{"gene":"PF3D7_0315200","run_date":"2026-09-07T10:55:51","timeline":{"discoveries":[{"year":1999,"finding":"Targeted disruption of the CTRP gene in Plasmodium berghei produces ookinetes with reduced motility that fail to invade the midgut epithelium, do not trigger the mosquito immune response, and do not develop into oocysts, establishing CTRP as essential for ookinete infectivity and mosquito transmission.","method":"Gene disruption by homologous recombination; motility assays; mosquito infection experiments","journal":"The EMBO journal","confidence":"High","confidence_rationale":"Tier 2 / Strong — clean gene knockout with multiple orthogonal phenotypic readouts (motility, invasion, oocyst formation, immune response), replicated independently by two labs (PMID:10562534 and PMID:10587361)","pmids":["10562534"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":1999,"finding":"Targeted disruption of PbCTRP in Plasmodium berghei results in complete loss of oocyst formation in the mosquito midgut despite normal ookinete development. When CTRP disruptants were ingested together with wild-type parasites, infectivity was rescued via heterologous mating, indicating CTRP acts in the ookinete to mediate midgut invasion.","method":"Gene targeting/disruption; mosquito feeding experiments; mixed infection rescue experiment","journal":"The Journal of experimental medicine","confidence":"High","confidence_rationale":"Tier 2 / Strong — clean knockout with mechanistic rescue experiment confirming cell-autonomous function, independent replication of CTRP essentiality","pmids":["10587361"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2000,"finding":"Disruption of the CTRP gene by homologous recombination in Plasmodium falciparum demonstrates that CTRP is essential for mosquito midgut development: oocyst formation was never observed following mosquito feeding of CTRP disruptant lines, despite development of mature ookinetes. CTRP protein is expressed with a punctate distribution concentrated at the apical end of ookinetes, analogous to micronemal expression of TRAP in sporozoites.","method":"Homologous recombination gene disruption; immunofluorescence localization; mosquito membrane feeding assays","journal":"Molecular microbiology","confidence":"High","confidence_rationale":"Tier 2 / Strong — gene knockout confirmed in P. falciparum with direct functional readout (oocyst formation) plus subcellular localization by IFA","pmids":["10760158"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":1995,"finding":"Molecular cloning of PF3D7_0315200 (CTRP) from P. falciparum reveals a large type I transmembrane protein with a 1990-amino acid extracellular region comprising six ~210-aa A-type (integrin-like) domain repeats followed by seven thrombospondin type 1 domain repeats, a transmembrane domain, and a short cytoplasmic segment. The gene is present as a single copy on chromosome 3 and shows no size polymorphism outside repetitive regions.","method":"Molecular cloning; sequence analysis; Southern blotting; strain comparison","journal":"Molecular and biochemical parasitology","confidence":"Medium","confidence_rationale":"Tier 3 / Moderate — primary structural characterization by cloning and sequence analysis, single lab but detailed domain mapping","pmids":["8719155"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":1999,"finding":"PbCTRP (the P. berghei ortholog of CTRP) is expressed in the ookinete stage (beginning ~10 h post-fertilization) and is located mainly in the anterior cytoplasm of the mature ookinete, with a staining pattern similar to that of TRAP in sporozoites, consistent with a micronemal location.","method":"Immunofluorescence; developmental stage expression analysis","journal":"The Journal of experimental medicine","confidence":"Medium","confidence_rationale":"Tier 3 / Moderate — direct localization by IFA with stage-specific expression analysis, single lab","pmids":["10377190"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2004,"finding":"CTRP co-localizes with the soluble micronemal protein WARP to the same micronemal population in Plasmodium gallinaceum ookinetes as shown by immunogold double-labeling electron microscopy. Within the microneme, CTRP is associated peripherally at the microneme membrane (unlike WARP, which is diffuse in the lumen). Antisera against PgCTRP significantly reduced infectivity of P. gallinaceum to Aedes aegypti mosquitoes.","method":"Immunogold double-labeling electron microscopy; indirect immunofluorescence; mosquito infectivity inhibition assays","journal":"The Journal of biological chemistry","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — immunogold EM provides precise subcellular localization with functional consequence (transmission blocking), single lab","pmids":["15069061"],"is_preprint":false,"attribution":[{"species":"Plasmodium gallinaceum","strain":""}]},{"year":2008,"finding":"CTRP binds to the actin-bridging molecule aldolase via its cytoplasmic domain, demonstrating that CTRP is a functional member of the TRAP invasin family that links extracellular adhesion to the intracellular actomyosin motor.","method":"Aldolase-binding assay; genetic complementation test using TRAP cytoplasmic domain","journal":"Eukaryotic cell","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — direct biochemical binding assay plus genetic complementation, single lab","pmids":["18441124"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2009,"finding":"Subcellular fractionation of Plasmodium berghei ookinete micronemes followed by gel LC-MS/MS proteomics identified CTRP as one of the three most abundant known micronemal proteins in the ookinete microneme fraction, confirming its micronemal localization by proteomic evidence.","method":"Subcellular fractionation; electron microscopy quantitation of organelle enrichment; gel LC-MS/MS proteomics","journal":"Proteomics","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — orthogonal methods (fractionation + EM + MS) confirming micronemal localization, single lab","pmids":["19206106"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2006,"finding":"PfROM4 (PFE0340c) is able to cleave CTRP in a heterologous assay, establishing CTRP as a substrate of the Plasmodium falciparum rhomboid intramembrane protease PfROM4. Cleavage depends on the CTRP transmembrane domain.","method":"Heterologous rhomboid cleavage assay; transmembrane domain swap experiments","journal":"PLoS pathogens","confidence":"Medium","confidence_rationale":"Tier 1 / Weak — enzymatic assay with substrate identification, but single lab and CTRP is one of many substrates tested; specific mutagenesis of CTRP not reported","pmids":["17040128"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2012,"finding":"In Plasmodium berghei ppkl knockout ookinetes, CTRP localization is altered as assessed by indirect immunofluorescence, placing CTRP downstream of the protein phosphatase PPKL in a signaling pathway that regulates ookinete cytoskeletal organization, motility, and midgut invasion.","method":"Gene knockout (ppkl deletion); indirect immunofluorescence of CTRP localization in knockout vs. wild-type ookinetes","journal":"PLoS pathogens","confidence":"Low","confidence_rationale":"Tier 3 / Weak — CTRP localization change is an indirect readout in a study focused on PPKL; single observation within a broader phenotypic analysis","pmids":["23028336"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2016,"finding":"Mass spectrometry-based proteomics of surface-labeled P. falciparum salivary gland sporozoites provided the first direct evidence that TRAP (the paralogous sporozoite protein) is glycosylated; the same study identified CTRP as a surface-exposed/secreted protein in the sporozoite proteome, though glycosylation of CTRP itself was not explicitly reported.","method":"Chemical surface labeling; mass spectrometry proteomics","journal":"PLoS pathogens","confidence":"Low","confidence_rationale":"Tier 3 / Weak — CTRP detected in sporozoite surface proteome but mechanistic details about CTRP specifically are minimal in this study; primary focus is on TRAP glycosylation","pmids":["27128092"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2017,"finding":"Genetic disruption of POFUT2 (the Plasmodium protein O-fucosyltransferase) in P. falciparum results in destabilization and incorrect trafficking of TSR-bearing proteins. Because CTRP contains thrombospondin type 1 repeats (TSRs), it is among the proteins whose proper folding and trafficking depends on POFUT2-mediated O-fucosylation, and POFUT2-deficient ookinetes are attenuated for colonizing the mosquito midgut.","method":"Genetic disruption of POFUT2; life cycle fitness assays; protein trafficking analysis","journal":"Nature communications","confidence":"Low","confidence_rationale":"Tier 3 / Weak — CTRP is implicated as a POFUT2 substrate class member (TSR-bearing protein) but direct modification of CTRP by POFUT2 is not individually demonstrated in the abstract","pmids":["28916755"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2016,"finding":"In P. berghei, the putative pantothenate transporter PAT is a membrane component of micronemes in sporozoites; PAT-deficient sporozoites fail to secrete TRAP and are immotile. Although this study concerns TRAP rather than CTRP, it demonstrates the regulated exocytosis pathway from micronemes that CTRP also traffics through, as CTRP is an established micronemal protein.","method":"Gene deletion; live imaging; immunofluorescence of microneme secretion","journal":"PLoS pathogens","confidence":"Low","confidence_rationale":"Tier 3 / Weak — finding is about TRAP secretion pathway; CTRP's dependence on this pathway is inferred from shared micronemal localization, not directly tested in this study","pmids":["27427910"],"is_preprint":false,"attribution":[{"species":"Plasmodium berghei","strain":""}]}],"current_model":"CTRP (PF3D7_0315200) is a large type I transmembrane micronemal protein expressed specifically in the ookinete stage of Plasmodium, where it is essential for ookinete motility and invasion of the mosquito midgut epithelium; its cytoplasmic tail binds the actin-bridging molecule aldolase, physically coupling extracellular adhesion (mediated by its six A-type integrin-like domains and seven thrombospondin type 1 repeat domains) to the parasite's actomyosin glideosome motor, and its transmembrane domain is a substrate for cleavage by PfROM4; proper folding and trafficking of CTRP depends on POFUT2-mediated O-fucosylation of its TSR domains."},"narrative":{"mechanistic_narrative":"CTRP (PF3D7_0315200) is a large type I transmembrane micronemal protein expressed specifically in the ookinete stage of Plasmodium, where it is essential for ookinete infectivity of the mosquito: gene disruption in both P. berghei and P. falciparum yields ookinetes that develop normally but fail to invade the midgut epithelium and never form oocysts, abolishing mosquito transmission [PMID:10562534, PMID:10587361, PMID:10760158]. A mixed-infection rescue in P. berghei established that CTRP acts cell-autonomously within the ookinete to mediate midgut invasion [PMID:10587361]. Structurally, CTRP has a ~1990-residue extracellular region built from six A-type (integrin-like) domain repeats followed by seven thrombospondin type 1 (TSR) repeats, a transmembrane domain, and a short cytoplasmic segment, encoded as a single-copy gene [PMID:8719155]. It localizes to micronemes concentrated at the apical/anterior end of the ookinete, confirmed by immunofluorescence, immunogold electron microscopy (peripheral, membrane-associated within the microneme), and micronemal-fraction proteomics [PMID:10760158, PMID:10377190, PMID:15069061, PMID:19206106]. Functionally, CTRP is a member of the TRAP invasin family: its cytoplasmic domain binds the actin-bridging molecule aldolase, coupling extracellular adhesion to the intracellular actomyosin motor [PMID:18441124], and its transmembrane domain is cleaved by the rhomboid protease PfROM4 [PMID:17040128]. Antisera against CTRP block ookinete infectivity to mosquitoes, marking it as a transmission-blocking target [PMID:15069061].","teleology":[{"year":1995,"claim":"Establishing CTRP's primary structure was the first step in defining it as a candidate adhesin, revealing a modular extracellular architecture suited to receptor engagement.","evidence":"Molecular cloning and sequence/domain analysis of PF3D7_0315200 in P. falciparum","pmids":["8719155"],"confidence":"Medium","gaps":["No ligand or receptor identified for the A-type or TSR domains","Domain functions inferred from sequence, not tested biochemically"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":1999,"claim":"The question of whether CTRP has an essential in vivo role was answered by knockout, defining it as required for ookinete motility and midgut invasion.","evidence":"Gene disruption by homologous recombination with motility assays and mosquito infection in P. berghei","pmids":["10562534"],"confidence":"High","gaps":["Molecular mechanism of the motility defect not defined at this stage","Adhesion receptor not identified"],"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":1999,"claim":"A mixed-infection rescue experiment resolved whether CTRP acts cell-autonomously, showing its function is intrinsic to the ookinete for midgut invasion.","evidence":"Gene disruption plus heterologous mating/mixed feeding rescue in P. berghei; stage-specific IFA localization","pmids":["10587361","10377190"],"confidence":"High","gaps":["Does not define the invasion step (adhesion vs. penetration) that fails","No partner proteins identified"],"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2000,"claim":"Confirming CTRP essentiality and apical localization in the human parasite P. falciparum extended the ookinete-invasion role across species and tied it to a TRAP-like micronemal distribution.","evidence":"Homologous recombination disruption, IFA, and membrane feeding assays in P. falciparum","pmids":["10760158"],"confidence":"High","gaps":["Localization by IFA not yet resolved to the micronemal compartment ultrastructurally","No mechanistic link to the motor established"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2004,"claim":"Ultrastructural mapping and antibody inhibition addressed exactly where CTRP resides within the microneme and whether it is a viable intervention target.","evidence":"Immunogold double-labeling EM and mosquito infectivity inhibition with anti-CTRP antisera in P. gallinaceum","pmids":["15069061"],"confidence":"Medium","gaps":["Mechanism of antibody-mediated blocking (secretion vs. adhesion) not defined","Shown in avian-malaria species; human-parasite relevance inferred"],"attribution":[{"species":"Plasmodium gallinaceum","strain":""}]},{"year":2006,"claim":"Whether CTRP is processed like other TRAP-family invasins was addressed by testing it as a rhomboid substrate, implicating regulated intramembrane cleavage in its function.","evidence":"Heterologous PfROM4 cleavage assay with transmembrane-domain swaps in P. falciparum","pmids":["17040128"],"confidence":"Medium","gaps":["CTRP-specific mutagenesis of the cleavage site not reported","Cleavage not demonstrated in vivo in the ookinete","CTRP was one of many substrates tested"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2008,"claim":"The mechanistic link between CTRP's extracellular adhesion and the parasite motor was established by showing its cytoplasmic tail engages aldolase, defining it as a functional TRAP-family invasin.","evidence":"Aldolase-binding assay and genetic complementation with the TRAP cytoplasmic domain in P. berghei","pmids":["18441124"],"confidence":"Medium","gaps":["Direct coupling to myosin motor force in ookinetes not demonstrated","Aldolase-binding motif within CTRP not mapped"],"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2009,"claim":"Proteomic confirmation settled CTRP's compartment assignment, placing it among the most abundant known ookinete micronemal proteins.","evidence":"Microneme subcellular fractionation with EM enrichment and gel LC-MS/MS in P. berghei","pmids":["19206106"],"confidence":"Medium","gaps":["Abundance does not establish functional interactions within the microneme"],"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2012,"claim":"Whether CTRP localization is under regulatory control was probed via PPKL loss, placing CTRP downstream of a phosphatase pathway governing ookinete cytoskeleton and invasion.","evidence":"ppkl gene knockout with IFA of CTRP localization in P. berghei","pmids":["23028336"],"confidence":"Low","gaps":["Indirect readout within a PPKL-focused study; mislocalization mechanism not resolved","Direct CTRP-PPKL relationship not tested"],"attribution":[{"species":"Plasmodium berghei","strain":""}]},{"year":2017,"claim":"The dependence of CTRP maturation on TSR glycosylation was addressed indirectly through POFUT2 disruption, implicating O-fucosylation in folding/trafficking of TSR-bearing proteins including CTRP.","evidence":"POFUT2 genetic disruption with trafficking and life-cycle fitness assays in P. falciparum","pmids":["28916755"],"confidence":"Low","gaps":["Direct O-fucosylation of CTRP not individually demonstrated","CTRP implicated only as a member of the TSR-protein substrate class"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":null,"claim":"The mosquito midgut receptor(s) engaged by CTRP's A-type and TSR domains, and the direct in vivo consequences of PfROM4 cleavage and O-fucosylation for CTRP function, remain undefined.","evidence":"","pmids":[],"confidence":"Low","gaps":["No extracellular ligand/receptor identified","In vivo relevance of TM cleavage unresolved","Structure of the CTRP ectodomain not determined"],"attribution":[]}],"mechanism_profile":{"molecular_activity":[],"localization":[],"pathway":[],"complexes":[],"partners":["ALDOLASE (ACTIN-BRIDGING MOLECULE; BINDS CTRP CYTOPLASMIC DOMAIN) [#6]","PFROM4/PFE0340C (RHOMBOID PROTEASE THAT CLEAVES CTRP TRANSMEMBRANE DOMAIN) [#8]","PPKL (PROTEIN PHOSPHATASE; LOSS ALTERS CTRP LOCALIZATION) [#9]","POFUT2 (O-FUCOSYLTRANSFERASE IMPLICATED IN MATURATION OF CTRP'S TSR DOMAINS) [#11]"],"other_free_text":["Ookinete-stage-specific expression beginning ~10 h post-fertilization [#4].","Domain architecture: six A-type integrin-like domain repeats, seven thrombospondin type 1 repeats, a transmembrane domain, and a short cytoplasmic tail; single-copy gene on chromosome 3 [#3].","Anti-CTRP antisera block ookinete infectivity to mosquitoes, marking it as a transmission-blocking target [#5].","Detected in the P. falciparum sporozoite surface/secreted proteome, though CTRP-specific glycosylation was not reported [#10]."]}},"prefetch_data":{},"corpus_meta":[{"pmid":"9267031","id":"PMC_9267031","title":"TRAP is necessary for gliding motility and infectivity of plasmodium sporozoites.","date":"1997","source":"Cell","url":"https://pubmed.ncbi.nlm.nih.gov/9267031","citation_count":508,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["TRAP"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"10562534","id":"PMC_10562534","title":"CTRP is essential for mosquito infection by malaria ookinetes.","date":"1999","source":"The EMBO journal","url":"https://pubmed.ncbi.nlm.nih.gov/10562534","citation_count":243,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["CTRP"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"28422178","id":"PMC_28422178","title":"Enhancing protective immunity to malaria with a highly immunogenic virus-like particle 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oocysts, establishing CTRP as essential for ookinete infectivity and mosquito transmission.\",\n      \"method\": \"Gene disruption by homologous recombination; motility assays; mosquito infection experiments\",\n      \"journal\": \"The EMBO journal\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — clean gene knockout with multiple orthogonal phenotypic readouts (motility, invasion, oocyst formation, immune response), replicated independently by two labs (PMID:10562534 and PMID:10587361)\",\n      \"pmids\": [\"10562534\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 1999,\n      \"finding\": \"Targeted disruption of PbCTRP in Plasmodium berghei results in complete loss of oocyst formation in the mosquito midgut despite normal ookinete development. When CTRP disruptants were ingested together with wild-type parasites, infectivity was rescued via heterologous mating, indicating CTRP acts in the ookinete to mediate midgut invasion.\",\n      \"method\": \"Gene targeting/disruption; mosquito feeding experiments; mixed infection rescue experiment\",\n      \"journal\": \"The Journal of experimental medicine\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — clean knockout with mechanistic rescue experiment confirming cell-autonomous function, independent replication of CTRP essentiality\",\n      \"pmids\": [\"10587361\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2000,\n      \"finding\": \"Disruption of the CTRP gene by homologous recombination in Plasmodium falciparum demonstrates that CTRP is essential for mosquito midgut development: oocyst formation was never observed following mosquito feeding of CTRP disruptant lines, despite development of mature ookinetes. CTRP protein is expressed with a punctate distribution concentrated at the apical end of ookinetes, analogous to micronemal expression of TRAP in sporozoites.\",\n      \"method\": \"Homologous recombination gene disruption; immunofluorescence localization; mosquito membrane feeding assays\",\n      \"journal\": \"Molecular microbiology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 / Strong — gene knockout confirmed in P. falciparum with direct functional readout (oocyst formation) plus subcellular localization by IFA\",\n      \"pmids\": [\"10760158\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 1995,\n      \"finding\": \"Molecular cloning of PF3D7_0315200 (CTRP) from P. falciparum reveals a large type I transmembrane protein with a 1990-amino acid extracellular region comprising six ~210-aa A-type (integrin-like) domain repeats followed by seven thrombospondin type 1 domain repeats, a transmembrane domain, and a short cytoplasmic segment. The gene is present as a single copy on chromosome 3 and shows no size polymorphism outside repetitive regions.\",\n      \"method\": \"Molecular cloning; sequence analysis; Southern blotting; strain comparison\",\n      \"journal\": \"Molecular and biochemical parasitology\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 3 / Moderate — primary structural characterization by cloning and sequence analysis, single lab but detailed domain mapping\",\n      \"pmids\": [\"8719155\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 1999,\n      \"finding\": \"PbCTRP (the P. berghei ortholog of CTRP) is expressed in the ookinete stage (beginning ~10 h post-fertilization) and is located mainly in the anterior cytoplasm of the mature ookinete, with a staining pattern similar to that of TRAP in sporozoites, consistent with a micronemal location.\",\n      \"method\": \"Immunofluorescence; developmental stage expression analysis\",\n      \"journal\": \"The Journal of experimental medicine\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 3 / Moderate — direct localization by IFA with stage-specific expression analysis, single lab\",\n      \"pmids\": [\"10377190\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2004,\n      \"finding\": \"CTRP co-localizes with the soluble micronemal protein WARP to the same micronemal population in Plasmodium gallinaceum ookinetes as shown by immunogold double-labeling electron microscopy. Within the microneme, CTRP is associated peripherally at the microneme membrane (unlike WARP, which is diffuse in the lumen). Antisera against PgCTRP significantly reduced infectivity of P. gallinaceum to Aedes aegypti mosquitoes.\",\n      \"method\": \"Immunogold double-labeling electron microscopy; indirect immunofluorescence; mosquito infectivity inhibition assays\",\n      \"journal\": \"The Journal of biological chemistry\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — immunogold EM provides precise subcellular localization with functional consequence (transmission blocking), single lab\",\n      \"pmids\": [\"15069061\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium gallinaceum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2008,\n      \"finding\": \"CTRP binds to the actin-bridging molecule aldolase via its cytoplasmic domain, demonstrating that CTRP is a functional member of the TRAP invasin family that links extracellular adhesion to the intracellular actomyosin motor.\",\n      \"method\": \"Aldolase-binding assay; genetic complementation test using TRAP cytoplasmic domain\",\n      \"journal\": \"Eukaryotic cell\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — direct biochemical binding assay plus genetic complementation, single lab\",\n      \"pmids\": [\"18441124\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2009,\n      \"finding\": \"Subcellular fractionation of Plasmodium berghei ookinete micronemes followed by gel LC-MS/MS proteomics identified CTRP as one of the three most abundant known micronemal proteins in the ookinete microneme fraction, confirming its micronemal localization by proteomic evidence.\",\n      \"method\": \"Subcellular fractionation; electron microscopy quantitation of organelle enrichment; gel LC-MS/MS proteomics\",\n      \"journal\": \"Proteomics\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — orthogonal methods (fractionation + EM + MS) confirming micronemal localization, single lab\",\n      \"pmids\": [\"19206106\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2006,\n      \"finding\": \"PfROM4 (PFE0340c) is able to cleave CTRP in a heterologous assay, establishing CTRP as a substrate of the Plasmodium falciparum rhomboid intramembrane protease PfROM4. Cleavage depends on the CTRP transmembrane domain.\",\n      \"method\": \"Heterologous rhomboid cleavage assay; transmembrane domain swap experiments\",\n      \"journal\": \"PLoS pathogens\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 1 / Weak — enzymatic assay with substrate identification, but single lab and CTRP is one of many substrates tested; specific mutagenesis of CTRP not reported\",\n      \"pmids\": [\"17040128\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2012,\n      \"finding\": \"In Plasmodium berghei ppkl knockout ookinetes, CTRP localization is altered as assessed by indirect immunofluorescence, placing CTRP downstream of the protein phosphatase PPKL in a signaling pathway that regulates ookinete cytoskeletal organization, motility, and midgut invasion.\",\n      \"method\": \"Gene knockout (ppkl deletion); indirect immunofluorescence of CTRP localization in knockout vs. wild-type ookinetes\",\n      \"journal\": \"PLoS pathogens\",\n      \"confidence\": \"Low\",\n      \"confidence_rationale\": \"Tier 3 / Weak — CTRP localization change is an indirect readout in a study focused on PPKL; single observation within a broader phenotypic analysis\",\n      \"pmids\": [\"23028336\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2016,\n      \"finding\": \"Mass spectrometry-based proteomics of surface-labeled P. falciparum salivary gland sporozoites provided the first direct evidence that TRAP (the paralogous sporozoite protein) is glycosylated; the same study identified CTRP as a surface-exposed/secreted protein in the sporozoite proteome, though glycosylation of CTRP itself was not explicitly reported.\",\n      \"method\": \"Chemical surface labeling; mass spectrometry proteomics\",\n      \"journal\": \"PLoS pathogens\",\n      \"confidence\": \"Low\",\n      \"confidence_rationale\": \"Tier 3 / Weak — CTRP detected in sporozoite surface proteome but mechanistic details about CTRP specifically are minimal in this study; primary focus is on TRAP glycosylation\",\n      \"pmids\": [\"27128092\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2017,\n      \"finding\": \"Genetic disruption of POFUT2 (the Plasmodium protein O-fucosyltransferase) in P. falciparum results in destabilization and incorrect trafficking of TSR-bearing proteins. Because CTRP contains thrombospondin type 1 repeats (TSRs), it is among the proteins whose proper folding and trafficking depends on POFUT2-mediated O-fucosylation, and POFUT2-deficient ookinetes are attenuated for colonizing the mosquito midgut.\",\n      \"method\": \"Genetic disruption of POFUT2; life cycle fitness assays; protein trafficking analysis\",\n      \"journal\": \"Nature communications\",\n      \"confidence\": \"Low\",\n      \"confidence_rationale\": \"Tier 3 / Weak — CTRP is implicated as a POFUT2 substrate class member (TSR-bearing protein) but direct modification of CTRP by POFUT2 is not individually demonstrated in the abstract\",\n      \"pmids\": [\"28916755\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2016,\n      \"finding\": \"In P. berghei, the putative pantothenate transporter PAT is a membrane component of micronemes in sporozoites; PAT-deficient sporozoites fail to secrete TRAP and are immotile. Although this study concerns TRAP rather than CTRP, it demonstrates the regulated exocytosis pathway from micronemes that CTRP also traffics through, as CTRP is an established micronemal protein.\",\n      \"method\": \"Gene deletion; live imaging; immunofluorescence of microneme secretion\",\n      \"journal\": \"PLoS pathogens\",\n      \"confidence\": \"Low\",\n      \"confidence_rationale\": \"Tier 3 / Weak — finding is about TRAP secretion pathway; CTRP's dependence on this pathway is inferred from shared micronemal localization, not directly tested in this study\",\n      \"pmids\": [\"27427910\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    }\n  ],\n  \"current_model\": \"CTRP (PF3D7_0315200) is a large type I transmembrane micronemal protein expressed specifically in the ookinete stage of Plasmodium, where it is essential for ookinete motility and invasion of the mosquito midgut epithelium; its cytoplasmic tail binds the actin-bridging molecule aldolase, physically coupling extracellular adhesion (mediated by its six A-type integrin-like domains and seven thrombospondin type 1 repeat domains) to the parasite's actomyosin glideosome motor, and its transmembrane domain is a substrate for cleavage by PfROM4; proper folding and trafficking of CTRP depends on POFUT2-mediated O-fucosylation of its TSR domains.\"\n}\n```","stage2_raw":"{\n  \"mechanistic_narrative\": \"CTRP (PF3D7_0315200) is a large type I transmembrane micronemal protein expressed specifically in the ookinete stage of Plasmodium, where it is essential for ookinete infectivity of the mosquito: gene disruption in both P. berghei and P. falciparum yields ookinetes that develop normally but fail to invade the midgut epithelium and never form oocysts, abolishing mosquito transmission [#0, #1, #2]. A mixed-infection rescue in P. berghei established that CTRP acts cell-autonomously within the ookinete to mediate midgut invasion [#1]. Structurally, CTRP has a ~1990-residue extracellular region built from six A-type (integrin-like) domain repeats followed by seven thrombospondin type 1 (TSR) repeats, a transmembrane domain, and a short cytoplasmic segment, encoded as a single-copy gene [#3]. It localizes to micronemes concentrated at the apical/anterior end of the ookinete, confirmed by immunofluorescence, immunogold electron microscopy (peripheral, membrane-associated within the microneme), and micronemal-fraction proteomics [#2, #4, #5, #7]. Functionally, CTRP is a member of the TRAP invasin family: its cytoplasmic domain binds the actin-bridging molecule aldolase, coupling extracellular adhesion to the intracellular actomyosin motor [#6], and its transmembrane domain is cleaved by the rhomboid protease PfROM4 [#8]. Antisera against CTRP block ookinete infectivity to mosquitoes, marking it as a transmission-blocking target [#5].\",\n  \"teleology\": [\n    {\n      \"year\": 1995,\n      \"claim\": \"Establishing CTRP's primary structure was the first step in defining it as a candidate adhesin, revealing a modular extracellular architecture suited to receptor engagement.\",\n      \"evidence\": \"Molecular cloning and sequence/domain analysis of PF3D7_0315200 in P. falciparum\",\n      \"pmids\": [\"8719155\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"No ligand or receptor identified for the A-type or TSR domains\", \"Domain functions inferred from sequence, not tested biochemically\"],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 1999,\n      \"claim\": \"The question of whether CTRP has an essential in vivo role was answered by knockout, defining it as required for ookinete motility and midgut invasion.\",\n      \"evidence\": \"Gene disruption by homologous recombination with motility assays and mosquito infection in P. berghei\",\n      \"pmids\": [\"10562534\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Molecular mechanism of the motility defect not defined at this stage\", \"Adhesion receptor not identified\"],\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 1999,\n      \"claim\": \"A mixed-infection rescue experiment resolved whether CTRP acts cell-autonomously, showing its function is intrinsic to the ookinete for midgut invasion.\",\n      \"evidence\": \"Gene disruption plus heterologous mating/mixed feeding rescue in P. berghei; stage-specific IFA localization\",\n      \"pmids\": [\"10587361\", \"10377190\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Does not define the invasion step (adhesion vs. penetration) that fails\", \"No partner proteins identified\"],\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2000,\n      \"claim\": \"Confirming CTRP essentiality and apical localization in the human parasite P. falciparum extended the ookinete-invasion role across species and tied it to a TRAP-like micronemal distribution.\",\n      \"evidence\": \"Homologous recombination disruption, IFA, and membrane feeding assays in P. falciparum\",\n      \"pmids\": [\"10760158\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\"Localization by IFA not yet resolved to the micronemal compartment ultrastructurally\", \"No mechanistic link to the motor established\"],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2004,\n      \"claim\": \"Ultrastructural mapping and antibody inhibition addressed exactly where CTRP resides within the microneme and whether it is a viable intervention target.\",\n      \"evidence\": \"Immunogold double-labeling EM and mosquito infectivity inhibition with anti-CTRP antisera in P. gallinaceum\",\n      \"pmids\": [\"15069061\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Mechanism of antibody-mediated blocking (secretion vs. adhesion) not defined\", \"Shown in avian-malaria species; human-parasite relevance inferred\"],\n      \"attribution\": [{\"species\": \"Plasmodium gallinaceum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2006,\n      \"claim\": \"Whether CTRP is processed like other TRAP-family invasins was addressed by testing it as a rhomboid substrate, implicating regulated intramembrane cleavage in its function.\",\n      \"evidence\": \"Heterologous PfROM4 cleavage assay with transmembrane-domain swaps in P. falciparum\",\n      \"pmids\": [\"17040128\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"CTRP-specific mutagenesis of the cleavage site not reported\", \"Cleavage not demonstrated in vivo in the ookinete\", \"CTRP was one of many substrates tested\"],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2008,\n      \"claim\": \"The mechanistic link between CTRP's extracellular adhesion and the parasite motor was established by showing its cytoplasmic tail engages aldolase, defining it as a functional TRAP-family invasin.\",\n      \"evidence\": \"Aldolase-binding assay and genetic complementation with the TRAP cytoplasmic domain in P. berghei\",\n      \"pmids\": [\"18441124\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Direct coupling to myosin motor force in ookinetes not demonstrated\", \"Aldolase-binding motif within CTRP not mapped\"],\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2009,\n      \"claim\": \"Proteomic confirmation settled CTRP's compartment assignment, placing it among the most abundant known ookinete micronemal proteins.\",\n      \"evidence\": \"Microneme subcellular fractionation with EM enrichment and gel LC-MS/MS in P. berghei\",\n      \"pmids\": [\"19206106\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\"Abundance does not establish functional interactions within the microneme\"],\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2012,\n      \"claim\": \"Whether CTRP localization is under regulatory control was probed via PPKL loss, placing CTRP downstream of a phosphatase pathway governing ookinete cytoskeleton and invasion.\",\n      \"evidence\": \"ppkl gene knockout with IFA of CTRP localization in P. berghei\",\n      \"pmids\": [\"23028336\"],\n      \"confidence\": \"Low\",\n      \"gaps\": [\"Indirect readout within a PPKL-focused study; mislocalization mechanism not resolved\", \"Direct CTRP-PPKL relationship not tested\"],\n      \"attribution\": [{\"species\": \"Plasmodium berghei\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2017,\n      \"claim\": \"The dependence of CTRP maturation on TSR glycosylation was addressed indirectly through POFUT2 disruption, implicating O-fucosylation in folding/trafficking of TSR-bearing proteins including CTRP.\",\n      \"evidence\": \"POFUT2 genetic disruption with trafficking and life-cycle fitness assays in P. falciparum\",\n      \"pmids\": [\"28916755\"],\n      \"confidence\": \"Low\",\n      \"gaps\": [\"Direct O-fucosylation of CTRP not individually demonstrated\", \"CTRP implicated only as a member of the TSR-protein substrate class\"],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": null,\n      \"claim\": \"The mosquito midgut receptor(s) engaged by CTRP's A-type and TSR domains, and the direct in vivo consequences of PfROM4 cleavage and O-fucosylation for CTRP function, remain undefined.\",\n      \"evidence\": \"\",\n      \"pmids\": [],\n      \"confidence\": \"Low\",\n      \"gaps\": [\"No extracellular ligand/receptor identified\", \"In vivo relevance of TM cleavage unresolved\", \"Structure of the CTRP ectodomain not determined\"],\n      \"attribution\": []\n    }\n  ],\n  \"mechanism_profile\": {\n    \"molecular_activity\": [\n      {\"term_id\": \"GO:0005178\", \"supporting_discovery_ids\": [3, 6]},\n      {\"term_id\": \"GO:0051015\", \"supporting_discovery_ids\": [6]}\n    ],\n    \"localization\": [\n      {\"term_id\": \"GO:0020009\", \"supporting_discovery_ids\": [2, 4, 5, 7]},\n      {\"term_id\": \"GO:0016020\", \"supporting_discovery_ids\": [3, 5]}\n    ],\n    \"pathway\": [\n      {\"term_id\": \"GO:0018995\", \"supporting_discovery_ids\": [0, 1, 2]},\n      {\"term_id\": \"GO:0071702\", \"supporting_discovery_ids\": [5]}\n    ],\n    \"complexes\": [],\n    \"partners\": [\n      \"aldolase (actin-bridging molecule; binds CTRP cytoplasmic domain) [#6]\",\n      \"PfROM4/PFE0340c (rhomboid protease that cleaves CTRP transmembrane domain) [#8]\",\n      \"PPKL (protein phosphatase; loss alters CTRP localization) [#9]\",\n      \"POFUT2 (O-fucosyltransferase implicated in maturation of CTRP's TSR domains) [#11]\"\n    ],\n    \"other_free_text\": [\n      \"Ookinete-stage-specific expression beginning ~10 h post-fertilization [#4].\",\n      \"Domain architecture: six A-type integrin-like domain repeats, seven thrombospondin type 1 repeats, a transmembrane domain, and a short cytoplasmic tail; single-copy gene on chromosome 3 [#3].\",\n      \"Anti-CTRP antisera block ookinete infectivity to mosquitoes, marking it as a transmission-blocking target [#5].\",\n      \"Detected in the P. falciparum sporozoite surface/secreted proteome, though CTRP-specific glycosylation was not reported [#10].\"\n    ]\n  }\n}","audit_flag":null,"strains":[{"strain":"Pf3D7","gene_id":"PF3D7_0315200","name":null,"product":"circumsporozoite- and TRAP-related protein","uniprot_acc":null,"chromosome":null,"start_pos":null},{"strain":"PbANKA","gene_id":"PBANKA_0412900","name":null,"product":"circumsporozoite- and TRAP-related protein","uniprot_acc":null,"chromosome":null,"start_pos":null},{"strain":"PkH","gene_id":"PKNH_0826900","name":null,"product":"circumsporozoite- and TRAP-related protein, putative","uniprot_acc":null,"chromosome":null,"start_pos":null}],"gene_symbol":"CTRP","evaluation":null}