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Saeij Lab Publications

Peer-reviewed Publications

($Denotes shared first author. *Denotes graduate students from my lab. #Denotes Co-corresponding authors)

From Saeij lab




  • Sarcocystis calchasi and other Sarcocystidae detected in predatory birds in California, USA. Rogers KH, Arranz-Solís D, Saeij JPJ, Lewis S, Mete A. Int J Parasitol Parasites Wildl. 2021 ;17:91-99. doi: 10.1016/j.ijppaw.2021.12.008.
  • The Toxoplasma Polymorphic Effector GRA15 Mediates Seizure Induction by Modulating Interleukin-1 Signaling in the Brain. Glausen TG, Carrillo GL, Jin RM, Boyle JP, Saeij JPJ, Wohlfert EA, Fox MA, Blader IJ. mBio. 2021 ;12(3):e0133121.
  • CRISPR screen to determine the in vivo fitness of Toxoplasma genes. Sangaré LO, Wang Y, Arranz-Solís D, Saeij JPJ. STAR Protoc. 2021 ;2(2):100520.
  • Toxoplasma gondii Matrix Antigen 1 Is a Secreted Immunomodulatory Effector. Tomita T, Mukhopadhyay D, Han B, Yakubu R, Tu V, Mayoral J, Sugi T, Ma Y, Saeij JPJ, Weiss LM. mBio. 2021 12(3):e00603-21.
  • Toxoplasma GRA Peptide-Specific Serologic Fingerprints Discriminate Among Major Strains Causing Toxoplasmosis. Arranz-Solís D, Carvalheiro CG, Zhang ER, Grigg ME, Saeij JPJ. Front Cell Infect Microbiol. 2021 ;11:621738.
  • Toxoplasma Effectors that Affect Pregnancy Outcome. Arranz-Solís D, Mukhopadhyay D, Saeij JPJ. Trends Parasitol. 2021 ;37(4):283-295.


  • Influence of the Host and Parasite Strain on the Immune Response During Toxoplasma Infection. Mukhopadhyay D, Arranz-Solís D, Saeij JPJ. Front Cell Infect Microbiol. 2020 ;10:580425.
  • Genome-wide screens identify Toxoplasma gondii determinants of parasite fitness in IFNγ-activated murine macrophages. Wang Y, Sangaré LO, Paredes-Santos TC, Hassan MA, Krishnamurthy S, Furuta AM, Markus BM, Lourido S, Saeij JPJ. Nat Commun. 2020 ;11(1):5258.
  • Naïve CD8 T cell IFNγ responses to a vacuolar antigen are regulated by an inflammasome-independent NLRP3 pathway and Toxoplasma gondii ROP5. Kongsomboonvech AK, Rodriguez F, Diep AL, Justice BM, Castallanos BE, Camejo A, Mukhopadhyay D, Taylor GA, Yamamoto M, Saeij JPJ, Reese ML, Jensen KDC. PLoS Pathog. 2020 ;16(8):e1008327.
  • Toxoplasma Mechanisms for Delivery of Proteins and Uptake of Nutrients Across the Host-Pathogen Interface. Wang Y, Sangaré LO, Paredes-Santos TC, Saeij JPJ. Annu Rev Microbiol. 2020 ;74:567-586.
  • Protozoal encephalitis associated with Sarcocystis calchasi and S. falcatula during an epizootic involving Brandt’s cormorants (Phalacrocorax penicillatus) in coastal Southern California, USA. Bamac OE, Rogers KH, Arranz-Solís D, Saeij JPJ, Lewis S, Duerr R, Skoglund J, Peronne L, Mete A. Int J Parasitol Parasites Wildl. 2020 ;12:185-191.
  • Toxoplasma  GRA15 and GRA24 are important activators of the host innate immune response in the absence of TLR11. Mukhopadhyay D, Arranz-Solís D, Saeij JPJ. PLoS Pathog. 2020 ;16(5):e1008586.
  • Comparative tachyzoite proteome analyses among six Neospora caninum isolates with different virulence. Rico-San Román L, Horcajo P, Regidor-Cerrillo J, Fernández-Escobar M, Collantes-Fernández E, Gutiérrez-Blázquez D, Hernáez-Sánchez ML, Saeij JPJ, Ortega-Mora LM. Int J Parasitol. 2020 ;50(5):377-388.
  • Toxoplasma GRA15 limits parasite growth in IFNγ-activated fibroblasts through TRAF ubiquitin ligases. Mukhopadhyay D, Sangaré LO, Braun L, Hakimi MA, Saeij JP. EMBO J. 2020 ;39(10):e103758.
  • Identification of a Master Regulator of Differentiation in Toxoplasma. Waldman BS, Schwarz D, Wadsworth MH 2nd, Saeij JP, Shalek AK, Lourido S. Cell. 2020 ;180(2):359-372.e16.
  • Serotyping of Toxoplasma gondii Infection Using Peptide Membrane Arrays. Arranz-Solís D, Cordeiro C, Young LH, Dardé ML, Commodaro AG, Grigg ME, Saeij JPJ. Front Cell Infect Microbiol. 2019 ;9:408.
  • Assays to Evaluate Toxoplasma-Macrophage Interactions. Mukhopadhyay D, Saeij JPJ. Methods Mol Biol. 2020;2071:347-370.


  • Sangaré LO, Ólafsson EB, Wang Y, Yang N, Julien L, Camejo A, Pesavento P, Sidik SM, Lourido S, Barragan A, Saeij JPJ. In Vivo CRISPR Screen Identifies TgWIP as a Toxoplasma Modulator of Dendritic Cell Migration. Cell Host Microbe. 26(4):478-492.2019.
  • Paredes-Santos T, Wang Y, Waldman B, Lourido S, Saeij JP. Front Cell Infect Microbiol. The GRA17 Parasitophorous Vacuole Membrane Permeability Pore Contributes to Bradyzoite Viability. 9:321. 2019.
  • Sangaré LO, Yang N, Konstantinou EK, Lu D, Mukhopadhyay D, Young LH, Saeij JPJ. Toxoplasma GRA15 Activates the NF-κB Pathway through Interactions with TNF Receptor-Associated Factors. mBio. 10(4):e00808-19. 2019.
  • Wang Y$, Cirelli KM$*, Barros PDC, Omar Sangaré L, Butty V, Hassan MA, Pesavento P, Mete A, Saeij JP. Three Toxoplasma gondii dense granule proteins are required for induction of Lewis rat macrophage pyroptosis. mBio. 8;10(1); 2019.
  • Hassan MA, Olijnik AA, Frickel EM, Saeij JP. Clonal and atypical Toxoplasma strain differences in virulence vary with mouse sub-species. Int J Parasitol. 49(1):63-70; 2019.


  • Krishnamurthy S, Saeij JP. Toxoplasma Does Not Secrete the GRA16 and GRA24 Effectors Beyond the Parasitophorous Vacuole Membrane of Tissue Cysts. Front Cell Infect Microbiol. 18;8:366; 2018.
  • Arranz-Solís D, Regidor-Cerrillo J, Lourido S, Ortega-Mora LM, Saeij JP. Toxoplasma CRISPR/Cas9 constructs are functional for gene disruption in Neospora caninum. Int J Parasitol. 48(8):597-600; 2018.


  • Saeij JP, Frickel EM. Exposing Toxoplasma gondii hiding inside the vacuole: a role for GBPs, autophagy and host cell death. Curr Opin Microbiol. 40:72-80; 2017. [Review Article]
  • Krishnamurthy S, Konstantinou EK, Young LH, Gold DA, Saeij JP. The human immune response to Toxoplasma: Autophagy versus cell death. PLoS Pathog. 9;13(3):e1006176; 2017. [Review Article]


  • Sidik SM, Huet D, Ganesan SM, Huynh MH, Wang T, Nasamu AS, Thiru P, Saeij JP, Carruthers VB, Niles JC, Lourido S. A Genome-wide CRISPR Screen in Toxoplasma Identifies Essential Apicomplexan Genes. Cell. S0092-8674(16)31070-4; 2016.
  • Fink A, Hassan MA, Okan NA, Sheffer M, Camejo A, Saeij JP, Kasper DL. Early Interactions of Murine Macrophages with Francisella tularensis Map to Mouse Chromosome 19. mBio. 15;7(2):e02243; 2016.
  • Lorenzi H, Khan A, Behnke MS, Namasivayam S, Swapna LS, Hadjithomas M, Karamycheva S, Pinney D, Brunk BP, Ajioka JW, Ajzenberg D, Boothroyd JC, Boyle JP, Dardé ML, Diaz-Miranda MA, Dubey JP, Fritz HM, Gennari SM, Gregory BD, Kim K, Saeij JP, Su C, White MW, Zhu XQ, Howe DK, Rosenthal BM, Grigg ME, Parkinson J, Liu L, Kissinger JC, Roos DS, Sibley LD. Local admixture of amplified and diversified secreted pathogenesis determinants shapes mosaic Toxoplasma gondii genomes. Nature Commun. 7:10147; 2016.


  • Hassan MA, Jensen KD, Butty V, Hu K, Boedec E, Prins P, Saeij JP. Transcriptional and Linkage Analyses Identify Loci that Mediate the Differential Macrophage Response to Inflammatory Stimuli and Infection. PLoS Genet. 11(10):e1005619; 2015.
  • Gold DA, Kaplan AD, Lis A, Bett GC, Rosowski EE*, Cirelli KM*, Bougdour A, Sidik SM, Beck JR, Lourido S, Egea PF, Bradley PJ, Hakimi MA, Rasmusson RL, Saeij JP. The Toxoplasma dense granule proteins GRA17 and GRA23 mediate the movement of small molecules between the host and the parasitophorous vacuole. Cell Host & Microbe. 17(5):642-52; 2015. Research Highlights: Nature Reviews Microbiology 13, 398 (2015) and Selected by Faculty of 1000.
  • Jensen KD$, Camejo A$, Melo MB, Cordeiro C, Grotenbreg GM, Frickel E, Ploegh HL, Young L, Saeij JP. Toxoplasma superinfection and virulence during secondary infection correlates with the exact ROP5/ROP18 allelic combination. mBio. 24;6(2). pii: e02280-14; 2015.


  • Hassan MA, Saeij JP. Incorporating alternative splicing and mRNA editing into the genetic analysis of complex traits. Bioessays. 36(11):1032-40; 2014. [Review article]
  • Murray PJ, Allen JE, Biswas SK, Fisher EA, Gilroy DW, Goerdt S, Gordon S, Hamilton JA, Ivashkiv LB, Lawrence T, Locati M, Mantovani A, Martinez FO, Mege JL, Mosser DM, Natoli G, Saeij JP, Schultze JL, Shirey KA, Sica A, Suttles J, Udalova I, van Ginderachter JA, Vogel SN, Wynn TA. Macrophage Activation and Polarization: Nomenclature and Experimental Guidelines. Immunity. 41(1):14-20; 2014.
  • Cirelli KM$*, Gorfu G, Hassan MA, Printz M, Crown D, Breen L, Leppla SH, Grigg ME#, Saeij JP#, Moayeri M#. Inflammasome sensor NLRP1 controls rat macrophage susceptibility to Toxoplasma gondii. PLoS Pathogens. 10(3):e1003927; 2014.
  • Cannella D, Brenier-Pinchart MP, Braun L, van Rooyen JM, Bougdour A, Bastien O, Behnke MS, Curt RL, Curt A, Saeij JP, Sibley LD, Pelloux H, Hakimi MA. miR-146a and miR-155 delineate a microRNA fingerprint associated with Toxoplasma persistence in the host brain. Cell Rep. pii: S2211-1247(14)00081-3; 2014.
  • Gorfu G$, Cirelli KM$*, Melo MB, Mayer-Barber K, Crown D, Koller BH, Masters S, Sher A, Leppla SH, Moayeri M#, Saeij JP#, Grigg ME#. A dual role for inflammasome sensors NLRP1 and NLRP3 in murine resistance to Toxoplasma gondii. mBio. 5(1); pii: e01117-13; 2014.
  • Hassan MA, Butty V, Jensen KD, Saeij JP. The genetic basis for individual differences in mRNA splicing and Apobec1 editing activity in murine macrophages. Genome Research. 24(3):377-89; 2014.
  • Rosowski EE*, Nguyen QP, Camejo A, Spooner E, Saeij JP. Toxoplasma gondii inhibits gamma interferon (IFN-γ)- and IFN-β-induced host cell STAT1 transcriptional activity by increasing the association of STAT1 with DNA. Infection and Immunity. 82(2): 706-19; 2014.
  • Camejo A, Gold DA, Lu D*, McFetridge K, Julien L, Yang N*, Jensen KD, Saeij JP. Identification of three novel Toxoplasma gondii rhoptry proteins. International Journal for Parasitology. 44(2): 147-60; 2014.


  • Melo MB, Nguyen QP, Cordeiro C, Hassan MA, Yang N*, McKell R*, Rosowski EE*, Julien L, Butty V, Darde M-L, Ajzenberg D, Fitzgerald K. Young LH, Saeij JP. Transcriptional analysis of murine macrophages infected with different Toxoplasma strains identifies novel regulation of host signaling pathways. PLoS Pathogens. 9(12): e1003779; 2013.
  • Lim D, Gold DA, Julien L, Rosowski EE*, Niedelman W*, Yaffe MB#, Saeij JP#. Structure of the Toxoplasma gondii ROP18 kinase domain reveals a second ligand binding pocket required for acute virulence. The Journal of Biological Chemistry. 288(48): 34968-80; 2013.
  • Niedelman W*, Sprokholt JK, Clough B, Frickel EM, Saeij JP. Cell death of interferon-gamma stimulated human fibroblasts upon Toxoplasma gondii infection induces early parasite egress and limits parasite replication. Infection and Immunity. 81(12): 4341-9; 2013.


  • Hassan MA, Melo MB, Haas B, Jensen KD, Saeij JP. De novo reconstruction of the Toxoplasma gondii transcriptome improves on the current genome annotation and reveals alternatively spliced transcripts and putative long non-coding RNAs. BMC Genomics. 13(1):696; 2012.
  • Minot S$, Melo MB$, Li F, Lu D*, Niedelman W*, Levine SS, Saeij JP. Admixture and recombination among Toxoplasma gondii lineages explains global genome diversity. PNAS. 14;109(33):13458-63; 2012.
  • Niedelman W*, Gold DA, Rosowski EE*, Sprokholt J, Lim D, Farid A, Melo MB, Spooner E, Yaffe MB, Saeij JP. The rhoptry proteins ROP18 and ROP5 mediate Toxoplasma gondii evasion of the murine, but not the human, interferon-gamma response. PLoS Pathogens. 8(6):e1002784; 2012.
  • Szatanek T, Anderson-White BR, Faugno-Fusci DM, White M, Saeij JP, Gubbels MJ. Cactin is essential for G1 progression in Toxoplasma gondii. Molecular Microbiology. 84(3):566-77; 2012.
  • Lim DC, Cooke BM, Doerig C, Saeij JP. Toxoplasma and Plasmodium protein kinases: roles in invasion and host cell remodelling. International Journal for Parasitology. 42(1):21-32; [Review article]


  • Virreira-Winter S, Niedelman W*, Jensen KD, Rosowski EE*, Julien L, Spooner E, Caradonna K, Burleigh BA, Saeij JP, Ploegh HL, Frickel E. Determinants of GBP recruitment to Toxoplasma gondii vacuoles and the parasitic factors that control it. PLoS One. 6(9):e24434; 2011.
  • Reese ML, Zeiner GM, Saeij JP, Boothroyd JC, Boyle JP. Polymorphic family of injected pseudokinases is paramount in Toxoplasma. PNAS 108(23):9625-30; 2011. Selected by Faculty of 1000.
  • Jensen KD, Wang Y, Tait ED, Shastri AJ, Hu K, Cornel L, Boedec E, Ong Y, Chien Y, Hunter CA, Boothroyd JC, Saeij JP. Toxoplasma polymorphic effectors determine macrophage polarization and intestinal inflammation. Cell Host & Microbe 9(6):472-83; 2011. Highlighted in: Macrophages as a battleground for Toxoplasma pathogenesis. Cell Host & Microbe 9(6):445-7; 2011.
  • Melo MB$, Jensen KD$, Saeij JP. Toxoplasma gondii effectors are master regulators of the inflammatory response. Trends in Parasitology. 27(11):487-95; 2011. [Review article]
  • Rosowski EE$*, Lu D$*, Julien L, Rodda L, Gaiser R, Jensen KD, Saeij JP. Strain-specific activation of the NF-κB pathway by GRA15, a novel Toxoplasma dense granule protein. Journal of Experimental Medicine 208(1):195-212; 2011. Selected by Faculty of 1000. Highlighted in: It takes II to induce NF-κ Nature Reviews Microbiology 9, 147, 2011.


  • Blader IJ, Saeij JPJ. Communication between Toxoplasma gondii and its host: impact on parasite growth, development, immune evasion, and virul APMIS (acta pathologica, microbiologica, et immunologica Scandinavica). 117(5-6):458-76; 2009. [Review article]


From Postdoctoral work

  • Boyle JP, Saeij JP, Harada SY, Ajioka JW, Boothroyd JC. Expression quantitative trait locus mapping of Toxoplasma genes reveals multiple mechanisms for strain-specific differences in gene expression. Eukaryotic Cell. 7(8):1403-14; 2008.
  • Saeij JP, Arrizabalaga G, Boothroyd JC. A cluster of four surface antigen genes specifically expressed in bradyzoites, SAG2CDXY, plays an important role in Toxoplasma gondii. Infection and Immunity 76(6): 2402-2410; 2008.
  • Boyle JP, Saeij JP, Boothroyd JC. Inconsistent dissemination patterns of Toxoplasma gondii following oral infection. Experimental Parasitology 116(3):302-305; 2007.
  • Saeij JP$, Coller S$, Boyle JP, Jerome ME, White MW, Boothroyd JC. Toxoplasma co-opts host gene expression by injection of a polymorphic kinase homologue. Nature 445(7125): 324-327; 2007. Selected by Faculty of 1000. Highlighted in: Toxo researchers spot the difference. Nature Reviews Microbiology 5, 86, 2007.
  • Saeij JP$, Boyle JP$, Coller SC, Taylor S, Sibley LD, Brooke-Powell ET, Ajioka JW, Boothroyd JC. Polymorphic secreted kinases are key virulence factors in toxoplasmosis. Science 314(5806):1780-1783; 2006. Selected by Faculty of 1000. Highlighted in: Toxo researchers spot the difference. Nature Reviews Microbiology 5, 86, 2007; Nature Immunology 8, 129, 2007.
  • Boyle, JP, Rajasekar B, Saeij JP, Ajioka JW, Berriman M, Paulsen I, Roos DS, Sibley D, White M, Boothroyd JC. Just one cross appears capable of dramatically altering the population biology of a eukaryotic pathogen like Toxoplasma gondii. PNAS 103(27): 10514-10519; 2006. Selected by Faculty of 1000.
  • Boyle JP, Saeij JP, Cleary MD, Boothroyd JC. Analysis of gene expression during development: lessons from the Apicomplexa. Microbes and Infection 8(6): 1623-1630; 2006. [Review article]
  • Saeij JP, Boyle JP, Boothroyd JC. Differences among the three major strains of Toxoplasma gondii and their specific interactions with the infected host. Trends in Parasitology. 21(10): 476-81; 2005.
  • Saeij JP$, Boyle JP$, Grigg ME, Arrizabalaga G, Boothroyd JC. Bioluminescence imaging of Toxoplasma gondii infection in living mice reveals dramatic differences between strains. Infection and Immunity. 73(2): 695-702; 2005. Selected by Faculty of 1000.

From graduate work

  • Huttenhuis HBT, Taverne-Thiele AJ, Grou CPO, Bergsma J, Saeij JP, Nakayasu C, Rombout JHWM. Ontogeny of the common carp (Cyprinus carpio) innate immune system. Developmental & Comparative Immunology. 30(6): 557-574; 2006.
  • Joerink M, Saeij JP, Stafford JL, Belosevic M, Wiegertjes GF. Animal models for the study of innate immunity: protozoan infections in fish. Symposia of the Society for Experimental Biology. 55:67-89; 2004.
  • Saeij JP, Groeneveld A, Van Rooijen N, Haenen OLM, Wiegertjes GF. Minor effect of depletion of resident macrophages from peritoneal cavity on resistance of common carp Cyprinus carpio to blood flagellates. Diseases of aquatic organisms 57: 67-75; 2003.
  • Engelsma MY, Stet RJM, Saeij JP, Verburg-van Kemenade BML. Differential expression and haplotypic variation of two interleukin-1 beta genes in the common carp (Cyprinus carpio). Cytokine 22: 21-32; 2003.
  • Saeij JP, Stet RJM, de Vries B, van Muiswinkel WB, Wiegertjes GF. Molecular and functional characterization of carp TNF: a link between TNF polymorphism and trypanotolerance? Developmental & Comparative Immunology. 27: 29-41; 2003.
  • Saeij JP, van Muiswinkel WB, van de Meent M, Amaral C, Wiegertjes GF. Different capacities of carp leukocytes to encounter nitric oxide-mediated stress: a role for the intracellular reduced glutathione pool. Developmental & Comparative Immunology. 27: 555-568; 2003.
  • Saeij JP, Verburg-van Kemenade BML, van Muiswinkel WB, Wiegertjes GF. Daily handling reduces resistance of carp to Trypanoplasma borreli: in vitro modulatory effects of cortisol on leukocyte function and apoptosis. Developmental & Comparative Immunology. 27: 233-245; 2003.
  • Saeij JP, de Vries BJ, Wiegertjes GF. The immune response of carp to Trypanoplasma borreli: kinetics of immune gene expression and polyclonal lymphocyte activation. Developmental & Comparative Immunology. 27: 859-874; 2003.
  • Saeij JP, Stet RJM, Wiegertjes GF. Immune modulation by fish kinetoplastid parasites: a role for nitric oxide. Parasitology 124: 77-86; 2002.
  • Saeij JP, Stet RJM, Groeneveld A, Verburg-van Kemenade BML, van Muiswinkel WB, Wiegertjes GF. Molecular and functional characterization of a fish inducible-type nitric oxide synthase. Immunogenetics 51: 339-346; 2000.
  • Saeij JP, Wiegertjes GF, Stet RJM. Identification and characterization of a fish natural resistance-associated macrophage protein (NRAMP) cDNA. Immunogenetics 50: 60-66; 1999.
  • Stet RJM, Kruiswijk CP, Saeij JP, Wiegertjes GF. Major histocompatibility genes in cyprinid fishes: theory and practice. Immunological Reviews. 166: 301-316; 1998. [Review article]
  • Verburg-van Kemenade BML, Saeij JP, Flik G, Willems PHGM. Ca2+ signals during early lymphocyte activation in carp Cyprinus carpio L. The Journal of Experimental Biology 201: 591-598; 1998.