochemical parasitology 33 17418907
2011 Toxoplasma gondii HLA-B*0702-restricted GRA7(20-28) peptide with adjuvants and a universal helper T cell epitope elicits CD8(+) T cells producing interferon-γ and reduces parasite burden in HLA-B*0702 mice. Human immunology 30 22027386
2005 Toxoplasma gondii dense granule protein 3 (GRA3) is a type I transmembrane protein that possesses a cytoplasmic dilysine (KKXX) endoplasmic reticulum (ER) retrieval motif. Parasitology 30 16149193
1995 Sporozoites of Toxoplasma gondii lack dense-granule protein GRA3 and form a unique parasitophorous vacuole. Molecular and biochemical parasitology 28 8720177
2018 Discovery of new Toxoplasma gondii antigenic proteins using a high throughput protein microarray approach screening sera of murine model infected orally with oocysts and tissue cysts. Parasites & vectors 24 29973272
2005 Host cell binding of GRA10, a novel, constitutively secreted dense granular protein from Toxoplasma gondii. Biochemical and biophysical research communications 22 15850804
2019 Rhoptry and Dense Granule Secreted Effectors Regulate CD8+ T Cell Recognition of Toxoplasma gondii Infected Host Cells. Frontiers in immunology 21 31555296
2019 The Toxoplasma gondii dense granule protein TgGRA3 interacts with host Golgi and dysregulates anterograde transport. Biology open 19 30814066
2011 Development and clinical evaluation of a rapid serodiagnostic test for toxoplasmosis of cats using recombinant SAG1 antigen. The Korean journal of parasitology 19 22072819
2017 Genotyping of polymorphic effectors of Toxoplasma gondii isolates from China. Parasites & vectors 13 29157292
2022 Toxoplasma gondii dense granule protein 3 promotes endoplasmic reticulum stress-induced apoptosis by activating the PERK pathway. Parasites & vectors 12 35918751
2021 Toxoplasma GRA Peptide-Specific Serologic Fingerprints Discriminate Among Major Strains Causing Toxoplasmosis. Frontiers in cellular and infection microbiology 10 33680990
2018 Analysis of Toxoplasma gondii clonal type-specific antibody reactions in experimentally infected turkeys and chickens. International journal for parasitology 10 29969590
2017 Suppressors for Human Epidermal Growth Factor Receptor 2/4 (HER2/4): A New Family of Anti-Toxoplasmic Agents in ARPE-19 Cells. The Korean journal of parasitology 6 29103264
2023 Dense granule protein 3 of Toxoplasma gondii plays a crucial role in the capability of the tissue cysts of the parasite to persist in the presence of anti-cyst CD8+ T cells during the chronic stage of infection. Frontiers in immunology 4 37868957
2014 High expression of water-soluble recombinant antigenic domains of Toxoplasma gondii secretory organelles. The Korean journal of parasitology 4 25246715
2024 A combination of GRA3, GRA6 and GRA7 peptides offer a useful tool for serotyping type II and III Toxoplasma gondii infections in sheep and pigs. Frontiers in cellular and infection microbiology 3 38720960
2021 Proteomics analysis reveals that the proto-oncogene eIF-5A indirectly influences the growth, invasion and replication of Toxoplasma gondii tachyzoite. Parasites & vectors 3 34039408
2025 A Structural In Silico Analysis of Novel Epitopes from Toxoplasma gondii Proteins for the Serodiagnosis of Toxoplasmosis. International journal of molecular sciences 2 40429830
2024 Comparison of the performance of SAG2, GRA6, and GRA7 for serological diagnosis of Toxoplasma gondii infection in cats. Frontiers in veterinary science 2 38898997
2023 Preparation and Preliminary Application of Epitope Peptide-Based Antibody against Toxoplasma gondii GRA3. Tropical medicine and infectious disease 2 36977144
2026 Toxoplasma gondii GRA3 activates interferon-stimulated genes and STAT6 by the cGAS-STING pathway to promote parasite proliferation. PLoS neglected tropical diseases 0 41712610
2025 [Generation of a dense granule protein 3 gene-deficient strain of Toxoplasma gondii and its virulence testing]. Zhongguo xue xi chong bing fang zhi za zhi = Chinese journal of schistosomiasis control 0 40730529