Diane McVey Ward

Diane McVey Ward, PhD

Languages spoken: English

Academic Information

Departments Primary - Pathology

Academic Office Information

diane.mcveyward@path.utah.edu

Lab

Research Statement

Our research focuses on two major areas. First, we are interested in iron metabolism. Iron is an element required by virtually all organisms. The facile ability of iron to gain and loose electrons renders this metal an essential cofactor in redox reactions. Organisms as disparate as prokaryotes and mammals have developed a variety of mechanisms to obtain iron and regulate its storage and utilization. We utilize yeast as a model system to study iron metabolism because of the relative ease of genetic manipulation. We have utilized a variety of genetic screens to identify yeast genes required for iron transport across the plasma membrane for iron storage in the yeast vacuole. We have also identified molecules that respond to both high and low iron and regulate the transcription of iron transporters. Our studies in yeast have led to the identification of both plant and mammalian iron transporters.

We have also identified the mechanisms underlying both the regulation and malregulation of mammalian iron-linked disorders. We determined that entry of iron from cells into plasma is dependent upon the interaction of the iron exporter ferroportin with the peptide hormone hepcidin. Hepcidin is synthesized by the liver in response to inflammation and iron stores and is a negative regulator of plasma iron. We determined that hepcidin binds to ferroportin inducing its internalization and degradation. Mammalian iron overload disease is the result of either inadequate hepcidin production or mutations in ferroportin that lead to hepcidin resistance. Our current studies have identified the mechanism of hepcidin-mediated ferroportin internalization and degradation.

Our second area of research is the study of membrane trafficking. We are interested in identifying molecules that regulate the fusion and fission of endocytic vesicles. Many human diseases result from alterations vesicle trafficking and delivery to the lysosomes. One such disease is Chediak-Higashi syndrome in which lysosomes are abnormally large. We have identified the gene responsible for this disorder and have ongoing studies to identify the biochemical defect responsible for the enlargement of lysosomes.

Education History

Undergraduate Brigham Young University
 BS
Doctoral Training University of Utah
 PhD
Postdoctoral Fellowship University of Utah
 Postdoctoral Fellow

Selected Publications

Journal Article

  1. Lin H, BurtonD, Li L, Warner DE, Phillips J, Ward DM, and Kaplan J (2009). Gain-of-function mutations identify amino acids within transmembrane domains of theyeast vacuolar transporter Zrc1 that determine metal specificity. Biochemistry, 422, 273-283.
  2. Ward DM, and Kaplan (2012). Ferroportin-mediated iron transport: expression and regulation. Biochimica et biophysica acta, 1823, 1426-1433.
  3. De Domenico I, Ward DM, Musci G, Kaplan (2007). Evidence for the multimeric structure of ferroportin. Blood, 109(5), 2205-9.
  4. Zohn IE, De Domenico I, Pollock A, Ward DM, Goodman JF, Liang X, Sanchez AJ, Niswander L, Kaplan (2007). The flatiron mutation in mouse ferroportin acts as a dominant negative to cause ferroportin disease. Blood, 109(10), 4174-80.
  5. De Domenico I, Ward DM, Kaplan (2009). Specific iron chelators determine the route of ferritin degradation. Blood, 114(20), 4546-51.
  6. Troadec MB, Warner D, Wallace J, Thomas K, Spangrude GJ, Phillips J, Khalimonchuk O, Paw BH, Ward DM, Kaplan (2011). Targeted deletion of the mouse Mitoferrin1 gene: from anemia to protoporphyria. Blood, 117(20), 5494-502.
  7. DeDomenico I, Vaughn MB, Yoon D, Kushner JP, Kaplan J, Ward D (2007). Zebrafish as a model for defining the functional impact of mammalian ferroportin mutations. Blood, 110, 3780-3783.
  8. Koening CL, Miller JC, Nelson JM, Ward DM, Kushner JP, Bockenstedt LK, Weis JJ, Kaplan J, and DeDomenico I (2009). Toll-like receptors mediate induction of hepcidin in mice infected with Borrelia Burgdorferi. Blood, 114, 1913-1918.
  9. Paradkar PN, DeDomenico I, Zohn I, Kaplan J, Ward D (2008). Iron-depletion limits intracellular bacterial growth in macrophages. Blood, 112, 866-874.
  10. Troadec MB, Warner DE, Wallace J, Phillips J, Paw BH, Ward DM, and Kaplan (2011). Selective loss of mitoferrin 1 in mouse tissues reveals organ specific defects. Blood, 117, 5494-5502.
  11. Troadec MB, Ward DM, Lo E, Kaplan J, De Domenico I (2010). Induction of FPN1 transcription by MTF-1 reveals a role for ferroportin in transition metal efflux. Blood, 116(22), 4657-4664.
  12. De Domenico I, Ward DM, Langelier C, Vaughn MB, Nemeth E, Sundquist WI, Ganz T, Musci G, Kaplan (2007). The molecular mechanism of hepcidin-mediated ferroportin down-regulation. Molecular biology of the cell, 18(7), 2569-78.
  13. Runkel F, Bussow H, Seburn KL, Cox GA, Ward DM, Kaplan J, Franz (2006). Grey, a novel mutation in the murine Lyst gene, causes the beige phenotype by skipping of exon 25. Mammalian genome, 17(3), 203-10.
  14. Wang F, Custodio AO, Paradkar P, Ward DM, Fleming MD, Campagna D, Roberts KA, Boyartchuk V, Kaplan J, Dietrich WF, Andrews N (2007). Mon 1 a, a modifer of spleen iron loading, regulates macrophage trafficking of the iron transporter ferroportin. Nature genetics,
  15. Wang F, Paradka P, Custodio AO, Ward DM, Fleming MD, Campagna D, Roberts KA, Boyartchuk V, Dietrich WF, Kaplan J. and Andrews N (2007). Genetic variation in Mon1a affects protein trafficking and modifies macrophage iron loading in mice. Nature genetics, 39, 1025-1032.
  16. De Domenico I, Vaughn MB, Li L, Bagley D, Musci G, Ward DM, Kaplan (2006). Mobilization of ferritin iron does not require ferritin degradation. The EMBO journal, 25, 5396-5404.
  17. De Domenico I, Ward DM, Carmela Bonaaccorsi di Patti M, Musci G, Kaplan (2007). Hepcidin-independent regulation of ferroportin: a role for ceruloplasmin in ferroportin-mediated iron export. The EMBO journal,
  18. De Domenico I, Ward DM, di Patti MC, Jeon SY, David S, Musci G, and Kaplan (2007). Ferroxidase activity is required for the stability of cell surface ferroportin in cells expressing GPI-ceruloplasmin.PMID:17541408. The EMBO journal, 26, 2823-2831.
  19. Mohlig H, Mathieu S, Thon L, Frederiksen MC, Ward DM, Kaplan J, Schutze S, Kabelitz D, Adam (2007). The WD repeat protein FAN regulates lysosome size independent from abnormal downregulation/membrane recruitment of protein kinase C. Experimental cell research,
  20. De Domenico I, Ward DM, Musci G, Kaplan (2006). Iron overload due to mutations in ferroportin. Haematologica, 91(1), 92-5.
  21. DeDomenico J, Ward DM, Nemeth E, Ganz T, Corradini E, Ferrara F, Musci G, Pietrangelo A., Kaplan (2006). Molecular and clinical correlates in iron overload associated with mutations in ferroportin. Hematologic pathology, 91, 1092-1095.
  22. Davis-Kaplan SR, Ward DM, Shiflett SL, Kaplan (2004). Genome-wide analysis of iron-dependent growth reveals a novel yeast gene required for vacuolar acidification. The Journal of biological chemistry, 279(6), 4322-9.
  23. Shiflett SL, Ward DM, Huynh D, Vaughn MB, Simmons JC, Kaplan (2004). Characterization of Vta1p, a class E Vps protein in Saccharomyces cerevisiae. The Journal of biological chemistry, 279(12), 10982-90.
  24. Felice MR, De Domenico I, Li L, Ward DM, Bartok B, Musci G, Kaplan (2005). Post-transcriptional regulation of the yeast high affinity iron transport system. The Journal of biological chemistry, 280(23), 22181-90.
  25. Kumanovics A, Poruk KE, Osborn KA, Ward DM, Kaplan (2006). YKE4 (YIL023C) encodes a bidirectional zinc transporter in the endoplasmic reticulum of Saccharomyces cerevisiae. The Journal of biological chemistry, 281(32), 22566-74.
  26. Davis-Kaplan SR, Compton MA, Flannery AR, Ward DM, Kaplan J, Stevens TH, Graham L (2006). PKR1 encodes an assembly factor for the yeast V-type ATPase. The Journal of biological chemistry, 281(42), 32025-35.
  27. Lin H, Li L, Jia X, Ward DM, Kaplan (2011). Genetic and biochemical analysis of high iron toxicity in yeast: iron toxicity is due to the accumulation of cytosolic iron and occurs under both aerobic and anaerobic conditions. The Journal of biological chemistry, 286(5), 3851-62.
  28. Ward DM, Chen OS, Li L, Kaplan J, Bhuiyan SA, Natarajan SK, Bard M, Cox J (2018). Altered sterol metabolism in budding yeast affects mitochondrial iron-sulfur (Fe-S) cluster synthesis. The Journal of biological chemistry, 293(27), 10782-10795.
  29. Yien YY, Shi J, Chen C, Cheung JTM, Grillo AS, Shrestha R, Li L, Zhang X, Kafina MD, Kingsley PD, King MJ, Ablain J, Li H, Zon LI, Palis J, Burke MD, Bauer DE, Orkin SH, Koehler CM, Phillips JD, Kaplan J, Ward DM, Lodish HF, Paw B (2018). FAM210B is an erythropoietin target and regulates erythroid heme synthesis by controlling mitochondrial iron import and ferrochelatase activity. The Journal of biological chemistry, 293(51), 19797-19811.
  30. Ward DM, Vaughn MB, Shiflett SL, Pollock AL, White P, Schnegelberger R, Sundquist WI, Kaplan (2005). The Role of LIP5 and CHMP5 in Multivesicular Body Formation and HIV-1 Budding in Mammalian Cells. The Journal of biological chemistry, 280, 10548-55.
  31. Lin H, Kumanovics A, Warner D, Nelson J, Ward DM,and Kaplan (2008). A single amino acid change in the yeast vacuolar metal transporters zrc1 and cot1 alters their substrate specificity. The Journal of biological chemistry, 283, 33865-33873.
  32. Li L, Jia X, Ward DM, and Kaplan (2011). Identification of Yap5-regulated high iron responsive genes in yeast. The Journal of biological chemistry, 286, 38488-38497.
  33. Kumanovics A, Chen O, Li L, Adkins E, Bagely D, Lin H, Outten C, Keller G, Winge D, Ward DM, and Kaplan, (2008). Identification of FRA1 and FRA2 as genes involved in regulating the yeast iron regulon in response to decreased mitochondrial iron-sulfur cluster synthesis. The Journal of biological chemistry, 283, 10276-10286.
  34. Li L, Murdock G, Bagley D, Jia X, Ward DM,and Kaplan, (2010). Genetic dissection of a mitochonria to vacuole signaling pathway in yeast reveals a link between chronic oxidative stress and vacuolar iron transport. The Journal of biological chemistry, 285(14), 10232-10242.
  35. Bagley DC, Paradkar PN, Durchfort N, Kaplan J, Ward D (2012). Mon1a protein acts in trafficking through the secretory apparatus. The Journal of biological chemistry, 287, 25577-25588.
  36. Li l, Miao R, Bertram S, Jia X, Ward DM ,and Kaplan (2012). A role for iron-sulfur clusters in the regulation of Yap5-dependent high iron transcriptional responses in yeast. The Journal of biological chemistry, 287, 35709-35721.
  37. Li L, Miao R, Jia X, Ward DM, and Kaplan J (2014). Expression of the yeast cation diffusion facilitators Mmt1 and Mmt2 affects mitochondrial and cellular iron homeostasis: evidence for mitochondrial iron export. The Journal of biological chemistry, 289(24), 17132-17141.
  38. Chung J, Anderson SA, Gwynn B, Deck KM, Chen MJ, Langer NB, Shaw GC, Huston NC, Boyer LA, Datta S, Paradkar PN, Li L, Wei Z, Lambert AJ, Sahr K, Wittig JG, Chen W, Lu W, Galy B, Schlaeger TM, Hentze MW, Ward DM, Kaplan J, Eisenstein RS, Peters LL, Paw B (2014). Iron Regulatory Protein-1 protects against mitoferrin1-deficient Porphyria. The Journal of biological chemistry, 289(11), 7835-7843.
  39. Seguin A, Takahashi-Makise N, Yien YY, Huston NC, Whitman JC, Musso G, Wallace JA, Bradley T, Bergonia H, Kafina MD, Matsumoto M, Igarashi K, Phillips JD, Paw BH, Kaplan J, and Ward DM (2017). Reductions in the mitochondrial ABC transporter Abcb10 affect the transcriptional profile of heme biosynthesis genes. The Journal of biological chemistry, 292(39), 16284-16299.
  40. Li L, Kaplan J, and Ward D (2018). The glucose sensor Snf1 and the transcription factors Msn2 and Msn4 regulate transcription of the vacuolar iron importer gene CCC1 and iron resistance in yeast. The Journal of biological chemistry, 292(37), 15577-15586.
  41. Li L, Bertram S, Kaplan J, Jia X, Ward D (2020). The mitochondrial iron exporter genes MMT1 and MMT2 in yeast are transcriptionally regulated by Aft1 and Yap1. The Journal of biological chemistry, 295(6), 1716-1726.
  42. Seguin A, Jia X, Wallace JA, Qiu A, Bradley T, Shrestha R, Li L, Hockin M, Titen S, Warner D, Dowdle PT, Wohlfahrt ME, Hillas E, Firpo M, Phillips JD, Paw BH, Barasch J, Kaplan J, Ward DM (2020). Targeted deletion of mouse Slc25a28 (Mitoferrin2) reveals Mitoferrin1 and Mitoferrin2 are necessary for cell proliferation. The Journal of biological chemistry, 295, 11002-11020.
  43. De Domenico I, Zhang TY, Koening CL, Branch RW, London N, Lo E, Daynes RA, Kushner JP, Li D, Ward DM, Kaplan J (2010). Hepcidin mediates transcriptional changes that modulate acute cytokine-induced inflammatory responses in mice. The Journal of clinical investigation, 120(7), 2395-2405.
  44. Yien YY Robledo RF, Schultz IJ, Takahashi-Makise N, Gwynn B, Bauer DE, Dass A, Yi G, Li L, Hildick-Smith GJ, Cooney JD, Pierce EL, Mohler K, Dailey TA, Miyata N, Kingsley PD, Garone C, Hattangadi SM, Huang H, Chen W, Keenan EM, Shah DI, Schlaeger TM, DiMauro S, Orkin SH, Cantor AB, Palis J, Koehler CM, Lodish HF, Kaplan J, Ward DM, Dailey HA, Phillips JD, Peters LL, Paw BH (2014). TMEM14C is required for erythroid mitochondrial heme metabolism. The Journal of clinical investigation, 124(10), 4294-4304.
  45. Zhu W, Shi DS, Winter JM, Rich BE, Tong Z, Sorensen LK, Zhao H, Huang Y, Tai Z, Mleynek TM, Yoo JH, Dunn C, Ling J, Bergquist JA, Richards JR, Jiang A, Hartnett ME, Ward DM, Mueller AL, Ostanin K, Thomas KR, Odelberg SJ and Li DY (2017). ARF6 is an actionable node that controls VEGFR2 trafficking and signaling in Diabetic Retinopathy. The Journal of clinical investigation,
  46. Girelli D, DeDomenico I, Bozzini C, Campostrini N, Busti F, Castagna A, Soriani N, Cremonesi L, Ferrari M, Colombari R, Ward DM, Kaplan J and Corrocher R (2008). Clinical, pathological, and molecular correlates in ferroportin disease. A study of two novel mutations. Journal of hepatology, 49, 664-671.
  47. Alexander M, Ramstead A, Bauer K, Runtsch MC, Wallace J, Huffaker TB, Larsen DK, Tolmachova T, Seabra MC, Round JL, Ward DM, and O'Connell RM (2017). Rab27-dependent exosome production inhibits chronic inflammation and enables acute responses to inflammatory stimuli. Journal of immunology (Baltimore, Md., 199(10), 3559-3570.
  48. MacQueen BC, Christense RD, Ward DM, Yaish HM, Bennett ST, O'Brien EA, Sheffied MJ, Baer VL, Now GL, Weaver Lewis K, Fleming RE, and Kaplan J (2016). The iron status at birth of neonates who are "at risk" for developing iron deficiency; a pilot study. Journal of perinatology, 1-5.
  49. Gerday E, Brereton JB, Bahr TM, Elmont JO, Fullmer S, Middleton BA, Ward DM, Ohls RK, Christensen R (2020). Urinary ferritin; a potential noninvasive way to screen NICU patients for iron deficiency. Journal of perinatology, 41(6), 1419-1425.
  50. Bahr TM, Baer VL, Ohls RK, Christensen TR, Ward DM, Bennett ST, Christensen R (2020). Reconciling markedly discordant values of serum ferritin versus reticulocyte hemoglobin content. Journal of perinatology, 41(3), 619-626.
  51. Langelier C, von Schwedler UK, Fisher RD, De Domenico I, White PL, Hill CP, Kaplan J, Ward D, Sundquist W (2006). Human ESCRT-II complex and its role in human immunodeficiency virus type 1 release. Journal of virology, 80(19), 9465-80.
  52. Baldauf L, Endres T, Scholz J, Kirches E, Ward DM, Lessmann V, Borucki K, Mawrin (2019). Mitoferrin-1 is required for brain energy metabolism and hippocampus-dependent memory. Neuroscience letters, 713, 134521.
  53. Paradkar PN, Zumbrennan KB, Paw BH, Ward DM, Kaplan J (2009). Regulation of mitochondrial iron import through differential turnover of mitoferrin1 and mitoferrin2. Molecular and cellular biology, 29, 1007-1016.
  54. Shah DI, Takahashi-Makise N, Cooney JD, Seguin A, Li L, Schultz IJ, Pierce EL, Narla A, Hattangadi SM, Langer NB, Chen W, Dailey TA, Medlock AE, Hurst SN, Faccenda D, Wiwczar JM, Vogin G, Brugnara C, Zhou Y, Ward DM, Ebert BL, Campanella M, Danial NN, Dailey HA, Kaplan J and Paw B (2012). Mitochondrial Atpif1 regultes heme synthesis in developing erythroblasts. Nature, 491, 608-612.
  55. Huynh C, Roth D, Ward DM, Kaplan J, Andrews N (2004). Defective Lysosomal Exocytosis and Plasma Membrane Repair in Chediak-Higashi/Beige Cells. Proceedings of the National Academy of Sciences of the United States of America, 101, 16795-80.
  56. De Domenico I, Ward DM, Nemeth E, Vaughn MB, Musci G, Ganz T, Kaplan (2005). Molecular Basis for Ferroportin-Linked Hemochromatosis. Proceedings of the National Academy of Sciences of the United States of America, 102, 8955-60.
  57. DeDomenico I, Ward DM, and Kaplan J (2009). Serum ferritin regulates blood vessel formation: a role beond iron storage. Proceedings of the National Academy of Sciences of the United States of America, 106, 1683-1684.
  58. Chen W, Paradkar PN, Li L, Pierce EL, Langer NB, Takahashi-Makise N, Hyde BB, Shirihai OS, Ward DM, Kaplan J, Paw BH (2009). Abcb10 physically interacts with mitoferrin-1 (Slc25a37) to enhance its stability and function in the erythroid mitochondria. Proceedings of the National Academy of Sciences of the United States of America, 106, 16263-16268.
  59. Nemeth E, Tuttle MS, Powelson J, Vaughn MB, Donovan A, Ward DM, Ganz T, Kaplan (2004). Hepcidin regulates cellular iron efflux by binding to ferroportin and inducing its internalization. Science (New York, N.Y.), 306(5704), 2090-3.
  60. Meznarich JA, Draper L, Christensen RD, Yaish HM, Luem ND, Pysher TJ, Jung G, Nemeth E, Ganz T, Ward D (2018). Fetal presentation of congenital dyserythropoietic anemia type 1 with novel compound heterozygous CDAN1 mutations. Blood cells, molecules & diseases, 71, 63-66.
  61. Bahr TM, Christensen RD, Ward DM, Meng F, Jackson LK, Doyle K, Christensen DR, Harvey AG, Yaish H (2019). Ferritin in serum and urine: A pilot study. Blood cells, molecules & diseases, 76, 59-62.
  62. Yaish HM, Farrell CP, Christensen RD, MacQueen BC, Jackson LK, Trochez-Encisco J, Kaplan J, Ward DM, Salah WK, Phillips J (2017). Two novel mutations in TMPRSS6 associated with irion-refractory iron deficiency anemia in a mother and child. Blood cells, molecules & diseases, 65, 38-40.
  63. MacQueen BC, Christensen RD, Baer VL, Ward DM, Snow G (2019). Screening umbilical cord blood for congenital Iron deficiency. Blood cells, molecules & diseases, 77, 95-100.
  64. Bahr TM, Ward DM, Jia X, Ohls RK, German KR, Christensen R (2021). Is the erythropoietin-erythroferrone-hepcidin axis intact in human neonates?. Blood cells, molecules & diseases, 88, 102536.
  65. Li L, Bagley D, Ward DM, and Kaplan (2008). Yap5p is an iron-responsive transcriptional activator that regulates vacuolar iron storage. Molecular cell biology research communications, 4, 1326-1337.
  66. Shiflett SL, Vaughn MB, Huynh D, Kaplan J, Ward D (2004). Bph1p, the Saccharomyces cerevisiae homologue of CHS1/beige, functions in cell wall formation and protein sorting. Traffic (Copenhagen, Denmark), 5(9), 700-10.
  67. Durchfort N, Verhoef S, Vaughn MB, Shrestha R, Adam D, Kaplan J, Ward D (2012). The enlarged lysosomes in beige cells result from decreased lysosome fission and not increased lysosome fusion. Traffic (Copenhagen, Denmark), 13, 108-119.
  68. Kieffer C, Skalicky JJ, Morita E, DeDomenico I, Ward DM, Kaplan J, and Sundquist W (2008). A new functionally important ESCRT-III recognition mode for the VPS4 ATPases. Developmental cell, 15, 62-73.
  69. Koening CL, Mu HH, Van Schelt A, Lo E, Ward DM, Kaplan J, De Domenico I (2009). Hepcidin is elevated in mice injected with Mycoplasma arthritidis. Journal of inflammation (London, England), 6 (1), 33.
  70. White C, Yuan X, Schmidt PJ, Bresciani E, Samuel TK, Campagna D, Hall C, Bishop K, Calicchio ML, Lapierre A, Ward DM, Liu P, Fleming MD, Hamza (2013). HRG1 is essential for heme transport from the phagolysosome of macrophages during erythrophagocytosis. Cell metabolism, 17(2), 261-70.
  71. Nilsson RH, Schultz IJ, Pierce EL, Soltis KA, Naranuntarat A, Ward DM, Baughman JM, Paradkar PN, Kingsley PD, Culotta VC, Kaplan J, Palis J, Paw BH and Mootha V (2009). Identifying genes essential for heme biosynthesis through large-scale gene expression screening. Cell metabolism, 2, 119-130.
  72. Chen, C,Garcia-Santos, D, Ishikawa, Y, Sequin, A, Fegan, KH, Hildick-Smith, GJ, Cooney, JD, Chen, W, King, MJ, Schultz, IJ, Dalton, AJ, Kingsley, PD, Palis, J, Hattangadi, SM, Lodish, HF, Ward DM, Kaplan, J, Maeda, T, Ponka, P, Paw, B (2013). Snx3 regulates recycling of the transferring receptor and iron assimilation. Cell metabolism, 17, 343-352.
  73. Wilson J, Huynh C, Ward DM, Kaplan J, Kennedy K, Aderem A, and Andrews N.W (2008). A LYST/Beige-mediated innate response restricts parasitophorous vacuole expansion and inhibits Leishmania growth. PLoS pathogens, 4, 1-9.
  74. Ben-Othman R, Flannery A Miguel, Ward DM, Kaplan J and Andrews N (2014). Leishmania-mediated inhibition of iron export promotes parasite replication in macrophages. PLoS pathogens, 10(1), e1003901.
  75. Alexander M, Hu R, Runtsch MC, Kaegle DA, Round JL, Ward DM, and O'Connell R (2015). Exosome-delivered microRNAs reprogram the inflammatory response to endotoxin. Nature communications, 6, 7321.
  76. Kaplan J, Ward D (2015). Muscle specific iron deficiency has systemic consequences. EBioMedicine, 11, 1582-1583.
  77. Feldkamp ML, Ward DM, Pysher TJ, and Chambers C (2017). Chlamydia trachomatis is responsible for lipid vacuolation in the amniotic epithelium of fetal gastroschisis. Birth defects research, 109(13), 1003-1010.
  78. Klag KA, Bell R, Jia X, Seguin A, Maschek JA, Bronner M, Cox JE, Round JL, Ward D (2024). Low-Iron Diet-Induced Fatty Liver Development Is Microbiota Dependent and Exacerbated by Loss of the Mitochondrial Iron Importer Mitoferrin2. Nutrients, 16(12),
  79. Miljkovic M, Seguin A, Jia X, Cox JE, Catrow JL, Bergonia H, Phillips JD, Stephens WZ, Ward D (2023). Loss of the mitochondrial protein Abcb10 results in altered arginine metabolism in MEL and K562 cells and nutrient stress signaling through ATF4. The Journal of biological chemistry, 299(7), 104877.
  80. Bauer KM, Nelson MC, Tang WW, Chiaro TR, Brown DG, Ghazaryan A, Lee SH, Weis AM, Hill JH, Klag KA, Tran VB, Thompson JW, Ramstead AG, Monts JK, Marvin JE, Alexander M, Voth WP, Stephens WZ, Ward DM, Petrey AC, Round JL, O'Connell R (2022). CD11c+ myeloid cell exosomes reduce intestinal inflammation during colitis. JCI insight, 7(19),
  81. Hackett PT, Jia X, Li L, Ward D (2022). Posttranslational regulation of mitochondrial frataxin and identification of compounds that increase frataxin levels in Friedreich's ataxia. The Journal of biological chemistry, 298(6), 101982.
  82. Meng F, Fleming BA, Jia X, Rousek AA, Mulvey MA, Ward D (2022). Lysosomal iron recycling in mouse macrophages is dependent upon both LcytB and Steap3 reductases. Blood advances, 6(6), 1692-1707.
  83. Bahr TM, Christensen TR, Henry E, Wilkes J, Ohls RK, Bennett ST, Ward DM, Pysher TJ, Christensen R (2021). Neonatal Reference Intervals for the Complete Blood Count Parameters MicroR and HYPO-He: Sensitivity Beyond the Red Cell Indices for Identifying Microcytic and Hypochromic Disorders. The Journal of pediatrics, 239, 95-100.e2.
  84. Cox HD, Miller GD, Manandhar A, Husk JD, Jia X, Marvin J, Ward DM, Phillips J, Eichner (2021). Measurement of Immature Reticulocytes in Dried Blood Spots by Mass Spectrometry. Clinical chemistry, 67(8), 1071-1079.
  85. Ward DM, Gillis S, Yien Y (2024). Radiolabeling Heme and Porphyrin with (14)C-Glycine or (14)C ¿-Aminolevulinic Acid. Methods in molecular biology (Clifton, N.J.), 2839, 225-231.
  86. Sun W, Jia X, Liesa M, Tantin D, Ward D (2022). ABCB10 Loss Reduces CD4(+) T Cell Activation and Memory Formation. Journal of immunology (Baltimore, Md., 208(2), 328-337.
  87. Christensen RD, Bahr TM, Ward D (2022). Iron deficiency in newborn infants: global rewards for recognizing and treating this silent malady. Newborn (Clarksville, Md.), 1(1), 97-103.
  88. Bahr TM, Carr NR, Christensen TR, Wilkes J, O'Brien EA, German KR, Ohls RK, Ward DM, Christensen R (2021). Early iron supplementation and iron sufficiency at one month of age in NICU patients at-risk for iron deficiency. Blood cells, molecules & diseases, 90, 102575.
  89. Seguin A, Jia X, Earl AM, Li L, Wallace J, Qiu A, Bradley T, Shrestha R, Troadec MB, Hockin M, Titen S, Warner DE, Dowdle PT, Wohlfahrt ME, Hillas E, Firpo MA, Phillips JD, Kaplan J, Paw BH, Barasch J, Ward D (2020). The mitochondrial metal transporters mitoferrin1 and mitoferrin2 are required for liver regeneration and cell proliferation in mice. The Journal of biological chemistry, 295(32), 11002-11020.
  90. De Domenico I, Vaughn MB, Paradkar PN, Lo E, Ward DM, Kaplan (2011). Decoupling ferritin synthesis from free cytosolic iron results in ferritin secretion. Cell metabolism, 13(1), 57-67.
  91. Troadec MB, Ward DM, Lo E, Kaplan J, De Domenico (2010). Induction of FPN1 transcription by MTF-1 reveals a role for ferroportin in transition metal efflux. Blood, 116(22), 4657-64.
  92. De Domenico I, Zhang TY, Koening CL, Branch RW, London N, Lo E, Daynes RA, Kushner JP, Li D, Ward DM, Kaplan (2010). Hepcidin mediates transcriptional changes that modulate acute cytokine-induced inflammatory responses in mice. The Journal of clinical investigation, 120(7), 2395-405.
  93. Li L, Murdock G, Bagley D, Jia X, Ward DM, Kaplan (2010). Genetic dissection of a mitochondria-vacuole signaling pathway in yeast reveals a link between chronic oxidative stress and vacuolar iron transport. The Journal of biological chemistry, 285(14), 10232-42.
  94. De Domenico I, Ward DM, Kaplan (2010). Autophagy, ferritin and iron chelation. Autophagy, 6(1), 157.
  95. Koening CL, Mu HH, Van Schelt A, Lo E, Ward DM, Kaplan J, De Domenico (2009). Hepcidin is elevated in mice injected with Mycoplasma arthritidis. Journal of inflammation (London, England), 6, 33.
  96. Koening CL, Miller JC, Nelson JM, Ward DM, Kushner JP, Bockenstedt LK, Weis JJ, Kaplan J, De Domenico (2009). Toll-like receptors mediate induction of hepcidin in mice infected with Borrelia burgdorferi. Blood, 114(9), 1913-8.
  97. Lin H, Burton D, Li L, Warner DE, Phillips JD, Ward DM, Kaplan (2009). Gain-of-function mutations identify amino acids within transmembrane domains of the yeast vacuolar transporter Zrc1 that determine metal specificity. The Biochemical journal, 422(2), 273-83.
  98. De Domenico I, Lo E, Ward DM, Kaplan (2009). Hepcidin-induced internalization of ferroportin requires binding and cooperative interaction with Jak2. Proceedings of the National Academy of Sciences of the United States of America, 106(10), 3800-5.
  99. De Domenico I, Ward DM, Kaplan (2009). Serum ferritin regulates blood vessel formation: a role beyond iron storage. Proceedings of the National Academy of Sciences of the United States of America, 106(6), 1683-4.
  100. Paradkar PN, Zumbrennen KB, Paw BH, Ward DM, Kaplan (2009). Regulation of mitochondrial iron import through differential turnover of mitoferrin 1 and mitoferrin 2. Molecular and cellular biology, 29(4), 1007-16.
  101. Lin H, Kumánovics A, Nelson JM, Warner DE, Ward DM, Kaplan (2008). A single amino acid change in the yeast vacuolar metal transporters ZRC1 and COT1 alters their substrate specificity. The Journal of biological chemistry, 283(49), 33865-73.
  102. De Domenico I, Nemeth E, Nelson JM, Phillips JD, Ajioka RS, Kay MS, Kushner JP, Ganz T, Ward DM, Kaplan (2008). The hepcidin-binding site on ferroportin is evolutionarily conserved. Cell metabolism, 8(2), 146-56.
  103. Paradkar PN, De Domenico I, Durchfort N, Zohn I, Kaplan J, Ward D (2008). Iron depletion limits intracellular bacterial growth in macrophages. Blood, 112(3), 866-74.
  104. Kieffer C, Skalicky JJ, Morita E, De Domenico I, Ward DM, Kaplan J, Sundquist W (2008). Two distinct modes of ESCRT-III recognition are required for VPS4 functions in lysosomal protein targeting and HIV-1 budding. Developmental cell, 15(1), 62-73.
  105. Kumánovics A, Chen OS, Li L, Bagley D, Adkins EM, Lin H, Dingra NN, Outten CE, Keller G, Winge D, Ward DM, Kaplan (2008). Identification of FRA1 and FRA2 as genes involved in regulating the yeast iron regulon in response to decreased mitochondrial iron-sulfur cluster synthesis. The Journal of biological chemistry, 283(16), 10276-86.
  106. Li L, Bagley D, Ward DM, Kaplan (2008). Yap5 is an iron-responsive transcriptional activator that regulates vacuolar iron storage in yeast. Molecular and cellular biology, 28(4), 1326-37.
  107. De Domenico I, Vaughn MB, Yoon D, Kushner JP, Ward DM, Kaplan (2007). Zebrafish as a model for defining the functional impact of mammalian ferroportin mutations. Blood, 110(10), 3780-3.
  108. De Domenico I, Ward DM, Kaplan (2007). Hepcidin regulation: ironing out the details. The Journal of clinical investigation, 117(7), 1755-8.
  109. De Domenico I, Ward DM, di Patti MC, Jeong SY, David S, Musci G, Kaplan (2007). Ferroxidase activity is required for the stability of cell surface ferroportin in cells expressing GPI-ceruloplasmin. The EMBO journal, 26(12), 2823-31.
  110. De Domenico I, Vaughn MB, Li L, Bagley D, Musci G, Ward DM, Kaplan (2006). Ferroportin-mediated mobilization of ferritin iron precedes ferritin degradation by the proteasome. The EMBO journal, 25(22), 5396-404.
  111. Ward DM, Vaughn MB, Shiflett SL, White PL, Pollock AL, Hill J, Schnegelberger R, Sundquist WI, Kaplan (2005). The role of LIP5 and CHMP5 in multivesicular body formation and HIV-1 budding in mammalian cells. The Journal of biological chemistry, 280(11), 10548-55.
  112. Ardon O, Bussey H, Philpott C, Ward DM, Davis-Kaplan S, Verroneau S, Jiang B, Kaplan (2001). Identification of a Candida albicans ferrichrome transporter and its characterization by expression in Saccharomyces cerevisiae. The Journal of biological chemistry, 276(46), 43049-55.

Review

  1. Ward DM, Cloonan S (2019). Mitochondrial Iron in Human Health and Disease. Annual review of physiology, 81, 453-482.
  2. Kaplan J, Ward DM, Crisp RJ, Philpott C (2006). Iron-dependent metabolic remodeling in S. cerevisiae. Biochimica et biophysica acta, 1763(7), 646-51.
  3. Ward DM, Kaplan (2012). Ferroportin-mediated iron transport: expression and regulation. Biochimica et biophysica acta, 1823(9), 1426-33.
  4. Kaplan J, Ward DM, De Domenico (2011). The molecular basis of iron overload disorders and iron-linked anemias. International journal of hematology, 93(1), 14-20.
  5. Li L and Ward D (2017). Iron Toxicity in Yeast: Transcriptional Regulation of the Vacuolar Iron Importer Ccc1. Current genetics,
  6. De Domenico I, Ward DM, and Kaplan, (2011). Hepcidin and Ferroportin: The new players in iron metabolism. Seminars in liver disease, 31, 272-279.
  7. Kaplan J, Ward D (2013). The essential nature of iron usage and regulation: variations on a theme. Current biology, 15, :R642-64.
  8. Sequin A, Ward DM, and Kaplan, (2011). Regulation of ribonucleotide reductase during iron limitation. Molecular cell, 44, 683-684.
  9. DeDomenico I, Ward DM, and Kaplan (2008). The genetic and cellular basis of iron-linked disorders. Nature reviews. Molecular cell biology, 1, 72-81.
  10. Troadec MB, Ward DM, Kaplan (2009). A Tf-independent iron transport system required for organogenesis. Developmental cell, 16(1), 3-4.
  11. Takahashi-Makise N, Ward DM, and Kaplan (2009). On the mechanism of iron sensing by IRP2: new players, new paradigms. Nature chemical biology, 5, 874-875.
  12. DeDomenico I, Ward DM, and Kaplan (2009). Autophagy, ferritin and iron chelation. Autophagy, 6, 157.
  13. Klionsky DJ, Adballa FC, Abeliovich H, Abraham RT et al., Ward DM, et al., Zuckerbrau (2012). Guidelines for the use and interpretation of assays for monitoring autophagy. Autophagy, 8, 445-544.
  14. Klionsky DJ, Abdelmohsen K, Abe A, Abedin MJ, Abeliovich H., Ward DM, et a (2016). Guidelines for the use and interpretation of assays for monitoring autophagy (3rd edition). Autophagy, 12(1), 1-222.
  15. Seguin A and Ward D (2019). Mitochondrial ABC Transporters and Iron Metabolism. Journal of clinical & experimental pathology,
  16. Kaplan J, De Domenico I, Ward D (2008). Chediak-Higashi syndrome. Current opinion in hematology, 15(1), 22-9.
  17. Ward DM, Griffiths GM, Stinchcombe JC, Kaplan (2000). Analysis of the lysosomal storage disease Chediak-Higashi syndrome. Traffic (Copenhagen, Denmark), 1(11), 816-22.

Book Chapter

  1. Shrestha R, Kaplan J, Ward D (2017). Conventional and Secretory Lysosomes. 2, 225-234.

Commentary

  1. Bahr TM, Benson AE, Kling PJ, Ohls RK, Ward DM, Christensen R (2021). Maternal obesity and impaired offspring neurodevelopment: could fetal iron deficiency be a pathogenic link?. Journal of perinatology, doi: 10.1038/s41372-021-00951-9. Epub ahead of print. PMID: 33594227,

Letter

  1. Bahr TM, Benson AE, Kling PJ, Ohls RK, Ward DM, Christensen R (2021). Maternal obesity and impaired offspring neurodevelopment: could fetal iron deficiency be a pathogenic link?. Journal of perinatology, 41(5), 1199-1200.

Video/Film/CD/Web/Podcast

  1. Kaplan J, Ward D (2013). Molecular Genetics of Chediak'Higashi Syndrome.