{
"METABOLOMICS WORKBENCH":{"STUDY_ID":"ST001279","ANALYSIS_ID":"AN002120","VERSION":"1","CREATED_ON":"November 18, 2019, 3:27 pm"},

"PROJECT":{"PROJECT_TITLE":"K13 mutations driving artemisinin resistance rewrite Plasmodium falciparum’s programmed intra-erythrocytic development and transform mitochondrial physiology","PROJECT_SUMMARY":"The emergence of artemisinin resistance in Southeast Asia, dictated by mutations in the Plasmodium falciparum k13 gene, has compromised antimalarial efficacy and created a core vulnerability in the global malaria elimination campaign. Applying quantitative transcriptomics, proteomics, and metabolomics to a panel of isogenic K13 mutant or wild-type P. falciparum lines, we observe that K13 mutations reprogram multiple aspects of intra-erythrocytic parasite biology. These changes impact its cell cycle periodicity, the unfolded protein response and protein degradation, vesicular trafficking and endocytosis, and mitochondrial functions including the TCA cycle, the electron transport chain, and redox regulation. Ring-stage artemisinin resistance mediated by the K13 R539T mutation was neutralized using atovaquone, an electron transport chain inhibitor. Our data suggest that modification of mitochondrial physiology, accompanied by other processes to reduce artemisinin’s proteotoxic effects, help protect parasites against this pro-oxidant drug, allowing resumption of growth once the rapidly-cleared artemisinins have reached sub-therapeutic levels.","INSTITUTE":"Pennsylvania State University","LAST_NAME":"Llinas","FIRST_NAME":"Manuel","ADDRESS":"W126 Millennium Science Complex, University Park, PENNSYLVANIA, 16802, USA","EMAIL":"mul27@psu.edu","PHONE":"(814) 867-3527"},

"STUDY":{"STUDY_TITLE":"K13 mutations driving artemisinin resistance rewrite Plasmodium falciparum’s programmed intra-erythrocytic development and transform mitochondrial physiology","STUDY_SUMMARY":"The emergence of artemisinin resistance in Southeast Asia, dictated by mutations in the Plasmodium falciparum k13 gene, has compromised antimalarial efficacy and created a core vulnerability in the global malaria elimination campaign. Applying quantitative transcriptomics, proteomics, and metabolomics to a panel of isogenic K13 mutant or wild-type P. falciparum lines, we observe that K13 mutations reprogram multiple aspects of intra-erythrocytic parasite biology. These changes impact its cell cycle periodicity, the unfolded protein response and protein degradation, vesicular trafficking and endocytosis, and mitochondrial functions including the TCA cycle, the electron transport chain, and redox regulation. Ring-stage artemisinin resistance mediated by the K13 R539T mutation was neutralized using atovaquone, an electron transport chain inhibitor. Our data suggest that modification of mitochondrial physiology, accompanied by other processes to reduce artemisinin’s proteotoxic effects, help protect parasites against this pro-oxidant drug, allowing resumption of growth once the rapidly-cleared artemisinins have reached sub-therapeutic levels.","INSTITUTE":"Pennsylvania State University","LAST_NAME":"Llinas","FIRST_NAME":"Manuel","ADDRESS":"W126 Millennium Science Complex, University Park, PENNSYLVANIA, 16802, USA","EMAIL":"mul27@psu.edu","PHONE":"(814) 867-3527"},

"SUBJECT":{"SUBJECT_TYPE":"Cultured cells","SUBJECT_SPECIES":"Plasmodium falciparum","TAXONOMY_ID":"5833"},
"SUBJECT_SAMPLE_FACTORS":[
{
"Subject ID":"-",
"Sample ID":"MTZH4DM.1_t3",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH4DM.2_t3",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH4DM.3_t3",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH470.1_t3",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH470.2_t3",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH470.3_t3",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH4350.1_t3",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH4350.2_t3",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"MTZH4350.3_t3",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2ND.1_t4",
"Factors":{"Treatment":"None"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2ND.2_t4",
"Factors":{"Treatment":"None"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2ND.3_t4",
"Factors":{"Treatment":"None"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2DM.1_t4",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2DM.2_t4",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2DM.3_t4",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC270.1_t4",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC270.2_t4",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC270.3_t4",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2350.1_t4",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2350.2_t4",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"MTZC2350.3_t4",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"H4ND_t6",
"Factors":{"Treatment":"None"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM_t6",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350_t6",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"C2ND_t6",
"Factors":{"Treatment":"None"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM_t6",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350_t6",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.1_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.2_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.3_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.1_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.2_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.3_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.1_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.2_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.3_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.1_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.2_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.3_t7",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.1_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.2_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.3_t7",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.1_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.2_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.3_t7",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.1_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.2_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4DM.3_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.1_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.2_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H470.3_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.1_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.2_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"H4350.3_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIC580Y"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.1_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.2_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2DM.3_t8",
"Factors":{"Treatment":"DMSO"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.1_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.2_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C270.3_t8",
"Factors":{"Treatment":"70nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.1_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.2_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
},
{
"Subject ID":"-",
"Sample ID":"C2350.3_t8",
"Factors":{"Treatment":"350nM DHA"},
"Additional sample data":{"Genotype":"Cam3.IIWT"}
}
],
"COLLECTION":{"COLLECTION_SUMMARY":"P. falciparum parasites were cultured at 3% hematocrit in human O+ RBCs (Interstate blood bank, USA) and P. falciparum culture media comprising of RPMI1640 (Thermo Fisher Scientific) supplemented with 0.5% (w/v) Albumax II, 50mg/L hypoxanthine, 0.2% NaHCO3, 25mM HEPES and 10mg/L gentamycin (Fidock et al., 1998). Parasites were cultured at 37ºC in 5% O2, 5%CO2 and 90% N2. For the collection of RNA, proteins and metabolite extracts, parasite cultures at 3% hematocrit and 20 mL or 200 mL volumes were kept in T75 or T225 flasks respectively, with daily media changes. Parasite lines were genotyped by Sanger sequencing for the k13 gene to verify their identities before the start of an experiment.","SAMPLE_TYPE":"Parasite"},

"TREATMENT":{"TREATMENT_SUMMARY":"Mycoplasma-free Cam3.IIC580Y and Cam3.IIWT parasites were doubly synchronized by 5% D-Sorbitol in each generation for at least two generations. 0-3 hpi early rings of each parasite line were treated for 3h at 350 nM or 70 nM DHA along with vehicle-treated 0.05% DMSO controls in two to three independent experiments. 24 hpi trophozoites were similarly treated with DHA or DMSO control in a single experiment for each parasite line and subsequently magnetically enriched using MACS CS columns on the SuperMACS™ II Separator (Miltenyi Biotec, Inc.) to remove uninfected RBCs."},

"SAMPLEPREP":{"SAMPLEPREP_SUMMARY":"The K13 mutant and WT ring-stage parasites were run in parallel for each metabolomic experiment to allow direct comparisons of their metabolomic states and the effects of DHA. Metabolites from saponin-lysed parasites were extracted with cold methanol along with spike-in control [13C4, 15N1]-Aspartate (Cambridge Isotope) as an internal LC/MS standard to correct for technical variation arising from sample processing in the data analysis phase, as described previously (Allman et al., 2016).Samples were injected on a Thermo Exactive Plus Orbitrap in negative ion mode (Lu, W. et al., Anal. Chem. 2010.)."},

"CHROMATOGRAPHY":{"CHROMATOGRAPHY_TYPE":"Reversed phase","INSTRUMENT_NAME":"Thermo Dionex Ultimate 3000","COLUMN_NAME":"Phenomenex Synergi Hydro RP 100 A (100 x 2mm, 2.5um)"},

"ANALYSIS":{"ANALYSIS_TYPE":"MS"},

"MS":{"INSTRUMENT_NAME":"Thermo Exactive Plus Orbitrap","INSTRUMENT_TYPE":"Orbitrap","MS_TYPE":"ESI","ION_MODE":"NEGATIVE","MS_COMMENTS":"Raw data files from the Thermo Exactive Plus orbitrap (.raw) were converted to a format compatible with our analysis software (.raw¡.mzXML) and spectral data (.mzXML files) were visualized in MAVEN. The labeled [13C4, 15N1]-Aspartate internal standard intensity was assessed for technical reproducibility. Peaks for each metabolite in the targeted library were identified based on proximity to standard retention time, the observed mass falling within 10 ppm of the expected m/z (calculated from the monoisotopic mass), and the signal/blank ratio (minimum, 10,000 ions). Based upon the above criteria, peaks were manually inspected and demarcated as good or bad based on peak shape. Peak areas were exported into an R working environment (http://www.R-project.org) to calculate log2 fold changes for each sample compared to an untreated control. Metabolites that were not reliably detected across 90% of all the trials were removed prior to additional analysis to minimize subsequent imputation bias. The peak areas for any remaining metabolites not detected were imputed to have 10,000 ions, and metabolites detected below background levels (negative after blank subtraction) were maintained as “0” prior to averaging and log2 calculation. Because our extraction method did not include a wash step we excluded metabolites found in the RPMI-based medium."},

"MS_METABOLITE_DATA":{
"Units":"Peak Area",

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}

}