Professor Kate Loveland, Research Group Head, Testis Development and Male Germ Cell Biology and Hudson Institute

Professor Kate Loveland, Senior Principal Research Fellow

Professor Loveland received her undergraduate and PhD degrees at Duke University in the United States, studying mammalian fertilization. Her laboratory based in Hudson Institute investigates how events in fetal and postnatal life affect adult male fertility.  Her team has identified new mechanisms that underpin mammalian testis development and sperm production, including the molecular switches that regulate cell fate decisions in sperm precursor cells (germ cells) and in the somatic cells that support them.  Her longstanding interest in signalling by activin/ TGFβ superfamily underpins the current work addressing how exposure to environmental factors, such as medications and endocrine disruptor chemicals, underpin testicular dysgenesis and the risk of testicular tumours in young men.  How this complicated signalling family is regulated to generate normal and pathological outcomes is a key feature of her interests.  This extends to include studies evaluating how the proteins that are responsible for transporting factors into the cell nucleus work to control cellular development and stress responses.

She held a National Health and Medical Research Council of Australia Senior Research Fellowship (2000-2020) and continuous competitive research funding for over 30 years. She received the American Society for Andrology Young Andrologist Award in 2004, the Monash University Vice Chancellor’s Award for Excellence in Postgraduate Supervision in 2010, an Honorary Liebig Professorship from the Justus-Liebig University in Giessen, Germany for her joint role in leading the establishment of an International PhD Research Training Group in the Pathogenesis of Male Reproduction (2014-2022), and the inaugural Society for the Study of Reproduction (SSR) Fuller W Bazer International Scientist Award in 2018.

Prof Loveland also serves at the Head of Graduate Research for the Monash University School of Clinical Sciences.  Professional roles for her discipline include Society for Reproductive Biology: Program Chair, and Treasurer; American Society for Andrology: President (2023-2024), Major Awards Committee Chair, Annual Meeting Program co-Chair (2010), Founder and Chair of the Basic Science Workshop, President of Women in Andrology and Council member; Society for the Study of Reproduction: International Members Subcommittee Chair, Director, and Annual Meeting Program co-Chair.

Prof Loveland has published over 175 peer-reviewed manuscripts, and she is currently serving as an Associate and Guest Editor for Andrology. She has supervised over 30 PhD students and over 30 Honours and Masters students to successful completion of the degree.

Selected publications

  • Reviews

    O’Donnell L, Whiley PAF, Loveland KL (2022) Activin A and Sertoli Cells:  Key to fetal testis steroidogenesis.  Frontiers in Endocrinology (Reproduction). 24 May 2022. Mini Review article.

  • Nathaniel B, Whiley PAF, Miyamoto Y, Loveland KL (2021) Importins: Diverse roles in male fertility. Semin Cell Dev Biol. Aug 20;S1084-9521(21)00206-8.

  • Loveland KL, Klein B, Pueschl D, Indumathy S, Bergmann M, Loveland BE, Hedger MP, Schuppe HC (2017) Cytokines in male fertility and reproductive pathologies:  immunoregulation and beyond. Front Endocrinol (Lausanne) 8:307.

  • Loveland KL, Hedger MP (2015) Activins and inhibins in Sertoli cell biology: Implications for testis development and function. In Sertoli Cell Biology. Griswold M ed.  Elsevier Science (USA). pp 201-221.

  • Loveland K, Major A, Butler R, Young J, Jans DA, Miyamoto Y (2015) Putting things in place for fertilization: Discovering roles for importin proteins in cell fate and spermatogenesis. Asian Journal of Andrology 17(4):537-544.

  • De Kretser DM, O’Bryan MK, Loveland KL (2015) Spermatogenesis. In: eds Jameson/DeGroot: Endocrinology: Adult & Pediatric, 7/E. Elsevier.

  • McIver SC, Roman SD, Nixon B, Loveland KL, McLaughlin EA (2013) The rise of testicular germ cell tumours: the search for causes, risk factors and novel therapeutic targets. F1000 Research 2:55.

  • Itman C, Loveland KL (2013) Smads and cell fate: distinct roles in specification, development and tumourigenesis in the testis. IUBMB Life 65(2):85-97.

  • Barakat B, Itman C, Mendis S, Loveland KL (2012) Activins and inhibins in mammalian testis development: new models, new insights. Invited review. Molecular and Cellular Endocrinology 359:66-77.

  • Miyamoto Y, Boag P, Hime G, Loveland KL (2012) Regulated nucleocytoplasmic transport during gametogenesis. BBA – Gene Regulatory Mechanisms 1819:616-30.

  • Primary Data Publications

    Radhakrishnan K, Luu M, Iaria J, Sutherland JM, McLaughlin EA, Zhu H-J, Loveland KL (2023) Activin and BMP signalling in human testicular cancer cell lines, and a role for the neucleocytoplasmmic transport protein Importin-5 in their crosstalk. Cell Signaling. Accepted for publication on 9 March 2023.


    Hosseini S, Moody SC, Fietz D, Indumathy S, Schuppe H-C, Hedger MP, Loveland KL (2022) The changing landscape of immune cells in the fetal mouse testis. Histochemistry and Cell Biology. Published online 12 July 2022. doi/org/10.1007/s00418-022-02129-6.

  • Moody SC, Whiley PAF, Western PS, Loveland KL (2022) The impact of activin A on fetal gonocytes: chronic versus acute exposure outcomes. Front Endocrinol (Lausanne) May 31;13:896747.

  • Young JC, Kerr G, Micati DJ, Nielsen JE, Rajpert-DeMeyts, Abud HE*, Loveland KL* (2020) WNT signaling in the normal human adult testis and in male germ cell neoplasms. Human Reproduction. Jul 15:deaa150. Online ahead of print. *Equal senior co-authors.

  • Moody SC, Wakitani S, Young JC, Western PS, Loveland KL (2020) Evidence that activin A directly modulates early human male germline differentiation status. Reproduction. Jul;160(1):141-154.

  • Whiley P, O’Donnell L, Moody SC, Handelsman DJ, Young JC, Richards EA, Almstrup K, Western PS, Loveland KL (2020) Activin A determines steroid levels and composition in the fetal testis. Endocrinology. Jul 1;161(7):bqaa058.

  • Micati DJ, Radhakrishnan K, Young JC, Rajpert-de Meyts E, Hime GR, Abud HE*, Loveland KL* (2020) Snail factors in testicular germ cell tumours and their regulation by the BMP4 signaling pathway. Andrology. May 22. doi: 10.1111/andr.12823. *Equal senior co-authors.

  • Indumathy S, Pueschl D, Klein B, Fietz D, Bergmann M, Schuppe HC, Da Silva N, Loveland BE, Hickey MJ, Hedger MP, Loveland KL (2020) Testicular immune cell populations and macrophage polarisation in adult male mice and the influence of altered activin A levels.  J Reprod Immunol Nov; 142:103204.

  • Major AT, Hogarth C, Young J, Kurihara Y, Jans DA, Loveland KL (2019) Dynamic paraspeckle component localization during spermatogenesis.  Reproduction July Sep;158(3):267-280.

  • Lepletier A, Hun, ML, Hammett MV, Hedger MP, Loveland KL, Chidgey AP (2019)  Interplay between Follistatin, Activin A and BMP4 signaling regulates postnatal thymic epithelial progenitor cell differentiation during aging. Cell Reports 27:3887-3901.

  • Klein B, Pant S, Bushan S, Kautz J, Rudat C, Kispert A, Pilatz A, Wijayarathna R, Middendorff R, Loveland KL, Hedger MP, Meinhardt A (2019) Dexamethasone improves therapeutic outcomes in a preclinical bacterial epididymitis mouse model. Hum Reprod Jul 8;34(7):1195-1205.


  • Szarek M, Bergmann M, Konrad L, Schuppe HC, Kliesch S, Hedger MP, Loveland KL (2018) Activin A target genes are differentially expressed between normal and neoplastic adult human testes:  clues to gonocyte fate choice.  Andrology Oct 13. doi: 10.1111/andr.12553.

  • Miyamoto Y, Whiley PAF, Goh HY, Wong C, Higgins G, Tachibana T, McMenamin P, Mayne L, Loveland KL (2018) The Stk35 locus contributes to normal gametogenesis and encodes a IncRNA responsive to oxidative stress. Biol Open Jul 3 pii:bio 032631. doi: 10.1242/bio.032631.

  • Micati DJ, Hime GR, McLaughlin EA, Abud HE, Loveland KL (2018) Differential expression profiles of conserved Snail transcription factors in the mouse testis. Andrology 6(2):362-373.

  • Major AT, Miyanoto Y, Jans DA, Loveland KL (2017) Development of a pipeline for automated, high-throughput analysis of paraspeckle proteins reveals specific roles for importin α proteins. Sci Rep 7, Article number: 43323. doi: 10.1038/srep43323.

  • Klein B, Schuppe HC, Bergmann M, Hedger MP, Loveland BE, Loveland KL (2017) An in vitro model demonstrates the potential of neoplastic human germ cells to influence the tumour microenvironment. Andrology  5(4):763-770.

  • Itman C, Bielanowicz A, Goh H, Lee Q, Fulcher AJ, Moody SC, Doery JCG, Martin J, Eyre S, Hedger MP, Loveland KL (2015) Murine inhibin alpha subunit haploinsufficiency causes transient abnormalities in prepubertal testis development followed by adult testicular decline. Endocrinology 156(6):2254-2268.

  • Major AT, Hogarth CA, Miyamoto Y, Sarraj MA, Smith CL, Koopman P, Kurihara Y, Jans DA, Loveland KL (2015) Specific interaction with the nuclear transporter importin alpha 2 can modulate paraspeckle protein 1 delivery to nuclear paraspeckles. Biol Cell 26:1543-1558.

  • Arjomand A, Baker MA, Li C, Buckle AM, Jans DA, *Loveland KL, *Miyamoto Y. (2014) The α-importome of mammalian germ cell maturation provides novel insights for importin biology. FASEB J 28(8):3480-93. * Equal contributors. ^Primary supervisor of first author.

  • Jørgensen A, Young J, Nielsen JE, Joensen UN, Toft BG, *Rajpert-De Meyts E, *Loveland KL (2014). Hanging drop cultures of human testis and testis cancer samples used to investigate activin treatment effects in a preserved niche. * Equal contributors.  Br J Cancer 110(10):2604-2014.

  • Sahin Z, Szezepny A, McLaughlin EA, Meistrich ML, Zhou W, Unstunel I, Loveland, KL (2014) Dynamic Hedgehog signalling pathway activity in germline stem cells. Andrology 2(2):267-274.

  • Kerr G, Young, J, Horvay K, Abud H, Loveland KL (2014) Intrinsic regulation of Wnt/beta-catenin signaling is required for adult spermatogenesis. Biology of Reproduction. 90(1):3. doi: 10.1095/biolreprod.112.105809.

  • Moody S, Goh H, Loveland KL, Itman, C. (2013) Prepubertal mouse testis growth and maturation and androgen production are acutely sensitive to di-n-butyl phthalate. Endocrinology 154(9):3460-3475.

  • Young JC, Ly-Hyunh J, Lescesen H, Miyamoto Y, Browne C, Yoneda Y, Koopman P, Loveland KL*, Jans DA* (2013) The nuclear import factor importin a4 can protect against oxidative stress. Biochim Biophys Acta – Molecular Cell Research 1833(10):2348-2356. * equal contributors.

  • Tarulli GA, Stanton PG, LovelandKL, Rajpert-De Meyts E, McLachlan RI,  Meachem SJ (2013) A survey of Sertoli cell differentiation in men after gonadotropin suppression and in testicular cancer. Spermatogenesis 3:1-7.