{"gene":"PF3D7_0704600","run_date":"2026-09-08T12:51:27","timeline":{"discoveries":[{"year":2014,"finding":"PF3D7_0704600 (UT/MAL7P1.19), a HECT ubiquitin-protein ligase, was associated with altered quinine and quinidine sensitivity in P. falciparum; episomal transfection experiments demonstrated a role in quinine and quinidine sensitivity in a subset of genetic backgrounds, with moderate (~2-fold) changes in IC50 values, indicating a contributing but background-dependent role in drug response.","method":"Quantitative trait locus analysis on F1 progeny of a genetic cross, genetic screens of field isolates and laboratory strains, episomal transfection with phenotypic IC50 measurement","journal":"PLoS genetics","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — genetic cross QTL mapping plus functional episomal transfection in multiple genetic backgrounds, single lab, effect moderate and background-dependent","pmids":["24830312"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2019,"finding":"Overexpression of PF3D7_0704600 (PfUT) in P. falciparum resulted in an S/M phase-associated lengthening of the intraerythrocytic developmental cycle and a reduced merozoite invasion efficiency, establishing a role for PfUT in controlling cell cycle progression and merozoite invasion.","method":"glmS ribozyme system-mediated unintended overexpression (2.5-fold increase in RNA and protein), cell cycle staging, invasion efficiency assays, partial phenotypic rescue with glucosamine","journal":"Scientific reports","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — defined cellular phenotypes (cell cycle lengthening, invasion defect) with partial rescue, single lab, two orthogonal phenotypic readouts","pmids":["31797898"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2020,"finding":"PF3D7_0704600 (a HECT E3 protein-ubiquitin ligase) was identified as a functional target of PfPP1 in the context of parasite egress from host erythrocytes; phosphoproteomic and chemical-genetic analysis placed it downstream of PfPP1 in the egress pathway.","method":"Phosphoproteomic analysis and chemical-genetic analysis of PfPP1","journal":"Nature communications","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — phosphoproteomics combined with chemical-genetic analysis, single lab, two orthogonal methods, but PF3D7_0704600 is one of two identified targets and further mechanistic detail is not provided in the abstract","pmids":["32669539"],"is_preprint":false,"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2017,"finding":"The P. yoelii ortholog of PF3D7_0704600 (Pyheul, a HECT-like E3 ubiquitin ligase) was shown to influence parasitemia and host mortality; allelic exchange of partial Pyheul sequences between two lethal parasite strains and modification of its expression altered parasite growth and host mortality, placing it in a virulence-regulatory pathway.","method":"Genetic cross of P. yoelii strains, genome-wide QTL scan, allelic exchange, gene expression modification with phenotypic readout (parasitemia, host mortality)","journal":"Nature communications","confidence":"Medium","confidence_rationale":"Tier 2 / Moderate — genetic cross, QTL mapping, and allelic exchange in P. yoelii; multiple orthogonal methods, single lab","pmids":["28790316"],"is_preprint":false,"attribution":[{"species":"Plasmodium yoelii","strain":""}]}],"current_model":"PF3D7_0704600 (UT/PfUT) is a HECT-type E3 ubiquitin ligase in P. falciparum that controls intraerythrocytic cell cycle progression (S/M phase duration) and merozoite invasion efficiency when overexpressed, functions as a downstream substrate/target of PfPP1 in the egress pathway, and contributes in a genetic-background-dependent manner to quinine/quinidine sensitivity; the P. yoelii ortholog similarly regulates parasite growth and virulence through allele-dependent mechanisms."},"narrative":{"mechanistic_narrative":"PF3D7_0704600 (UT/PfUT) is a HECT-type E3 ubiquitin ligase of Plasmodium falciparum that regulates intraerythrocytic cell cycle progression and merozoite invasion and functions within the parasite egress pathway [PMID:31797898, PMID:32669539]. Overexpression of PfUT lengthens the S/M phase of the intraerythrocytic developmental cycle and reduces merozoite invasion efficiency, establishing it as a positive controller of cell cycle timing and invasion [PMID:31797898]. Phosphoproteomic and chemical-genetic analyses place PfUT downstream of the protein phosphatase PfPP1 in the pathway governing parasite egress from host erythrocytes, indicating its activity is phosphoregulated [PMID:32669539]. Genetically, PfUT contributes in a background-dependent manner to quinine and quinidine sensitivity, with episomal expression producing moderate (~2-fold) shifts in IC50 in a subset of strains [PMID:24830312]. The P. yoelii ortholog (Pyheul) similarly acts in a virulence-regulatory pathway, where allele-dependent differences and altered expression modulate parasitemia and host mortality [PMID:28790316]. The specific ubiquitination substrates of PfUT, its structural basis, and the mechanistic link between its ligase activity and each phenotype have not been characterized in the available corpus.","teleology":[{"year":2014,"claim":"Establishing whether PF3D7_0704600 contributes to antimalarial drug response addressed whether this HECT ligase has any measurable phenotypic role in P. falciparum.","evidence":"QTL analysis of a genetic cross plus episomal transfection with IC50 measurement across multiple genetic backgrounds","pmids":["24830312"],"confidence":"Medium","gaps":["Effect is moderate (~2-fold) and only seen in a subset of genetic backgrounds","The molecular basis linking ubiquitin ligase activity to quinine/quinidine sensitivity is unknown","No substrate or interacting partner identified for this drug phenotype"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2017,"claim":"Analysis of the P. yoelii ortholog Pyheul tested whether this HECT ligase influences parasite growth and virulence in an in vivo rodent malaria model.","evidence":"Genetic cross of P. yoelii strains, genome-wide QTL scan, and allelic exchange with parasitemia and host-mortality readouts","pmids":["28790316"],"confidence":"Medium","gaps":["Findings are in the P. yoelii ortholog, not PF3D7_0704600 itself","The allele-specific molecular differences driving virulence are not defined","No ubiquitination substrate linked to the virulence phenotype"],"attribution":[{"species":"Plasmodium yoelii","strain":""}]},{"year":2019,"claim":"Defining the cellular consequence of altered PfUT levels revealed a direct role in intraerythrocytic cell cycle timing and merozoite invasion, moving beyond drug-response association.","evidence":"glmS ribozyme-mediated overexpression (2.5-fold), cell cycle staging, invasion efficiency assays, and partial glucosamine rescue in P. falciparum","pmids":["31797898"],"confidence":"Medium","gaps":["Phenotype derives from overexpression rather than loss-of-function","The substrates ubiquitinated by PfUT to control S/M phase are unidentified","Mechanistic link between invasion defect and cell cycle lengthening not resolved"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":2020,"claim":"Positioning PfUT downstream of PfPP1 established that its activity is embedded in the phosphoregulated egress signaling pathway.","evidence":"Phosphoproteomic and chemical-genetic analysis of PfPP1 in P. falciparum","pmids":["32669539"],"confidence":"Medium","gaps":["PfUT is one of two identified PfPP1 targets; direct regulation not fully dissected in the abstract","Whether PfPP1 dephosphorylates PfUT directly and how this modulates ligase activity is unknown","The egress-relevant substrates of PfUT are not identified"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]},{"year":null,"claim":"The ubiquitination substrates of PfUT, its catalytic and structural basis, and how its HECT ligase activity mechanistically drives cell cycle timing, egress, invasion, and drug response remain unresolved.","evidence":"","pmids":[],"confidence":"Medium","gaps":["No direct ubiquitination substrate identified","No structural or biochemical characterization of ligase activity","Loss-of-function phenotype in P. falciparum not defined"],"attribution":[{"species":"Plasmodium falciparum","strain":""}]}],"mechanism_profile":{"molecular_activity":[],"localization":[],"pathway":[],"complexes":[],"partners":["PFPP1 (FUNCTIONAL UPSTREAM REGULATOR IN THE EGRESS PATHWAY) [#2]"],"other_free_text":["Overexpression lengthens the S/M phase of the intraerythrocytic developmental cycle and reduces merozoite invasion efficiency [#1]","Contributes in a genetic-background-dependent manner to quinine and quinidine sensitivity (~2-fold IC50 shifts) [#0]","P. yoelii ortholog Pyheul modulates parasitemia and host mortality in an allele-dependent manner [#3]","No direct ubiquitination substrate, structural model, or loss-of-function phenotype characterized in the available corpus"]}},"prefetch_data":{},"corpus_meta":[{"pmid":"18545708","id":"PMC_18545708","title":"Deciphering the ubiquitin-mediated pathway in apicomplexan parasites: a potential strategy to interfere with parasite virulence.","date":"2008","source":"PloS one","url":"https://pubmed.ncbi.nlm.nih.gov/18545708","citation_count":85,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["curated_ref"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"32669539","id":"PMC_32669539","title":"Co-option of Plasmodium falciparum PP1 for egress from host erythrocytes.","date":"2020","source":"Nature communications","url":"https://pubmed.ncbi.nlm.nih.gov/32669539","citation_count":40,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"24830312","id":"PMC_24830312","title":"A HECT ubiquitin-protein ligase as a novel candidate gene for altered quinine and quinidine responses in Plasmodium falciparum.","date":"2014","source":"PLoS genetics","url":"https://pubmed.ncbi.nlm.nih.gov/24830312","citation_count":31,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"28790316","id":"PMC_28790316","title":"A Plasmodium yoelii HECT-like E3 ubiquitin ligase regulates parasite growth and virulence.","date":"2017","source":"Nature communications","url":"https://pubmed.ncbi.nlm.nih.gov/28790316","citation_count":22,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"31797898","id":"PMC_31797898","title":"Overexpression of the HECT ubiquitin ligase PfUT prolongs the intraerythrocytic cycle and reduces invasion efficiency of Plasmodium falciparum.","date":"2019","source":"Scientific reports","url":"https://pubmed.ncbi.nlm.nih.gov/31797898","citation_count":9,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"35464842","id":"PMC_35464842","title":"Identification of Co-Existing Mutations and Gene Expression Trends Associated With K13-Mediated Artemisinin Resistance in Plasmodium falciparum.","date":"2022","source":"Frontiers in genetics","url":"https://pubmed.ncbi.nlm.nih.gov/35464842","citation_count":9,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false},{"pmid":"40249735","id":"PMC_40249735","title":"Activity-based protein profiling reveals both canonical and novel ubiquitin pathway enzymes in Plasmodium.","date":"2025","source":"PLoS pathogens","url":"https://pubmed.ncbi.nlm.nih.gov/40249735","citation_count":2,"is_preprint":false,"source_species":["Pf"],"role":"primary","match_terms":["HECT"],"competing_organism_flag":false,"competing_organism_match":"","competing_organism_only":false}],"cost":{"stage1":{"model":"claude-sonnet-4-6","input_tokens":6142,"output_tokens":1247,"usd":0.018565,"stage1_stop_reason":"end_turn"},"stage2":{"model":"claude-opus-4-8","input_tokens":6164,"output_tokens":2339,"usd":0.044647,"stage2_stop_reason":"end_turn"},"total_usd":0.063212,"stage1_batch_id":"msgbatch_01DisAaTypwmC8QwdMBuePHo","stage2_batch_id":"msgbatch_0184zxZTqxiEfdYYfPMfkqjb","note":"batch pricing = 50% of standard"},"stage1_raw":"```json\n{\n  \"discoveries\": [\n    {\n      \"year\": 2014,\n      \"finding\": \"PF3D7_0704600 (UT/MAL7P1.19), a HECT ubiquitin-protein ligase, was associated with altered quinine and quinidine sensitivity in P. falciparum; episomal transfection experiments demonstrated a role in quinine and quinidine sensitivity in a subset of genetic backgrounds, with moderate (~2-fold) changes in IC50 values, indicating a contributing but background-dependent role in drug response.\",\n      \"method\": \"Quantitative trait locus analysis on F1 progeny of a genetic cross, genetic screens of field isolates and laboratory strains, episomal transfection with phenotypic IC50 measurement\",\n      \"journal\": \"PLoS genetics\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — genetic cross QTL mapping plus functional episomal transfection in multiple genetic backgrounds, single lab, effect moderate and background-dependent\",\n      \"pmids\": [\"24830312\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2019,\n      \"finding\": \"Overexpression of PF3D7_0704600 (PfUT) in P. falciparum resulted in an S/M phase-associated lengthening of the intraerythrocytic developmental cycle and a reduced merozoite invasion efficiency, establishing a role for PfUT in controlling cell cycle progression and merozoite invasion.\",\n      \"method\": \"glmS ribozyme system-mediated unintended overexpression (2.5-fold increase in RNA and protein), cell cycle staging, invasion efficiency assays, partial phenotypic rescue with glucosamine\",\n      \"journal\": \"Scientific reports\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — defined cellular phenotypes (cell cycle lengthening, invasion defect) with partial rescue, single lab, two orthogonal phenotypic readouts\",\n      \"pmids\": [\"31797898\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2020,\n      \"finding\": \"PF3D7_0704600 (a HECT E3 protein-ubiquitin ligase) was identified as a functional target of PfPP1 in the context of parasite egress from host erythrocytes; phosphoproteomic and chemical-genetic analysis placed it downstream of PfPP1 in the egress pathway.\",\n      \"method\": \"Phosphoproteomic analysis and chemical-genetic analysis of PfPP1\",\n      \"journal\": \"Nature communications\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — phosphoproteomics combined with chemical-genetic analysis, single lab, two orthogonal methods, but PF3D7_0704600 is one of two identified targets and further mechanistic detail is not provided in the abstract\",\n      \"pmids\": [\"32669539\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2017,\n      \"finding\": \"The P. yoelii ortholog of PF3D7_0704600 (Pyheul, a HECT-like E3 ubiquitin ligase) was shown to influence parasitemia and host mortality; allelic exchange of partial Pyheul sequences between two lethal parasite strains and modification of its expression altered parasite growth and host mortality, placing it in a virulence-regulatory pathway.\",\n      \"method\": \"Genetic cross of P. yoelii strains, genome-wide QTL scan, allelic exchange, gene expression modification with phenotypic readout (parasitemia, host mortality)\",\n      \"journal\": \"Nature communications\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 / Moderate — genetic cross, QTL mapping, and allelic exchange in P. yoelii; multiple orthogonal methods, single lab\",\n      \"pmids\": [\"28790316\"],\n      \"is_preprint\": false,\n      \"attribution\": [{\"species\": \"Plasmodium yoelii\", \"strain\": \"\"}]\n    }\n  ],\n  \"current_model\": \"PF3D7_0704600 (UT/PfUT) is a HECT-type E3 ubiquitin ligase in P. falciparum that controls intraerythrocytic cell cycle progression (S/M phase duration) and merozoite invasion efficiency when overexpressed, functions as a downstream substrate/target of PfPP1 in the egress pathway, and contributes in a genetic-background-dependent manner to quinine/quinidine sensitivity; the P. yoelii ortholog similarly regulates parasite growth and virulence through allele-dependent mechanisms.\"\n}\n```","stage2_raw":"```json\n{\n  \"mechanistic_narrative\": \"PF3D7_0704600 (UT/PfUT) is a HECT-type E3 ubiquitin ligase of Plasmodium falciparum that regulates intraerythrocytic cell cycle progression and merozoite invasion and functions within the parasite egress pathway [#1, #2]. Overexpression of PfUT lengthens the S/M phase of the intraerythrocytic developmental cycle and reduces merozoite invasion efficiency, establishing it as a positive controller of cell cycle timing and invasion [#1]. Phosphoproteomic and chemical-genetic analyses place PfUT downstream of the protein phosphatase PfPP1 in the pathway governing parasite egress from host erythrocytes, indicating its activity is phosphoregulated [#2]. Genetically, PfUT contributes in a background-dependent manner to quinine and quinidine sensitivity, with episomal expression producing moderate (~2-fold) shifts in IC50 in a subset of strains [#0]. The P. yoelii ortholog (Pyheul) similarly acts in a virulence-regulatory pathway, where allele-dependent differences and altered expression modulate parasitemia and host mortality [#3]. The specific ubiquitination substrates of PfUT, its structural basis, and the mechanistic link between its ligase activity and each phenotype have not been characterized in the available corpus.\",\n  \"teleology\": [\n    {\n      \"year\": 2014,\n      \"claim\": \"Establishing whether PF3D7_0704600 contributes to antimalarial drug response addressed whether this HECT ligase has any measurable phenotypic role in P. falciparum.\",\n      \"evidence\": \"QTL analysis of a genetic cross plus episomal transfection with IC50 measurement across multiple genetic backgrounds\",\n      \"pmids\": [\"24830312\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Effect is moderate (~2-fold) and only seen in a subset of genetic backgrounds\",\n        \"The molecular basis linking ubiquitin ligase activity to quinine/quinidine sensitivity is unknown\",\n        \"No substrate or interacting partner identified for this drug phenotype\"\n      ],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2017,\n      \"claim\": \"Analysis of the P. yoelii ortholog Pyheul tested whether this HECT ligase influences parasite growth and virulence in an in vivo rodent malaria model.\",\n      \"evidence\": \"Genetic cross of P. yoelii strains, genome-wide QTL scan, and allelic exchange with parasitemia and host-mortality readouts\",\n      \"pmids\": [\"28790316\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Findings are in the P. yoelii ortholog, not PF3D7_0704600 itself\",\n        \"The allele-specific molecular differences driving virulence are not defined\",\n        \"No ubiquitination substrate linked to the virulence phenotype\"\n      ],\n      \"attribution\": [{\"species\": \"Plasmodium yoelii\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2019,\n      \"claim\": \"Defining the cellular consequence of altered PfUT levels revealed a direct role in intraerythrocytic cell cycle timing and merozoite invasion, moving beyond drug-response association.\",\n      \"evidence\": \"glmS ribozyme-mediated overexpression (2.5-fold), cell cycle staging, invasion efficiency assays, and partial glucosamine rescue in P. falciparum\",\n      \"pmids\": [\"31797898\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Phenotype derives from overexpression rather than loss-of-function\",\n        \"The substrates ubiquitinated by PfUT to control S/M phase are unidentified\",\n        \"Mechanistic link between invasion defect and cell cycle lengthening not resolved\"\n      ],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": 2020,\n      \"claim\": \"Positioning PfUT downstream of PfPP1 established that its activity is embedded in the phosphoregulated egress signaling pathway.\",\n      \"evidence\": \"Phosphoproteomic and chemical-genetic analysis of PfPP1 in P. falciparum\",\n      \"pmids\": [\"32669539\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"PfUT is one of two identified PfPP1 targets; direct regulation not fully dissected in the abstract\",\n        \"Whether PfPP1 dephosphorylates PfUT directly and how this modulates ligase activity is unknown\",\n        \"The egress-relevant substrates of PfUT are not identified\"\n      ],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    },\n    {\n      \"year\": null,\n      \"claim\": \"The ubiquitination substrates of PfUT, its catalytic and structural basis, and how its HECT ligase activity mechanistically drives cell cycle timing, egress, invasion, and drug response remain unresolved.\",\n      \"evidence\": \"\",\n      \"pmids\": [],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"No direct ubiquitination substrate identified\",\n        \"No structural or biochemical characterization of ligase activity\",\n        \"Loss-of-function phenotype in P. falciparum not defined\"\n      ],\n      \"attribution\": [{\"species\": \"Plasmodium falciparum\", \"strain\": \"\"}]\n    }\n  ],\n  \"mechanism_profile\": {\n    \"molecular_activity\": [\n      {\"term_id\": \"GO:0061630\", \"supporting_discovery_ids\": [0, 1, 2, 3]}\n    ],\n    \"localization\": [],\n    \"pathway\": [\n      {\"term_id\": \"GO:0006511\", \"supporting_discovery_ids\": [0, 1, 2, 3]},\n      {\"term_id\": \"GO:0051301\", \"supporting_discovery_ids\": [1]}\n    ],\n    \"complexes\": [],\n    \"partners\": [\n      \"PfPP1 (functional upstream regulator in the egress pathway) [#2]\"\n    ],\n    \"other_free_text\": [\n      \"Overexpression lengthens the S/M phase of the intraerythrocytic developmental cycle and reduces merozoite invasion efficiency [#1]\",\n      \"Contributes in a genetic-background-dependent manner to quinine and quinidine sensitivity (~2-fold IC50 shifts) [#0]\",\n      \"P. yoelii ortholog Pyheul modulates parasitemia and host mortality in an allele-dependent manner [#3]\",\n      \"No direct ubiquitination substrate, structural model, or loss-of-function phenotype characterized in the available corpus\"\n    ]\n  }\n}\n```","audit_flag":null,"strains":[{"strain":"Pf3D7","gene_id":"PF3D7_0704600","name":null,"product":"HECT-type E3 ubiquitin ligase UT","uniprot_acc":null,"chromosome":null,"start_pos":null},{"strain":"PbANKA","gene_id":"PBANKA_0802300","name":null,"product":"HECT-type E3 ubiquitin ligase UT, putative","uniprot_acc":null,"chromosome":null,"start_pos":null},{"strain":"PkH","gene_id":"PKNH_0103100","name":null,"product":"HECT-type E3 ubiquitin ligase UT, putative","uniprot_acc":null,"chromosome":null,"start_pos":null}],"gene_symbol":"UT","evaluation":null}