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Diane McVey Ward

Diane McVey Ward, PhD

Languages spoken: English

Academic Information

Departments Primary - Pathology


Academic Office Information

Research Interests

  • Metal Homeostasis
  • Macrophage Iron Homeostasis
  • Mitochondrial Heme and Iron
  • Membrane Trafficking, Lysosomes and Exosomes
  • Iron Metabolism
  • Cell Biology

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

Postdoctoral Fellowship University of Utah
Postdoctoral Fellow
University of Utah
Undergraduate Brigham Young University

Selected Publications

Journal Article

  1. Bahr TM, Ward DM, Jia X, Ohls RK, German KR, Christensen RD (2021). Is the erythropoietin-erythroferrone-hepcidin axis intact in human neonates? Blood Cells Mol Dis, 88, 102536.
  2. Bahr TM, Baer VL, Ohls RK, Christensen TR, Ward DM, Bennett ST, Christensen RD (2020). Reconciling markedly discordant values of serum ferritin versus reticulocyte hemoglobin content. J Perinatol, 41(3), 619-626.
  3. Gerday E, Brereton JB, Bahr TM, Elmont JO, Fullmer S, Middleton BA, Ward DM, Ohls RK, Christensen RD (2020). Urinary ferritin; a potential noninvasive way to screen NICU patients for iron deficiency. J Perinatol, 41(6), 1419-1425.
  4. Li L, Bertram S, Kaplan J, Jia X, Ward DM (2020). The mitochondrial iron exporter genes MMT1 and MMT2 in yeast are transcriptionally regulated by Aft1 and Yap1. J Biol Chem, 295(6), 1716-1726.
  5. 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. J Biol Chem, 295, 11002-11020.
  6. Baldauf L, Endres T, Scholz J, Kirches E, Ward DM, Lessmann V, Borucki K, Mawrin C (2019). Mitoferrin-1 is required for brain energy metabolism and hippocampus-dependent memory. Neurosci Lett, 713, 134521.
  7. MacQueen BC, Christensen RD, Baer VL, Ward DM, Snow GL (2019). Screening umbilical cord blood for congenital Iron deficiency. Blood Cells Mol Dis, 77, 95-100.
  8. Bahr TM, Christensen RD, Ward DM, Meng F, Jackson LK, Doyle K, Christensen DR, Harvey AG, Yaish HM (2019). Ferritin in serum and urine: A pilot study. Blood Cells Mol Dis, 76, 59-62.
  9. 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 BH (2018). FAM210B is an erythropoietin target and regulates erythroid heme synthesis by controlling mitochondrial iron import and ferrochelatase activity. J Biol Chem, 293(51), 19797-19811.
  10. Ward DM, Chen OS, Li L, Kaplan J, Bhuiyan SA, Natarajan SK, Bard M, Cox JE (2018). Altered sterol metabolism in budding yeast affects mitochondrial iron-sulfur (Fe-S) cluster synthesis. J Biol Chem, 293(27), 10782-10795.
  11. Meznarich JA, Draper L, Christensen RD, Yaish HM, Luem ND, Pysher TJ, Jung G, Nemeth E, Ganz T, Ward DM (2018). Fetal presentation of congenital dyserythropoietic anemia type 1 with novel compound heterozygous CDAN1 mutations. Blood Cells Mol Dis, 71, 63-66.
  12. Li, L, Kaplan, J and Ward, DM (2017). 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. J Biol Chem, 292(37), 15577-15586.
  13. 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. J Clin Invest.
  14. Alexander, M, Ramstead, A, Bauer, K, Runtsch, MC, Wallace, J, Huffaker, TB, Larsen, DK, Tolmachova, T, Seabra, MC, Round, JL, Ward, DM and OConnell, RM (2017). Rab27-dependent exosome production inhibits chronic inflammation and enables acute responses to inflammatory stimuli. J Immunol, 199(10), 3559-3570.
  15. Feldkamp, ML, Ward, DM, Pysher, TJ and Chambers, CT (2017). Chlamydia trachomatis is responsible for lipid vacuolation in the amniotic epithelium of fetal gastroschisis. Birth Defects Res, 109(13), 1003-1010.
  16. 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. J Biol Chem, 292(39), 16284-16299.
  17. Yaish HM, Farrell CP, Christensen RD, MacQueen BC, Jackson LK, Trochez-Encisco J, Kaplan J, Ward DM, Salah WK, Phillips JD (2017). Two novel mutations in TMPRSS6 associated with irion-refractory iron deficiency anemia in a mother and child. Blood Cells Mol Dis, 65, 38-40.
  18. MacQueen BC, Christense RD, Ward DM, Yaish HM, Bennett ST, OBrien 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. J Perinatol, 1-5.
  19. Alexander M, Hu R, Runtsch MC, Kaegle DA, Round JL, Ward DM, and OConnell R (2015). Exosome-delivered microRNAs reprogram the inflammatory response to endotoxin. Nat Commun, 6, 7321.
  20. Kaplan J, Ward DM (2015). Muscle specific iron deficiency has systemic consequences. EBioMedicine, 11, 1582-1583.
  21. 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. J Clin Invest, 124(10), 4294-4304.
  22. 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. J Biol Chem, 289(24), 17132-17141.
  23. 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 BH (04/01/2014). Iron Regulatory Protein-1 protects against mitoferrin1-deficient Porphyria. J Biol Chem, 289(11), 7835-7843.
  24. Ben-Othman R, Flannery A Miguel, Ward DM, Kaplan J and Andrews NW (2014). Leishmania-mediated inhibition of iron export promotes parasite replication in macrophages. PLoS Pathog, 10(1), e1003901.
  25. 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, BH (2013). Snx3 regulates recycling of the transferring receptor and iron assimilation. Cell Metab, 17, 343-352.
  26. 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 I (2013). HRG1 is essential for heme transport from the phagolysosome of macrophages during erythrophagocytosis. Cell Metab, 17(2), 261-70.
  27. Li l, Miao R, Bertram S, Jia X, Ward DM, and Kaplan J (2012). A role for iron-sulfur clusters in the regulation of Yap5-dependent high iron transcriptional responses in yeast. J Biol Chem, 287, 35709-35721.
  28. 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 BH (2012). Mitochondrial Atpif1 regultes heme synthesis in developing erythroblasts. Nature, 491, 608-612.
  29. Ward DM, and Kaplan J (2012). Ferroportin-mediated iron transport: expression and regulation. Biochim Biophys Acta, 1823, 1426-1433.
  30. Bagley DC, Paradkar PN, Durchfort N, Kaplan J, Ward DM (2012). Mon1a protein acts in trafficking through the secretory apparatus. J Biol Chem, 287, 25577-25588.
  31. Durchfort N, Verhoef S, Vaughn MB, Shrestha R, Adam D, Kaplan J, Ward DM (2012). The enlarged lysosomes in beige cells result from decreased lysosome fission and not increased lysosome fusion. Traffic, 13, 108-119.
  32. Li L, Jia X, Ward DM, and Kaplan J (2011). Identification of Yap5-regulated high iron responsive genes in yeast. J Biol Chem, 286, 38488-38497.
  33. Troadec MB, Warner D, Wallace J, Thomas K, Spangrude GJ, Phillips J, Khalimonchuk O, Paw BH, Ward DM, Kaplan J (2011). Targeted deletion of the mouse Mitoferrin1 gene: from anemia to protoporphyria. Blood, 117(20), 5494-502.
  34. Troadec MB, Warner DE, Wallace J, Phillips J, Paw BH, Ward DM, and Kaplan J (2011). Selective loss of mitoferrin 1 in mouse tissues reveals organ specific defects. Blood, 117, 5494-5502.
  35. Lin H, Li L, Jia X, Ward DM, Kaplan J (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. J Biol Chem, 286(5), 3851-62.
  36. Troadec MB, Ward DM, Lo E, Kaplan J, De Domenico I (5/14/11). Induction of FPN1 transcription by MTF-1 reveals a role for ferroportin in transition metal efflux. Blood, 116(22), 4657-4664.
  37. De Domenico I, Zhang TY, Koening CL, Branch RW, London N, Lo E, Daynes RA, Kushner JP, Li D, Ward DM, Kaplan J (5/14/11). Hepcidin mediates transcriptional changes that modulate acute cytokine-induced inflammatory responses in mice. J Clin Invest, 120(7), 2395-2405.
  38. Li L, Murdock G, Bagley D, Jia X, Ward DM, and Kaplan, J (5/14/11). Genetic dissection of a mitochonria to vacuole signaling pathway in yeast reveals a link between chronic oxidative stress and vacuolar iron transport. J Biol Chem, 285(14), 10232-10242.
  39. De Domenico I, Ward DM, Kaplan J (2009). Specific iron chelators determine the route of ferritin degradation. Blood, 114(20), 4546-51.
  40. 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. J Inflamm (Lond), 6 (1), 33.
  41. Paradkar PN, Zumbrennan KB, Paw BH, Ward DM, Kaplan J (2009). Regulation of mitochondrial iron import through differential turnover of mitoferrin1 and mitoferrin2. Mol Cell Biol, 29, 1007-1016.
  42. DeDomenico I, Ward DM, and Kaplan J (2009). Serum ferritin regulates blood vessel formation: a role beond iron storage. Proc Natl Acad Sci U S A, 106, 1683-1684.
  43. 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.
  44. 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.
  45. 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 VK (2009). Identifying genes essential for heme biosynthesis through large-scale gene expression screening. Cell Metab, 2, 119-130.
  46. 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. Proc Natl Acad Sci U S A, 106, 16263-16268.
  47. Li L, Bagley D, Ward DM, and Kaplan J (2008). Yap5p is an iron-responsive transcriptional activator that regulates vacuolar iron storage. Mol Cell Biol Res Commun, 4, 1326-1337.
  48. Kumanovics A, Chen O, Li L, Adkins E, Bagely D, Lin H, Outten C, Keller G, Winge D, Ward DM, and Kaplan, J (2008). Identification of FRA1 and FRA2 as genes involved in regulating the yeast iron regulon in response to decreased mitochondrial iron-sulfur cluster synthesis. J Biol Chem, 283, 10276-10286.
  49. Paradkar PN, DeDomenico I, Zohn I, Kaplan J, Ward DM (2008). Iron-depletion limits intracellular bacterial growth in macrophages. Blood, 112, 866-874.
  50. 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. J Hepatol, 49, 664-671.
  51. Kieffer C, Skalicky JJ, Morita E, DeDomenico I, Ward DM, Kaplan J, and Sundquist WI (2008). A new functionally important ESCRT-III recognition mode for the VPS4 ATPases. Dev Cell, 15, 62-73.
  52. Wilson J, Huynh C, Ward DM, Kaplan J, Kennedy K, Aderem A, and Andrews NW (2008). A LYST/Beige-mediated innate response restricts parasitophorous vacuole expansion and inhibits Leishmania growth. PLoS Pathog, 4, 1-9.
  53. Lin H, Kumanovics A, Warner D, Nelson J, Ward DM, and Kaplan J (2008). A single amino acid change in the yeast vacuolar metal transporters zrc1 and cot1 alters their substrate specificity. J Biol Chem, 283, 33865-33873.
  54. Wang F, Paradka P, Custodio AO, Ward DM, Fleming MD, Campagna D, Roberts KA, Boyartchuk V, Dietrich WF, Kaplan J and Andrews NC (2007). Genetic variation in Mon1a affects protein trafficking and modifies macrophage iron loading in mice. Nat Genet, 39, 1025-1032.
  55. De Domenico I, Ward DM, Langelier C, Vaughn MB, Nemeth E, Sundquist WI, Ganz T, Musci G, Kaplan J (2007). The molecular mechanism of hepcidin-mediated ferroportin down-regulation. Mol Biol Cell, 18(7), 2569-78.
  56. Zohn IE, De Domenico I, Pollock A, Ward DM, Goodman JF, Liang X, Sanchez AJ, Niswander L, Kaplan J (2007). The flatiron mutation in mouse ferroportin acts as a dominant negative to cause ferroportin disease. Blood, 109(10), 4174-80.
  57. De Domenico I, Ward DM, Musci G, Kaplan J (2007). Evidence for the multimeric structure of ferroportin. Blood, 109(5), 2205-9.
  58. De Domenico I, Ward DM, Carmela Bonaaccorsi di Patti M, Musci G, Kaplan J (2007). Hepcidin-independent regulation of ferroportin: a role for ceruloplasmin in ferroportin-mediated iron export. EMBO J.
  59. Mohlig H, Mathieu S, Thon L, Frederiksen MC, Ward DM, Kaplan J, Schutze S, Kabelitz D, Adam D (2007). The WD repeat protein FAN regulates lysosome size independent from abnormal downregulation/membrane recruitment of protein kinase C. Exp Cell Res.
  60. Wang F, Custodio AO, Paradkar P, Ward DM, Fleming MD, Campagna D, Roberts KA, Boyartchuk V, Kaplan J, Dietrich WF, Andrews NC (2007). Mon 1 a, a modifer of spleen iron loading, regulates macrophage trafficking of the iron transporter ferroportin. Nat Genet.
  61. DeDomenico I, Vaughn MB, Yoon D, Kushner JP, Kaplan J, Ward DM (2007). Zebrafish as a model for defining the functional impact of mammalian ferroportin mutations. Blood, 110, 3780-3783.
  62. De Domenico I, Ward DM, di Patti MC, Jeon SY, David S, Musci G, and Kaplan J (01/01/2007). Ferroxidase activity is required for the stability of cell surface ferroportin in cells expressing GPI-ceruloplasmin.PMID:17541408. EMBO J, 26, 2823-2831.
  63. Davis-Kaplan SR, Compton MA, Flannery AR, Ward DM, Kaplan J, Stevens TH, Graham LA (2006). PKR1 encodes an assembly factor for the yeast V-type ATPase. J Biol Chem, 281(42), 32025-35.
  64. Langelier C, von Schwedler UK, Fisher RD, De Domenico I, White PL, Hill CP, Kaplan J, Ward D, Sundquist WI (2006). Human ESCRT-II complex and its role in human immunodeficiency virus type 1 release. J Virol, 80(19), 9465-80.
  65. Kumanovics A, Poruk KE, Osborn KA, Ward DM, Kaplan J (2006). YKE4 (YIL023C) encodes a bidirectional zinc transporter in the endoplasmic reticulum of Saccharomyces cerevisiae. J Biol Chem, 281(32), 22566-74.
  66. Runkel F, Bussow H, Seburn KL, Cox GA, Ward DM, Kaplan J, Franz T (2006). Grey, a novel mutation in the murine Lyst gene, causes the beige phenotype by skipping of exon 25. Mamm Genome, 17(3), 203-10.
  67. De Domenico I, Ward DM, Musci G, Kaplan J (2006). Iron overload due to mutations in ferroportin. Haematologica, 91(1), 92-5.
  68. De Domenico I, Vaughn MB, Li L, Bagley D, Musci G, Ward DM, Kaplan J (2006). Mobilization of ferritin iron does not require ferritin degradation. EMBO J, 25, 5396-5404.
  69. DeDomenico J, Ward DM, Nemeth E, Ganz T, Corradini E, Ferrara F, Musci G, Pietrangelo A, Kaplan J (2006). Molecular and clinical correlates in iron overload associated with mutations in ferroportin. Hematol Pathol, 91, 1092-1095.
  70. Felice MR, De Domenico I, Li L, Ward DM, Bartok B, Musci G, Kaplan J (2005). Post-transcriptional regulation of the yeast high affinity iron transport system. J Biol Chem, 280(23), 22181-90.
  71. Ward DM, Vaughn MB, Shiflett SL, Pollock AL, White P, Schnegelberger R, Sundquist WI, Kaplan J (2005). The Role of LIP5 and CHMP5 in Multivesicular Body Formation and HIV-1 Budding in Mammalian Cells. J Biol Chem, 280, 10548-55.
  72. De Domenico I, Ward DM, Nemeth E, Vaughn MB, Musci G, Ganz T, Kaplan J (2005). Molecular Basis for Ferroportin-Linked Hemochromatosis. Proc Natl Acad Sci U S A, 102, 8955-60.
  73. Nemeth E, Tuttle MS, Powelson J, Vaughn MB, Donovan A, Ward DM, Ganz T, Kaplan J (2004). Hepcidin regulates cellular iron efflux by binding to ferroportin and inducing its internalization. Science, 306(5704), 2090-3.
  74. Shiflett SL, Vaughn MB, Huynh D, Kaplan J, Ward DM (2004). Bph1p, the Saccharomyces cerevisiae homologue of CHS1/beige, functions in cell wall formation and protein sorting. Traffic, 5(9), 700-10.
  75. Shiflett SL, Ward DM, Huynh D, Vaughn MB, Simmons JC, Kaplan J (2004). Characterization of Vta1p, a class E Vps protein in Saccharomyces cerevisiae. J Biol Chem, 279(12), 10982-90.
  76. Davis-Kaplan SR, Ward DM, Shiflett SL, Kaplan J (2004). Genome-wide analysis of iron-dependent growth reveals a novel yeast gene required for vacuolar acidification. J Biol Chem, 279(6), 4322-9.
  77. Huynh C, Roth D, Ward DM, Kaplan J, Andrews NW (2004). Defective Lysosomal Exocytosis and Plasma Membrane Repair in Chediak-Higashi/Beige Cells. Proc Natl Acad Sci U S A, 101, 16795-80.


  1. Alexandra Seguin and Diane McVey Ward (Feb 2018). Mitochondrial ABC Transporters and Iron Metabolism. [Review]. J Clin Exp Pathol.
  2. Ward DM, Cloonan SM (2019). Mitochondrial Iron in Human Health and Disease. [Review]. Annu Rev Physiol, 81, 453-482.
  3. Li, L and Ward, DM (2017). Iron Toxicity in Yeast: Transcriptional Regulation of the Vacuolar Iron Importer Ccc1. [Review]. Curr Genet.
  4. Klionsky, DJ, Abdelmohsen K, Abe A, Abedin MJ, Abeliovich H, Ward, DM et al (2016). Guidelines for the use and interpretation of assays for monitoring autophagy (3rd edition). [Review]. Autophagy, 12(1), 1-222.
  5. Kaplan J, Ward DM (2013). The essential nature of iron usage and regulation: variations on a theme. [Review]. Curr Biol, 15, :R642-64.
  6. Ward, DM and Kaplan J (2012). Ferroportin-mediated iron transport: expression and regulation. [Review]. Biochim Biophys Acta, 1823(9), 1426-33.
  7. Klionsky DJ, Adballa FC, Abeliovich H, Abraham RT et al, Ward DM, et al, Zuckerbrau B (2012). Guidelines for the use and interpretation of assays for monitoring autophagy. [Review]. Autophagy, 8, 445-544.
  8. Sequin A, Ward DM, and Kaplan, J (2011). Regulation of ribonucleotide reductase during iron limitation. [Review]. Mol Cell, 44, 683-684.
  9. De Domenico I, Ward DM, and Kaplan, J (2011). Hepcidin and Ferroportin: The new players in iron metabolism. [Review]. Semin Liver Dis, 31, 272-279.
  10. Kaplan J, Ward DM, De Domenico I (2011). The molecular basis of iron overload disorders and iron-linked anemias. [Review]. Int J Hematol, 93(1), 14-20.
  11. Takahashi-Makise N, Ward DM, and Kaplan J (2009). On the mechanism of iron sensing by IRP2: new players, new paradigms. [Review]. Nat Chem Biol, 5, 874-875.
  12. DeDomenico I, Ward DM, and Kaplan J (2009). Autophagy, ferritin and iron chelation. [Review]. Autophagy, 6, 157.
  13. Troadec MB, Ward DM, Kaplan J (2009). A Tf-independent iron transport system required for organogenesis. [Review]. Dev Cell, 16(1), 3-4.
  14. DeDomenico I, Ward DM, and Kaplan J (2008). The genetic and cellular basis of iron-linked disorders. [Review]. Nat Rev Mol Cell Biol, 1, 72-81.
  15. Kaplan J, McVey Ward D, Crisp RJ, Philpott CC (2006). Iron-dependent metabolic remodeling in S. cerevisiae. [Review]. Biochim Biophys Acta, 1763(7), 646-51.

Book Chapter

  1. Shrestha R, Kaplan J, Ward DM (2017). Conventional and Secretory Lysosomes. In Encyclopedia of Cell Biology (2, pp. 225-234).


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


  1. Kaplan J, Ward DM (2013). Molecular Genetics of Chediak–Higashi Syndrome [Web]. Wiley Online Library- eLS (Invited Review). Available: