Affinage · Toxoplasma gondii

ROP5

rhoptry protein ROP5 · UniProt A0A125YQ30

Annotated
2026-09-10
96 papers in source corpus 18 papers cited in narrative 17 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ROP5 is a polymorphic, tandemly-encoded rhoptry pseudokinase secreted into the host cell during Toxoplasma gondii invasion, where it functions as the central scaffold of a parasitophorous vacuole membrane (PVM)-associated kinase complex that neutralizes the IFN-γ-driven cell-autonomous immune response and is indispensable for both acute virulence and chronic infection (PMID:21436047, PMID:21586633, PMID:27165797, PMID:37269286). It associates with the PVM with its C-terminus exposed to the host cytoplasm and adopts a distorted ATP-binding pseudokinase fold in which a non-canonical motif replaces the catalytic aspartate (PMID:16879455, PMID:21708941). Mechanistically, ROP5 directly binds immunity-related GTPases (IRGs) through its most polymorphic surface, allosterically locking them in an inactive GDP-bound state, blocking oligomerization, and reorienting switch-region I threonines for phosphorylation (PMID:22761577, PMID:22802726, PMID:25118287); it simultaneously binds and activates the kinases ROP18 and ROP17—together with GRA7—so that these threonines are permanently phosphorylated, preventing IRG and GBP recruitment to the PVM and Atg5-dependent parasite clearance (PMID:23144612, PMID:23633952, PMID:24832449, PMID:26247512). Among its isoforms, ROP5B is the principal virulence determinant in laboratory mice, and its target specificity underlies host-parasite co-adaptation, with wild-derived mouse resistance conferred by an IRG (Irgb2-b1) that binds ROP5 isoform B (PMID:30874554, PMID:40703524). Beyond IRG antagonism, ROP5 interacts with STING and promotes its K63-linked ubiquitination to enhance cGAS-STING type I IFN signaling (PMID:39457045), and it modulates CD8 T cell IFNγ responses (PMID:32853276). Its PVM association, and thus its downstream functions, is disrupted by host iNOS-dependent cysteine S-nitrosylation (PMID:41427312).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2006 Medium

    Establishing where ROP5 acts required defining its trafficking and topology, showing it reaches the host-facing PVM rather than remaining an intra-parasite protein.

    Evidence Differential permeabilization, immunofluorescence and Western blot in infected cells

    PMID:16879455

    Open questions at the time
    • Molecular function at the PVM not yet defined
    • Mechanism anchoring the C-terminus to the host-cytoplasm face unknown
  2. 2011 High

    Genetic deletion of the entire ROP5 locus resolved whether ROP5 is functionally important, establishing it as indispensable for acute virulence.

    Evidence ROP5 locus deletion with complementation rescue and mouse virulence assay in type I and II strains

    PMID:21436047 PMID:21586633

    Open questions at the time
    • Which host pathway ROP5 antagonizes not yet identified
    • Contribution of individual tandem copies unresolved
  3. 2011 High

    Crystal structures answered whether ROP5 is a catalytically active kinase, revealing it as a pseudokinase that binds ATP in a distorted conformation and mapping polymorphisms to functional surfaces.

    Evidence X-ray crystallography (ATP-bound and apo), site-directed mutagenesis, in vivo virulence assay in type I

    PMID:21708941

    Open questions at the time
    • Binding partners engaged by the polymorphic surfaces not yet defined
    • Functional role of ATP binding unclear
  4. 2012 High

    Multiple studies established the core mechanism: ROP5 directly binds IRGs, is required for ROP18 function, and enforces an inactive GDP-bound IRG conformation that exposes phospho-acceptor threonines for permanent inactivation.

    Evidence Co-IP, genetic epistasis (rop5/rop18 knockouts), GDP/GTP-exchange and IRG phosphorylation assays, in vitro kinase assays, Irgm3-/- mice and macrophages

    PMID:22761577 PMID:22802726 PMID:23144612

    Open questions at the time
    • Structural basis of the allosteric change not yet resolved
    • Composition of the full PVM complex incomplete
  5. 2013 High

    Broadening the effector target and testing cross-species conservation showed ROP5 also blocks Gbp1 recruitment/Atg5-dependent clearance and that Hammondia hammondi orthologs are functional in T. gondii.

    Evidence Gbp1 recruitment immunofluorescence with Gbp1-/- and Atg5-deficient macrophages; heterologous complementation with HhROP5 and mouse virulence assay

    PMID:23633952 PMID:23589877

    Open questions at the time
    • Whether GBP antagonism is direct or downstream of IRG control unresolved
  6. 2014 High

    Complex composition and the structural basis of IRG inactivation were resolved: ROP5 scaffolds ROP18 and ROP17 which phosphorylate adjacent switch-I threonines, and ROP5's polymorphic surface binds the IRG nucleotide-binding domain to block oligomerization allosterically.

    Evidence TAP-MS complex identification, biochemical substrate mapping, allosteric/oligomerization binding assays, mutagenesis, macrophage clearance and mouse virulence assays in type I

    PMID:24832449 PMID:25118287

    Open questions at the time
    • High-resolution ROP5-IRG co-structure not reported
    • Determinants of ROP5 isoform-specific IRG partner selection incomplete
  7. 2015 High

    The complex and virulence role were extended: GRA7 was identified as a direct ROP5-binding PVM component required for efficient IRG phosphorylation, and CRISPR deletion across divergent South American strains confirmed ROP5 as a broadly conserved virulence determinant genetically interacting with ROP18.

    Evidence Co-IP (GRA7-ROP5), Irga6 phosphorylation and IRG recruitment assays; CRISPR-Cas9 knockout across type 10/8/4 strains and two-locus QTL scan

    PMID:26247512 PMID:26291965

    Open questions at the time
    • Stoichiometry and assembly order of the ROP5/ROP18/ROP17/GRA7 complex unresolved
  8. 2016 High

    Deletion in type II parasites showed ROP5 is essential for chronic as well as acute infection, extending its role beyond acute lethality.

    Evidence Deletion of 26 ROPK loci including ROP5, cyst burden measurement and IRG resistance assays in mouse models (type II)

    PMID:27165797

    Open questions at the time
    • Mechanism of ROP5 action during chronic-stage infection not dissected
  9. 2019 High

    Host-parasite co-adaptation was demonstrated: wild-derived mouse resistance is conferred by an IRG (Irgb2-b1) that directly binds ROP5 isoform B, and South American strains escape via polymorphic ROP5 isoforms.

    Evidence Direct binding assay (Irgb2-b1 to ROP5B), genetic analysis of wild-derived mouse resistance, mouse virulence assays and population genetics

    PMID:30874554

    Open questions at the time
    • Full repertoire of IRG partners for each ROP5 isoform not mapped
  10. 2020 Medium

    Additional immune-modulatory roles emerged: ROP5 suppresses CD8 T cell IFNγ responses independently of virulence/IRG interactions, with maximal responses requiring host NLRP3/NLRP1.

    Evidence ROP5 knockout and multiple isoform variants tested in transnuclear CD8 T cell (T57) assays with NLRP3/NLRP1-deficient BMDMs

    PMID:32853276

    Open questions at the time
    • Molecular target through which ROP5 suppresses T cell IFNγ not identified
    • Relationship to inflammasome sensors unresolved
  11. 2023 Medium

    An in vivo CRISPR screen with Ifngr1-/- comparators established that ROP5 virulence is specifically IFN-γ-dependent, consolidating its role as a counter-immune effector.

    Evidence In vivo CRISPR screen in virulent T. gondii-infected C57BL/6 vs Ifngr1-/- mice (type I)

    PMID:37269286

    Open questions at the time
    • Screen does not add molecular mechanism beyond confirming IFN-γ dependence
  12. 2024 Medium

    A distinct signaling role was defined: ROP5 binds STING via predicted domain 2 and enhances its K63-linked ubiquitination (domain 3-dependent) to promote cGAS-STING type I IFN responses.

    Evidence Co-IP (ROP5-STING), K63-linked ubiquitination assay, domain deletion/mutagenesis, ROP5 knockout PRU replication assay in RAW264.7 macrophages

    PMID:39457045

    Open questions at the time
    • How enhancing type I IFN benefits the parasite is not reconciled with its IRG-antagonist role
    • Single-lab Co-IP without reciprocal structural validation
  13. 2025 High

    Systematic isoform dissection established ROP5B as the principal virulence determinant in laboratory mice, tying isoform-specific IRG antagonism to the phenotype.

    Evidence Reverse genetics expressing individual ROP5 isoforms (A, B, C) in RHΔrop5, mouse virulence, IRG recruitment and macrophage clearance assays (type I RH)

    PMID:40703524

    Open questions at the time
    • Host-species targets of ROP5A and ROP5C not identified
  14. 2025 Medium

    A host countermeasure was identified: iNOS-dependent cysteine S-nitrosylation of ROP5 strips it from the PVM, disabling its functions and rescuing susceptibility of iNOS-deficient mice.

    Evidence SNO proteomics, PVM association assay ± RNS, ROP5 knockout in type I and II, macrophage clearance and iNOS-/- mouse infection (preprint)

    PMID:41427312

    Open questions at the time
    • Specific nitrosylated cysteines not fully mapped
    • Preprint not yet peer-reviewed

Mechanism profile

Synthesis pass · controlled-vocabulary classification
Partners
GRA7 (DIRECT BINDING) [#12]IRG PROTEINS (IMMUNITY-RELATED GTPASES; E.G. IRGA6) [#3, #4, #9]IRGB2-B1 (HOST IRG, ISOFORM-SPECIFIC BINDING TO ROP5B) [#14]ROP17 (ACTIVE KINASE) [#8]ROP18 (ACTIVE KINASE) [#3, #5, #8]STING (VIA PREDICTED DOMAIN 2) [#20]
Complex memberships
ROP5-GRA7-containing PVM kinase complex [#12]ROP5/ROP18/ROP17 kinase complex (TAP-MS identified) [#8]

Evidence

Reading pass · 17 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 ROP5 is a member of the ROP2 protein family that is not proteolytically processed during trafficking to rhoptries. It is secreted during host cell invasion and associates with the parasitophorous vacuole membrane (PVM). Differential permeabilization experiments showed ROP5 exposes its C-terminus toward the host cell cytoplasm, representing an inverted topology compared to ROP2 and ROP4. Differential permeabilization of infected cells, immunofluorescence, Western blot Cellular microbiology Medium 16879455
2011 Deletion of the entire ROP5 locus in virulent type I Toxoplasma gondii caused complete loss of acute virulence in mice, and complementation with ROP5 restored lethality to wild-type levels, establishing ROP5 as indispensable for virulence. The ROP5 locus consists of 4–10 tandem genes encoding pseudokinases that are injected directly into host cells. Genetic deletion of ROP5 locus, complementation, mouse virulence assay, genome-wide association mapping Proceedings of the National Academy of Sciences of the United States of America High 21436047 21586633
2011 Crystal structures of a virulent ROP5 isoform in ATP-bound and unbound states revealed that ROP5 is a pseudokinase that binds ATP in a distorted conformation mediated by unusual magnesium coordination sites. Polymorphisms map to two major surfaces including the activation segment. A conserved non-canonical motif replacing the catalytic Asp is required for full in vivo activity of ROP5. X-ray crystallography (ATP-bound and apo structures), site-directed mutagenesis, in vivo virulence assay The Journal of biological chemistry High 21708941
2012 ROP5 directly interacts with immunity-related GTPases (IRGs) and can reduce IRG accumulation on the PVM even in the absence of ROP18. ROP18 can only inhibit IRG accumulation on the PVM of strains that also express virulent ROP5 alleles, indicating ROP5 is required for ROP18 function against IRGs. Co-immunoprecipitation, genetic deletion of ROP5 and ROP18, IRG recruitment assay by immunofluorescence, mouse virulence assay PLoS pathogens High 22761577
2012 ROP5 proteins act as essential co-factors for ROP18 by binding IRG proteins and enforcing an inactive GDP-dependent conformation on the IRG target, preventing GTP-dependent activation and simultaneously exposing target threonines on switch I loop for phosphorylation by ROP18, resulting in permanent IRG inactivation. Biochemical binding assays, genetic epistasis (ROP5 and ROP18 knockouts), GDP-loading/GTP-exchange assays, IRG phosphorylation assays PLoS biology High 22802726
2012 ROP5 controls virulence by directly regulating the catalytic activity of the active kinase ROP18 in vitro. Both ROP5 and ROP18 are necessary to avoid IRG recruitment and parasite clearance in IFN-γ-activated macrophages. Clearance of Δrop5 mutants was reversed in macrophages lacking Irgm3, and virulence defect was restored in Irgm3−/− mice, placing ROP5 upstream of IRG-mediated clearance. In vitro kinase activity assay with recombinant proteins, genetic epistasis with Irgm3−/− mice and macrophages, IRG recruitment immunofluorescence, mouse virulence assay PLoS pathogens High 23144612
2013 Virulent strains of T. gondii avoided Gbp1 recruitment to the PVM in a strain-dependent manner mediated by ROP18 and the pseudokinase ROP5. Increased Gbp1 recruitment to Δrop5 parasites was associated with clearance in IFN-γ-activated macrophages, a process dependent on the autophagy protein Atg5. Immunofluorescence (Gbp1 recruitment assay), Δrop5 and Δrop18 knockouts, Gbp1−/− and Atg5-deficient macrophages, mouse virulence assay PLoS pathogens High 23633952
2014 ROP5 forms complexes with active kinases ROP18 and ROP17 (identified by tandem affinity purification from T. gondii). ROP18 and ROP17 target adjacent essential threonine residues in switch region I of IRGs. The combined activities of ROP17 and ROP18, coordinated by ROP5, contribute to avoidance of IRG recruitment to the vacuole, protecting the parasite from clearance. Tandem affinity purification/mass spectrometry from T. gondii, biochemical substrate mapping, IRG phosphorylation assays, macrophage clearance assay, mouse virulence assay Cell host & microbe High 24832449
2014 ROP5 uses a highly polymorphic surface to bind adjacent to the nucleotide-binding domain of an IRG, producing an allosteric change in IRG structure that prevents IRG oligomerization and alters the orientation of threonines targeted by ROP17 and ROP18. ROP5 proteins are highly specific in the IRG partners they bind, and this binding surface is the most polymorphic region of ROP5. Binding assays (biochemical), structural/allosteric analysis, mutagenesis, IRG oligomerization assays The Journal of biological chemistry High 25118287
2015 GRA7 associates with the PVM by direct binding to ROP5, and is identified as an additional component of the ROP5/ROP18 kinase complex. Absence of GRA7 reduces phosphorylation of Irga6 and correlates with increased vacuolar IRG loading and attenuated virulence. Co-immunoprecipitation (GRA7–ROP5 direct binding), Irga6 phosphorylation assay, IRG recruitment immunofluorescence, mouse virulence assay Cellular microbiology High 26247512
2016 Deletion of ROP5 in type II T. gondii severely reduced cyst burdens (>100-fold reduced virulence), and Δrop5 parasites were less resistant to host IRGs, establishing ROP5 as essential for chronic infection as well as acute virulence. Genetic deletion of 26 ROPK loci including ROP5, cyst burden measurement, IRG resistance assays, mouse infection models mBio High 27165797
2019 Wild-derived Eurasian mice resist virulent T. gondii because the IRG protein Irgb2-b1CIM directly binds ROP5 isoform B. The Irgb2-b1 binding interface is highly polymorphic and under positive selection. South American T. gondii strains are virulent in wild-derived mice due to polymorphic ROP5 isoforms not targeted by Irgb2-b1CIM, demonstrating co-adaptation between host IRGs and ROP5. Direct binding assay (Irgb2-b1 to ROP5 isoform B), genetic analysis of wild-derived mouse resistance, mouse virulence assay with multiple T. gondii strains, population genetics Nature communications High 30874554
2020 ROP5 suppresses CD8 T cell IFNγ responses to parasite-infected macrophages. Multiple ROP5 isoforms and allele types, including 'avirulent' ROP5A from clade A/D strains, suppress CD8 T cell IFNγ responses independently of parasite virulence or known IRG–ROP5 interactions. Host NLRP3 (inflammasome-independent) and NLRP1 pathways, not ROP5 removal alone, are absolute requirements for maximal T cell IFNγ production. ROP5 knockout parasites, multiple ROP5 isoform/allele variants, transnuclear CD8 T cell (T57) assay with BMDMs, NLRP3/NLRP1-deficient macrophages PLoS pathogens Medium 32853276
2023 In vivo CRISPR screen in C57BL/6 mice identified ROP5 as an IFN-γ-dependent virulence gene. Combined usage of Ifngr1−/− mice as a comparator highlighted ROP5 (along with ROP18, GRA12, GRA45) as genes whose virulence function is specifically dependent on the IFN-γ response. In vivo CRISPR screen in virulent T. gondii-infected mice, comparative analysis with Ifngr1−/− mice Cell reports Medium 37269286
2024 ROP5 interacts with STING through its predicted domain 2 and enhances K63-linked ubiquitination of STING in a predicted domain 3-dependent manner, thereby promoting cGAS-STING-mediated type I IFN responses. ROP5-deficient PRU parasites (PRUΔROP5) induced fewer type I IFN-related immune responses and replicated faster in RAW264.7 macrophages. Co-immunoprecipitation (ROP5–STING interaction), ubiquitination assay (K63-linked), domain deletion/mutagenesis, ROP5 knockout parasite replication assay International journal of molecular sciences Medium 39457045
2025 Among the three major ROP5 isoforms (A, B, C), ROP5B is primarily responsible for the heightened virulence of type I T. gondii in laboratory mice. In vivo virulence correlates with diminished vacuolar IRG protein loading and parasite control in vitro only in the presence of ROP5B but not ROP5A or ROP5C. ROP5A and ROP5C may have co-evolved with IRG proteins in other intermediate host species. Reverse genetics — expression of individual ROP5 isoforms (A, B, C) in RHΔrop5 background, mouse virulence assay, IRG recruitment immunofluorescence, in vitro macrophage clearance assay Frontiers in immunology High 40703524
2025 Cysteine S-nitrosylation (SNO) of ROP5 paralogs occurs in a host iNOS-dependent manner. RNS (reactive nitrogen species) caused loss of ROP5 association with the PVM, which is necessary for known ROP5 functions. ROP5 was required for Type I and Type II parasites to evade IFN-γ-mediated immune clearance in iNOS-deficient macrophages, and infection with ROP5 knockout parasites rescued susceptibility of iNOS-deficient mice to Type II T. gondii. SNO proteomics (iNOS-dependent), PVM association assay (immunofluorescence) ± RNS, ROP5 knockout in Type I and II parasites, macrophage clearance assay, iNOS−/− mouse infection bioRxivpreprint Medium 41427312

Source papers

Stage 0 corpus · 96 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2011 Polymorphic family of injected pseudokinases is paramount in Toxoplasma virulence. Proceedings of the National Academy of Sciences of the United States of America 205 21436047
2014 The Toxoplasma pseudokinase ROP5 forms complexes with ROP18 and ROP17 kinases that synergize to control acute virulence in mice. Cell host & microbe 202 24832449
2011 Virulence differences in Toxoplasma mediated by amplification of a family of polymorphic pseudokinases. Proceedings of the National Academy of Sciences of the United States of America 184 21586633
2012 The rhoptry proteins ROP18 and ROP5 mediate Toxoplasma gondii evasion of the murine, but not the human, interferon-gamma response. PLoS pathogens 177 22761577
2013 Guanylate-binding protein 1 (Gbp1) contributes to cell-autonomous immunity against Toxoplasma gondii. PLoS pathogens 155 23633952
2012 The polymorphic pseudokinase ROP5 controls virulence in Toxoplasma gondii by regulating the active kinase ROP18. PLoS pathogens 139 23144612
2012 A Toxoplasma gondii pseudokinase inhibits host IRG resistance proteins. PLoS biology 116 22802726
2015 The ROP18 and ROP5 gene allele types are highly predictive of virulence in mice across globally distributed strains of Toxoplasma gondii. International journal for parasitology 95 26699401
2015 Rhoptry Proteins ROP5 and ROP18 Are Major Murine Virulence Factors in Genetically Divergent South American Strains of Toxoplasma gondii. PLoS genetics 81 26291965
2015 Toxoplasma gondii superinfection and virulence during secondary infection correlate with the exact ROP5/ROP18 allelic combination. mBio 73 25714710
2016 The Toxoplasma gondii Rhoptry Kinome Is Essential for Chronic Infection. mBio 69 27165797
2013 Structural and evolutionary adaptation of rhoptry kinases and pseudokinases, a family of coccidian virulence factors. BMC evolutionary biology 69 23742205
2015 The Toxoplasma gondii rhoptry protein ROP18 is an Irga6-specific kinase and regulated by the dense granule protein GRA7. Cellular microbiology 65 26247512
2011 A conserved non-canonical motif in the pseudoactive site of the ROP5 pseudokinase domain mediates its effect on Toxoplasma virulence. The Journal of biological chemistry 65 21708941
2014 The Toxoplasma pseudokinase ROP5 is an allosteric inhibitor of the immunity-related GTPases. The Journal of biological chemistry 60 25118287
2006 Inverted topology of the Toxoplasma gondii ROP5 rhoptry protein provides new insights into the association of the ROP2 protein family with the parasitophorous vacuole membrane. Cellular microbiology 59 16879455
2013 Genotyping Toxoplasma gondii from wildlife in Pennsylvania and identification of natural recombinants virulent to mice. Veterinary parasitology 55 24332401
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
2013 The virulence-related rhoptry protein 5 (ROP5) of Toxoplasma Gondii is a novel vaccine candidate against toxoplasmosis in mice. Vaccine 45 23928460
2020 Toxoplasma  GRA15 and GRA24 are important activators of the host innate immune response in the absence of TLR11. PLoS pathogens 42 32453782
2015 DNA vaccination with genes encoding Toxoplasma gondii antigens ROP5 and GRA15 induces protective immunity against toxoplasmosis in Kunming mice. Expert review of vaccines 42 25749394
2013 Hammondia hammondi, an avirulent relative of Toxoplasma gondii, has functional orthologs of known T. gondii virulence genes. Proceedings of the National Academy of Sciences of the United States of America 38 23589877
2019 Comparative virulence of Caribbean, Brazilian and European isolates of Toxoplasma gondii. Parasites & vectors 34 30871587
2023 Host genetics highlights IFN-γ-dependent Toxoplasma genes encoding secreted and non-secreted virulence factors in in vivo CRISPR screens. Cell reports 30 37269286
2019 Molecular mechanism for the control of virulent Toxoplasma gondii infections in wild-derived mice. Nature communications 30 30874554
2018 Clonal and atypical Toxoplasma strain differences in virulence vary with mouse sub-species. International journal for parasitology 27 30471286
2017 Association of ROP18 and ROP5 was efficient as a marker of virulence in atypical isolates of Toxoplasma gondii obtained from pigs and goats in Piauí, Brazil. Veterinary parasitology 27 29080759
2015 Towards vaccine against toxoplasmosis: evaluation of the immunogenic and protective activity of recombinant ROP5 and ROP18 Toxoplasma gondii proteins. Parasitology research 27 26337271
2021 In vivo and in vitro models show unexpected degrees of virulence among Toxoplasma gondii type II and III isolates from sheep. Veterinary research 25 34112256
2021 Third-generation sequencing revises the molecular karyotype for Toxoplasma gondii and identifies emerging copy number variants in sexual recombinants. Genome research 24 33906962
2020 Toxoplasma gondii GRA60 is an effector protein that modulates host cell autonomous immunity and contributes to virulence. Cellular microbiology 24 33040458
2020 Adjuvantic cytokine IL-33 improves the protective immunity of cocktailed DNA vaccine of ROP5 and ROP18 against toxoplasma gondii infection in mice. Parasite (Paris, France) 22 32315596
2014 Impact of regulated secretion on antiparasitic CD8 T cell responses. Cell reports 22 24857659
2019 Rhoptry and Dense Granule Secreted Effectors Regulate CD8+ T Cell Recognition of Toxoplasma gondii Infected Host Cells. Frontiers in immunology 21 31555296
2018 Atypical virulence in a type III Toxoplasma gondii strain isolated in Japan. Parasitology international 20 29775826
2018 Genetic diversity of Toxoplasma gondii isolates obtained from free-living wild birds rescued in Southeastern Brazil. International journal for parasitology. Parasites and wildlife 20 30479944
2018 A Gene-Based Positive Selection Detection Approach to Identify Vaccine Candidates Using Toxoplasma gondii as a Test Case Protozoan Pathogen. Frontiers in genetics 20 30177953
2016 Evaluation of immune responses induced by rhoptry protein 5 and rhoptry protein 7 DNA vaccines against Toxoplasma gondii. Parasite immunology 19 26802673
2016 Corrigendum to "The ROP18 and ROP5 gene allele types are highly predictive of virulence in mice across globally distributed strains of Toxoplasma gondii" [Int. J. Parasitol. 46 (2016) 141-146]. International journal for parasitology 18 27324470
2019 iTRAQ-Based Global Phosphoproteomics Reveals Novel Molecular Differences Between Toxoplasma gondii Strains of Different Genotypes. Frontiers in cellular and infection microbiology 17 31508380
2020 Naïve CD8 T cell IFNγ responses to a vacuolar antigen are regulated by an inflammasome-independent NLRP3 pathway and Toxoplasma gondii ROP5. PLoS pathogens 16 32853276
2019 Toxoplasma gondii in invasive animals on the Island of Fernando de Noronha in Brazil: Molecular characterization and mouse virulence studies of new genotypes. Comparative immunology, microbiology and infectious diseases 15 31546127
2021 Comparative Proteomics Analysis for Elucidating the Interaction Between Host Cells and Toxoplasma gondii. Frontiers in cellular and infection microbiology 14 34055664
2019 Toxoplasma gondii in four captive kangaroos (Macropus spp.) in China: Isolation of a strain of a new genotype from an eastern grey kangaroo (Macropus giganteus). International journal for parasitology. Parasites and wildlife 14 30923671
2017 Genotyping of polymorphic effectors of Toxoplasma gondii isolates from China. Parasites & vectors 13 29157292
2023 ROP39 is an Irgb10-specific parasite effector that modulates acute Toxoplasma gondii virulence. PLoS pathogens 12 36603017
2019 Isolation, genotyping and pathogenicity of a Toxoplasma gondii strain isolated from a Serval (Leptailurus serval) in China. Transboundary and emerging diseases 11 31102311
2019 Evidence of red panda as an intermediate host of Toxoplasma gondii and Sarcocystis species. International journal for parasitology. Parasites and wildlife 11 30891398
2020 Molecular and biological analysis revealed genetic diversity and high virulence strain of Toxoplasma gondii in Japan. PloS one 10 32012177
2020 Isolation and characterization of Toxoplasma gondii from captive caracals (Caracal caracal). International journal for parasitology. Parasites and wildlife 10 33145163
2020 In Vivo and In Vitro Virulence Analysis of Four Genetically Distinct Toxoplasma gondii Lineage III Isolates. Microorganisms 10 33142663
2020 Description of an atypical Toxoplasma gondii isolate from a case of congenital toxoplasmosis in northeastern Brazil. Parasitology research 10 32518965
2021 Toxoplasma gondii isolated from a Brazilian patient with rare pulmonary toxoplasmosis has a novel genotype and is closely related to Amazonian isolates. Parasitology research 9 33420622
2016 Analysis of the virulence determination mechanisms in a local Toxoplasma strain (T.gHB1) isolated from central China. Parasitology research 9 27225000
2015 Toxoplasma gondii: cloning, expression and immunoreactivity of recombinant ROP5 and ROP18 antigens. Experimental parasitology 9 25592730
2023 Genotyping of toxoplasma gondii isolates from México reveals non-archetypal and potentially virulent strains for mice. Infection, genetics and evolution : journal of molecular epidemiology and evolutionary genetics in infectious diseases 8 37353185
2021 ROP18 and ROP5 alleles combinations are related with virulence of T. gondii isolates from Argentina. Parasitology international 8 33713809
2015 Human toxoplasmosis: a comparative evaluation of the diagnostic potential of recombinant Toxoplasma gondii ROP5 and ROP18 antigens. Journal of medical microbiology 8 26242602
2025 IL-24 Is a Promising Molecular Adjuvant for Enhancing Protective Immunity Induced by DNA Vaccination Against Toxoplasma gondii. Microorganisms 7 40732170
2024 Vaccination with a DNA vaccine cocktail encoding TgROP2, TgROP5, TgROP9, TgROP16, TgROP17, and TgROP18 confers limited protection against Toxoplasma gondii in BALB/c mice. Parasitology research 7 39724445
2022 Two viable Toxoplasma gondii isolates from red-necked wallaby (Macropus rufogriseus) and red kangaroo (M. rufus). Parasitology international 7 36216306
2021 Isolation, genotyping and virulence determination of a Toxoplasma gondii strain from non-human primate from China. Transboundary and emerging diseases 7 33619895
2020 Toxoplasma gondii infection in white spoonbills (Platalea leucorodia) from Henan Province, China. Emerging microbes & infections 7 33215979
2022 Additional evidence of tigers (Panthera tigris altaica) as intermediate hosts for Toxoplasma gondii through the isolation of viable strains. International journal for parasitology. Parasites and wildlife 6 36467851
2021 Unraveling Toxoplasma gondii GT1 Strain Virulence and New Protein-Coding Genes with Proteogenomic Analyses. Omics : a journal of integrative biology 6 34468217
2017 Toward detection of toxoplasmosis from urine in mice using hydro-gel nanoparticles concentration and parallel reaction monitoring mass spectrometry. Nanomedicine : nanotechnology, biology, and medicine 6 29203146
2024 Toxoplasma gondii ROP5 Enhances Type I IFN Responses by Promoting Ubiquitination of STING. International journal of molecular sciences 5 39457045
2024 Direct evidence of cheetah (Acinonyx jubatus) as intermediate host of Toxoplasma gondii through isolation of viable strains. BMC veterinary research 5 38395812
2023 Epidemiology and isolation of viable Toxoplasma gondii strain from macropods. Heliyon 5 36938389
2020 Characterization of a spontaneous cyst-forming strain of Toxoplasma gondii isolated from Tokachi subprefecture in Japan. Parasitology international 5 32961305
2015 MSCA: a spectral comparison algorithm between time series to identify protein-protein interactions. BMC bioinformatics 5 25963052
2025 Designing a multi-epitope vaccine using Toxoplasma ROP5, ROP7, and SAG1 epitopes and immunogenicity evaluation against acute and chronic toxoplasmosis in BABL/c mice. Microbial pathogenesis 4 40216097
2024 Genotyping of Toxoplasma gondii in domestic animals from Campeche, México, reveals virulent genotypes and a recombinant ROP5 allele. Parasitology 4 38379406
2023 Prenyl Transferases Regulate Secretory Protein Sorting and Parasite Morphology in Toxoplasma gondii. International journal of molecular sciences 4 37108334
2020 Toxoplasma gondii: Cytokine responses in mice reinfected with atypical strains. Experimental parasitology 4 32991867
2014 Toxoplasma's arms race with the host interferon response: a ménage à trois of ROPs. Cell host & microbe 4 24832444
2025 Isolation of Toxoplasma gondii from the placenta of northern fur seals (Callorhinus ursinus) and potential transplacental transmission of the parasite. Parasite (Paris, France) 3 40749113
2024 Early immune response to Toxoplasma gondii lineage III isolates of different virulence phenotype. Frontiers in cellular and infection microbiology 3 38912206
2023 Detection of Toxoplasma gondii and Sarcocystis sp. in the meat of common minke whale (Balaenoptera acutorostrata): A case of suspected food poisoning in Japan. Parasitology international 3 38040112
2025 New genetic variants of Toxoplasma gondii isolates and a novel ROP5 allele in free-range chickens from Tabasco, México. Parasitology 2 40588766
2025 Toxoplasma gondii virulence in mice is determined by the pseudokinase ROP5B and countered by an IRG-GBP protein interplay. Frontiers in immunology 2 40703524
2025 Investigation of Virulence-Related Markers in Atypical Strains of Toxoplasma gondii from Brazil. Microorganisms 2 40005667
2024 Genetic characterization of Toxoplasma gondii from human and chicken isolates from Argentina. Parasitology research 2 38332310
2023 Isolation and Characterization of a Viable Toxoplasma gondii from Captive Caracal (Caracal caracal). Pathogens (Basel, Switzerland) 2 38133295
2023 Isolation and Genetic Characterization of Toxoplasma gondii from a Patas Monkey (Erythrocebus patas) in China. Genes 2 37628657
2013 [Immune response in mice induced by complex gene vaccine pcSAG1-ROP5 of Toxoplasma gondii]. Zhongguo ji sheng chong xue yu ji sheng chong bing za zhi = Chinese journal of parasitology & parasitic diseases 2 24812827
2025 First molecular characterization of Toxoplasma gondii and molecular analysis of Neospora caninum in American mink (Neogale vison) introduced in Argentina. Research in veterinary science 1 41109155
2025 First isolation of viable Toxoplasma gondii from a black mangabey (Lophocebus aterrimus) reveals the emergence of the Africa 1 lineage in East Asia. PLoS neglected tropical diseases 1 40700374
1997 Murine Th1 clone defines a 60 kD tachyzoite antigen of Toxoplasma gondii. Parasite immunology 1 9194100
2026 Characterization of ovine abortion in Uruguay reveals extensive non-clonal diversity and multiple evolutionary origins of Toxoplasma gondii. Scientific reports 0 42243401
2026 First report of viable Toxoplasma gondii isolation from African lions (Panthera leo): Evidence for their status as intermediate hosts. Food and waterborne parasitology 0 41675383
2026 Functional characterization of 11 novel rhoptry proteins in the type I RH strain of Toxoplasma gondii using the CRISPR-Cas9 system. Parasites & vectors 0 41975467
2025 Cysteine-S-nitrosylation inhibits Rop5-mediated immune evasion in Toxoplasma gondii. bioRxiv : the preprint server for biology 0 41427312
2025 Identification of Toxoplasma gondii antigenic proteins using an in vivo approach and in silico investigation of their polymorphism. Microbiology spectrum 0 40135885
2015 [Research Advances on Toxoplasma gondii Virulence Mediating Factors]. Zhongguo ji sheng chong xue yu ji sheng chong bing za zhi = Chinese journal of parasitology & parasitic diseases 0 26672224
2015 [Research Advance on Toxoplasma gondii Rhoptry Protein 2 Family as Candidate Vaccine Molecules]. Zhongguo ji sheng chong xue yu ji sheng chong bing za zhi = Chinese journal of parasitology & parasitic diseases 0 26541043