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Monika Ward,PhD

Title
Faculty RankProfessor
InstitutionUniversity of Hawaii at Manoa
DepartmentAnatomy, Biochemistry & Physiology
AddressJohn A. Burns School of Medicine
University of Hawaii at Manoa
1960 East-West Road
Honolulu HI 96822
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    Collapse Research 
    Collapse research activities and funding
    R03HD048446     (WARD, MONIKA A)Dec 5, 2004 - Nov 30, 2007
    NIH
    Preservation of ejaculated mouse spermatozoa
    Role: Principal Investigator
    R21HD048845     (WARD, MONIKA A)Jul 1, 2005 - Jun 30, 2008
    NIH
    Sperm DNA damage in fertilization
    Role: Principal Investigator
    R21HD058059     (WARD, MONIKA A)Apr 1, 2008 - Mar 31, 2012
    NIH
    Minimum Y gene complement necessary for successful ART
    Role: Principal Investigator
    R01HD072380     (WARD, MONIKA A)Aug 25, 2012 - Mar 31, 2024
    NIH
    Do we need Y chromosome for successful reproduction?
    Role: Principal Investigator
    R13HD097765     (WARD, MONIKA A)Jan 16, 2019 - Dec 31, 2019
    NIH
    2019 Germinal Stem Cell Biology Gordon Research Conference
    Role: Principal Investigator
    R03HD106936     (WARD, MONIKA A)Sep 27, 2021 - Sep 26, 2023
    NIH
    The role Y chromosome genes Prssly and Teyorf1 in male reproduction.
    Role: Principal Investigator
    R01HD114645     (WARD, MONIKA A)Aug 5, 2024 - May 31, 2029
    NIH
    The roles of mouse Y chromosome Zfy genes in male reproduction.
    Role: Principal Investigator
    R13HD121171     (WARD, MONIKA A)Sep 26, 2025 - Aug 31, 2026
    NIH
    Vertebrate Sex Determination 2026
    Role: Principal Investigator

    Collapse Bibliographic 
    Collapse selected publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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    PMC Citations indicate the number of times the publication was cited by articles in PubMed Central, and the Altmetric score represents citations in news articles and social media. (Note that publications are often cited in additional ways that are not shown here.) Fields are based on how the National Library of Medicine (NLM) classifies the publication's journal and might not represent the specific topic of the publication. Translation tags are based on the publication type and the MeSH terms NLM assigns to the publication. Some publications (especially newer ones and publications not in PubMed) might not yet be assigned Field or Translation tags.) Click a Field or Translation tag to filter the publications.
    1. Holmlund H, Coul?e M, Yamauchi Y, Yarbabaeva B, Tekayev M, Garcia IR, Feudjio OU, de la Iglesia A, Larcombe L, Ellis PJI, Cocquet J, Ward MA. Large-scale transcriptomic analyses reveal downstream target genes of ZFY1 and ZFY2 transcription factors in male germ cells. Cell Death Differ. 2026 Feb; 33(2):392-410. PMID: 40866672.
      Citations:    Fields:    
    2. Allan NP, Torres A, Corley MJ, Yamamoto BY, Balaan C, Yamauchi Y, Peres R, Qin Y, Khadka VS, Deng Y, Ward MA, Maunakea AK. Pre-Conception Maternal Obesity Confers Autism Spectrum Disorder-like Behaviors in Mice Offspring Through Neuroepigenetic Dysregulation. Cells. 2025 Aug 05; 14(15). PMID: 40801633.
      Citations:    Fields:    Translation:AnimalsCells
    3. Giannakopoulos S, Pak J, Bakse J, Ward MA, Nerurkar VR, Tallquist MD, Verma S. SARS-CoV-2-induced cytokine storm drives prolonged testicular injury and functional impairment in mice that are mitigated by dexamethasone. PLoS Pathog. 2025 Jan; 21(1):e1012804. PMID: 39775442.
      Citations: 2     Fields:    Translation:HumansAnimalsCells
    4. Holmlund H, Yamauchi Y, Ruthig VA, Cocquet J, Ward MA. Return of the forgotten hero: the role of Y chromosome-encoded Zfy in male reproduction. Mol Hum Reprod. 2023 07 31; 29(8). PMID: 37354519.
      Citations: 2     Fields:    Translation:HumansAnimalsCells
    5. Giannakopoulos S, Strange DP, Jiyarom B, Abdelaal O, Bradshaw AW, Nerurkar VR, Ward MA, Bakse J, Yap J, Vanapruks S, Boisvert WA, Tallquist MD, Shikuma C, Sadri-Ardekani H, Clapp P, Murphy SV, Verma S. In vitro evidence against productive SARS-CoV-2 infection of human testicular cells: Bystander effects of infection mediate testicular injury. PLoS Pathog. 2023 05; 19(5):e1011409. PMID: 37200377.
      Citations: 8     Fields:    Translation:HumansAnimalsCells
    6. Batdorj E, AlOgayil N, Zhuang QK, Galvez JH, Bauermeister K, Nagata K, Kimura T, Ward MA, Taketo T, Bourque G, Naumova AK. Genetic variation in the Y chromosome and sex-biased DNA methylation in somatic cells in the mouse. Mamm Genome. 2023 03; 34(1):44-55. PMID: 36454369.
      Citations: 4     Fields:    Translation:AnimalsCells
    7. Giannakopoulos S, Strange DP, Jiyarom B, Abdelaal O, Bradshaw AW, Nerurkar VR, Ward MA, Bakse J, Yap J, Vanapruks S, Boisvert W, Tallquist MD, Shikuma C, Sadri-Ardekani H, Clapp P, Murphy S, Verma S. In vitro evidence against productive SARS-CoV-2 infection of human testicular cells: Bystander effects of infection mediate testicular injury. bioRxiv. 2022 Sep 22. PMID: 36172118.
      Citations:    
    8. Holmlund H, Yamauchi Y, Durango G, Fujii W, Ward MA. Two acquired mouse Y chromosome-linked genes, Prssly and Teyorf1, are dispensable for male fertility?. Biol Reprod. 2022 09 12; 107(3):752-764. PMID: 35485405.
      Citations: 6     Fields:    Translation:AnimalsCells
    9. Hirose N, Blanchet G, Yamauchi Y, Snow AC, Friedman R, Khoo CY, Lary CW, Ward MA, Nadeau JH. Genetically-biased fertilization in APOBEC1 complementation factor (A1cf) mutant mice. Sci Rep. 2022 08 10; 12(1):13599. PMID: 35948620.
      Citations: 1     Fields:    Translation:AnimalsCells
    10. Yamauchi Y, Matsumura T, Bakse J, Holmlund H, Blanchet G, Carrot E, Ikawa M, Ward MA. Loss of mouse Y chromosome gene Zfy1 and Zfy2 leads to spermatogenesis impairment, sperm defects, and infertility. Biol Reprod. 2022 06 13; 106(6):1312-1326. PMID: 35293998.
      Citations: 9     Fields:    Translation:AnimalsCells
    11. Wang Y, Yamauchi Y, Wang Z, Zheng H, Yanagimachi R, Ward MA, Yan W. Both Cauda and Caput Epididymal Sperm Are Capable of Supporting Full-Term Development in FVB and CD-1 Mice. Dev Cell. 2020 12 21; 55(6):675-676. PMID: 33352141.
      Citations: 11     Fields:    Translation:AnimalsCells
    12. Moretti C, Blanco M, Ialy-Radio C, Serrentino ME, Gob? C, Friedman R, Battail C, Leduc M, Ward MA, Vaiman D, Tores F, Cocquet J. Battle of the Sex Chromosomes: Competition between X and Y Chromosome-Encoded Proteins for Partner Interaction and Chromatin Occupancy Drives Multicopy Gene Expression and Evolution in Muroid Rodents. Mol Biol Evol. 2020 12 16; 37(12):3453-3468. PMID: 32658962.
      Citations: 18     Fields:    Translation:AnimalsCells
    13. Tsai PS, Yamauchi Y, Riel JM, Ward MA. Pregnancy environment, and not preconception, leads to fetal growth restriction and congenital abnormalities associated with diabetes. Sci Rep. 2020 07 23; 10(1):12254. PMID: 32703993.
      Citations: 6     Fields:    Translation:HumansAnimalsPHPublic Health
    14. Yamauchi Y, Ajduk A, Ward MA. Oocytes from female mice on MF1 genetic background are not suitable for assisted reproduction?. Biol Reprod. 2020 03 13; 102(3):521-523. PMID: 31837135.
      Citations: 1     Fields:    Translation:AnimalsCells
    15. Ortega EA, Salvador Q, Fernandez M, Ward MA. Alterations of sex determination pathways in the genital ridges of males with limited Y chromosome genes?. Biol Reprod. 2019 03 01; 100(3):810-823. PMID: 30285093.
      Citations: 1     Fields:    Translation:AnimalsCells
    16. Riel JM, Yamauchi Y, Ruthig VA, Malinta QU, Blanco M, Moretti C, Cocquet J, Ward MA. Rescue of Sly Expression Is Not Sufficient to Rescue Spermiogenic Phenotype of Mice with Deletions of Y Chromosome Long Arm. Genes (Basel). 2019 02 12; 10(2). PMID: 30759861.
      Citations: 8     Fields:    Translation:AnimalsCells
    17. Ward MA, Ward WS. Contributions of Ryuzo Yanagimachi to the field of reproductive biology. Biol Reprod. 2019 01 01; 100(1):1-7. PMID: 30657895.
      Citations:    Fields:    Translation:HumansAnimals
    18. Balaan C, Corley MJ, Eulalio T, Leite-Ahyo K, Pang APS, Fang R, Khadka VS, Maunakea AK, Ward MA. Juvenile Shank3b deficient mice present with behavioral phenotype relevant to autism spectrum disorder. Behav Brain Res. 2019 01 01; 356:137-147. PMID: 30134148.
      Citations: 14     Fields:    Translation:Animals
    19. Ward MA, Yanagimachi R. Intracytoplasmic Sperm Injection in Mice. Cold Spring Harb Protoc. 2018 01 02; 2018(1). PMID: 29295903.
      Citations: 5     Fields:    Translation:AnimalsCells
    20. Ward MA. An Interview with Dr Marisa Bartolomei. Biol Reprod. 2018 02; 98(2):147-149. PMID: 29228104.
      Citations:    Fields:    
    21. Ward MA. An interview with Professor Mitinori Saitou. Biol Reprod. 2018 02; 98(2):150-152. PMID: 29228095.
      Citations:    Fields:    
    22. Ruthig VA, Nielsen T, Riel JM, Yamauchi Y, Ortega EA, Salvador Q, Ward MA. Testicular abnormalities in mice with Y chromosome deficiencies. Biol Reprod. 2017 03 01; 96(3):694-706. PMID: 28339606.
      Citations: 6     Fields:    Translation:Animals
    23. Ruthig VA, Labrash S, Lozanoff S, Ward MA. Macroscopic demonstration of the male urogenital system with evidence of a direct inguinal hernia utilizing room temperature plastination. Anatomy. 2016 Dec; 10(3):211-220. PMID: 28824276.
      Citations:    
    24. Yamauchi Y, Riel JM, Ruthig VA, Ortega EA, Mitchell MJ, Ward MA. Two genes substitute for the mouse Y chromosome for spermatogenesis and reproduction. Science. 2016 Jan 29; 351(6272):514-6. PMID: 26823431.
      Citations: 29     Fields:    Translation:AnimalsCells
    25. Yamauchi Y, Riel JM, Ruthig V, Ward MA. Mouse Y-Encoded Transcription Factor Zfy2 Is Essential for Sperm Formation and Function in Assisted Fertilization. PLoS Genet. 2015 Dec; 11(12):e1005476. PMID: 26719889.
      Citations: 22     Fields:    Translation:AnimalsCells
    26. Ortega EA, Ruthig VA, Ward MA. Sry-Independent Overexpression of Sox9 Supports Spermatogenesis and Fertility in the Mouse. Biol Reprod. 2015 Dec; 93(6):141. PMID: 26536904.
      Citations: 6     Fields:    Translation:AnimalsCells
    27. Sato BL, Ward MA, Astern JM, Kendal-Wright CE, Collier AC. Validation of murine and human placental explant cultures for use in sex steroid and phase II conjugation toxicology studies. Toxicol In Vitro. 2015 Feb; 29(1):103-12. PMID: 25283089.
      Citations: 10     Fields:    Translation:HumansAnimals
    28. Collier AC, Yamauchi Y, Sato BL, Roug?e LR, Ward MA. UDP-glucuronosyltransferase 1a enzymes are present and active in the mouse blastocyst. Drug Metab Dispos. 2014 Nov; 42(11):1921-5. PMID: 25200869.
      Citations: 2     Fields:    Translation:Animals
    29. Sato BL, Sugawara A, Ward MA, Collier AC. Single blastomere removal from murine embryos is associated with activation of matrix metalloproteinases and Janus kinase/signal transducers and activators of transcription pathways of placental inflammation. Mol Hum Reprod. 2014 Dec; 20(12):1247-57. PMID: 25180268.
      Citations: 2     Fields:    Translation:AnimalsCells
    30. Vernet N, Mahadevaiah SK, Yamauchi Y, Decarpentrie F, Mitchell MJ, Ward MA, Burgoyne PS. Mouse Y-linked Zfy1 and Zfy2 are expressed during the male-specific interphase between meiosis I and meiosis II and promote the 2nd meiotic division. PLoS Genet. 2014 Jun; 10(6):e1004444. PMID: 24967676.
      Citations: 32     Fields:    Translation:AnimalsCells
    31. Comptour A, Moretti C, Serrentino ME, Auer J, Ialy-Radio C, Ward MA, Tour? A, Vaiman D, Cocquet J. SSTY proteins co-localize with the post-meiotic sex chromatin and interact with regulators of its expression. FEBS J. 2014 Mar; 281(6):1571-84. PMID: 24456183.
      Citations: 29     Fields:    Translation:AnimalsCells
    32. Yamauchi Y, Riel JM, Stoytcheva Z, Ward MA. Two Y genes can replace the entire Y chromosome for assisted reproduction in the mouse. Science. 2014 Jan 03; 343(6166):69-72. PMID: 24263135.
      Citations: 50     Fields:    Translation:HumansAnimalsCells
    33. Gawecka JE, Marh J, Ortega M, Yamauchi Y, Ward MA, Ward WS. Mouse zygotes respond to severe sperm DNA damage by delaying paternal DNA replication and embryonic development. PLoS One. 2013; 8(2):e56385. PMID: 23431372.
      Citations: 65     Fields:    Translation:AnimalsCells
    34. Riel JM, Yamauchi Y, Sugawara A, Li HY, Ruthig V, Stoytcheva Z, Ellis PJ, Cocquet J, Ward MA. Deficiency of the multi-copy mouse Y gene Sly causes sperm DNA damage and abnormal chromatin packaging. J Cell Sci. 2013 Feb 01; 126(Pt 3):803-13. PMID: 23178944.
      Citations: 16     Fields:    Translation:HumansAnimalsCells
    35. Sugawara A, Ward MA. Biopsy of embryos produced by in vitro fertilization affects development in C57BL/6 mouse strain. Theriogenology. 2013 Jan 15; 79(2):234-41. PMID: 23174776.
      Citations:    Fields:    Translation:AnimalsCells
    36. Sugawara A, Sato B, Bal E, Collier AC, Ward MA. Blastomere removal from cleavage-stage mouse embryos alters steroid metabolism during pregnancy. Biol Reprod. 2012 Jul; 87(1):4, 1-9. PMID: 22517623.
      Citations: 9     Fields:    Translation:HumansAnimalsCells
    37. Ward MA. Wild-derived inbred mice no longer ART-resistant. Biol Reprod. 2012 May; 86(5):168, 1-2. PMID: 22423046.
      Citations: 1     Fields:    Translation:Animals
    38. Sugawara A, Pearson BL, Blanchard DC, Ward MA. Mouse females devoid of exposure to males during fetal development exhibit increased maternal behavior. Psychoneuroendocrinology. 2012 Mar; 37(3):383-95. PMID: 21803500.
      Citations: 3     Fields:    Translation:AnimalsPHPublic Health
    39. Riel JM, Yamauchi Y, Huang TT, Grove J, Ward MA. Short-term storage of human spermatozoa in electrolyte-free medium without freezing maintains sperm chromatin integrity better than cryopreservation. Biol Reprod. 2011 Sep; 85(3):536-47. PMID: 21593474.
      Citations: 8     Fields:    Translation:HumansAnimalsCells
    40. Yamauchi Y, Riel JM, Ward MA. Paternal DNA damage resulting from various sperm treatments persists after fertilization and is similar before and after DNA replication. J Androl. 2012 Mar-Apr; 33(2):229-38. PMID: 21546611.
      Citations: 20     Fields:    Translation:AnimalsCells
    41. Raunig JM, Yamauchi Y, Ward MA, Collier AC. Assisted reproduction technologies alter steroid delivery to the mouse fetus during pregnancy. J Steroid Biochem Mol Biol. 2011 Aug; 126(1-2):26-34. PMID: 21193037.
      Citations: 17     Fields:    Translation:Animals
    42. Cocquet J, Ellis PJ, Yamauchi Y, Riel JM, Karacs TP, Rattigan A, Ojarikre OA, Affara NA, Ward MA, Burgoyne PS. Deficiency in the multicopy Sycp3-like X-linked genes Slx and Slxl1 causes major defects in spermatid differentiation. Mol Biol Cell. 2010 Oct 15; 21(20):3497-505. PMID: 20739462.
      Citations: 41     Fields:    Translation:AnimalsCells
    43. Yamauchi Y, Riel JM, Stoytcheva Z, Burgoyne PS, Ward MA. Deficiency in mouse Y chromosome long arm gene complement is associated with sperm DNA damage. Genome Biol. 2010; 11(6):R66. PMID: 20573212.
      Citations: 22     Fields:    Translation:AnimalsCells
    44. Cocquet J, Ellis PJ, Yamauchi Y, Mahadevaiah SK, Affara NA, Ward MA, Burgoyne PS. The multicopy gene Sly represses the sex chromosomes in the male mouse germline after meiosis. PLoS Biol. 2009 Nov; 7(11):e1000244. PMID: 19918361.
      Citations: 105     Fields:    Translation:AnimalsCells
    45. Yamauchi Y, Ward MA, Ward WS. Asynchronous DNA replication and origin licensing in the mouse one-cell embryo. J Cell Biochem. 2009 May 15; 107(2):214-23. PMID: 19301258.
      Citations: 13     Fields:    Translation:AnimalsCells
    46. Yamauchi Y, Riel JM, Wong SJ, Ojarikre OA, Burgoyne PS, Ward MA. Live offspring from mice lacking the Y chromosome long arm gene complement. Biol Reprod. 2009 Aug; 81(2):353-61. PMID: 19420387.
      Citations: 23     Fields:    Translation:AnimalsCells
    47. Collier AC, Miyagi SJ, Yamauchi Y, Ward MA. Assisted reproduction technologies impair placental steroid metabolism. J Steroid Biochem Mol Biol. 2009 Aug; 116(1-2):21-8. PMID: 19406239.
      Citations: 29     Fields:    Translation:Animals
    48. Riel JM, Huang TT, Ward MA. Freezing-free preservation of human spermatozoa--a pilot study. Arch Androl. 2007 Sep-Oct; 53(5):275-84. PMID: 18309900.
      Citations: 3     Fields:    Translation:HumansAnimalsCells
    49. Yamauchi Y, Doe B, Ajduk A, Ward MA. Genomic DNA damage in mouse transgenesis. Biol Reprod. 2007 Nov; 77(5):803-12. PMID: 17652664.
      Citations: 16     Fields:    Translation:AnimalsCells
    50. Yamauchi Y, Ajduk A, Riel JM, Ward MA. Ejaculated and epididymal mouse spermatozoa are different in their susceptibility to nuclease-dependent DNA damage and in their nuclease activity. Biol Reprod. 2007 Oct; 77(4):636-47. PMID: 17596560.
      Citations: 12     Fields:    Translation:AnimalsCells
    51. Yamauchi Y, Ward MA. Preservation of ejaculated mouse spermatozoa from fertile C57BL/6 and infertile Hook1/Hook1 mice collected from the uteri of mated females. Biol Reprod. 2007 Jun; 76(6):1002-8. PMID: 17314312.
      Citations: 9     Fields:    Translation:AnimalsCells
    52. Ajduk A, Yamauchi Y, Ward MA. Sperm chromatin remodeling after intracytoplasmic sperm injection differs from that of in vitro fertilization. Biol Reprod. 2006 Sep; 75(3):442-51. PMID: 16775225.
      Citations: 46     Fields:    Translation:AnimalsCells
    53. Ward MA, Burgoyne PS. The effects of deletions of the mouse Y chromosome long arm on sperm function--intracytoplasmic sperm injection (ICSI)-based analysis. Biol Reprod. 2006 Apr; 74(4):652-8. PMID: 16354792.
      Citations: 26     Fields:    Translation:HumansAnimalsCells
    54. Ward MA. Intracytoplasmic sperm injection effects in infertile azh mutant mice. Biol Reprod. 2005 Jul; 73(1):193-200. PMID: 15800178.
      Citations: 10     Fields:    Translation:AnimalsCells
    55. Ward MA, Ward WS. A model for the function of sperm DNA degradation. Reprod Fertil Dev. 2004; 16(5):547-54. PMID: 15355763.
      Citations: 15     Fields:    Translation:HumansCells
    56. Ward MA, Kaneko T, Kusakabe H, Biggers JD, Whittingham DG, Yanagimachi R. Long-term preservation of mouse spermatozoa after freeze-drying and freezing without cryoprotection. Biol Reprod. 2003 Dec; 69(6):2100-8. PMID: 12930716.
      Citations: 30     Fields:    Translation:AnimalsCells
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