MRC Centre for Reproductive Health
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Professor Lee Smith group - recent publications

Publication list.


Livigni A, O’Hara L, Polak ME, Angus T, Smith LB and Freeman TC (2018) A Graphical and Computational Modelling Platform for Biological Pathways Nature Protocols (In Press)

Neirjinck Y Calvel P, Kilcoyne KR, Kühne F, Stévant I, Griffeth RJ, Pitetti JL, Andric S, Hu MC, Pralong F, Smith LB,  and Nef S. (2018) Insulin and IGF1 receptors are essential for the development and steroidogenic function of adult Leydig cells The FASEB Journal (In Press)

Stévant I, Neirjinck Y, Borel C, Escoffier J, Smith LB, Antonarakis SE, Dermitzakis ET and Nef S (2018) Deciphering cell lineage specification during male sex determination with single-cell RNA sequencing Cell Reports (In Press)

Smith LB and McEwan I (2018) Androgen Receptor, In Encyclopedia of Reproduction - Second Edition. Eds Skinner M, Jegou B, Elsevier Press. (In Press)

Rebourcet D, Darbey A, Curley M, O’Shaughnessy PJ and Smith LB (2018) Testicular Cell Selective Ablation Using Diphtheria Toxin Receptor Transgenic Mice. In Sertoli Cells: Methods and Protocols; Methods Molecular Biology, Vol. 1748, Marco G. Alves and Pedro F. Oliveira (Eds): Springer Press (In Press)


Hutka M, Smith LB and Mitchell RT (2017) Xenotransplantation as a model for human testicular development Differentiation 97:44-53

Patel SH, O'Hara L, Atanassova N, Smith SE, Curley MK, Rebourcet D, Darbey AL, Gannon A-L, Sharpe RM, Smith LB (2017) Low-dose tamoxifen treatment in juvenile males has long-term adverse effects on the reproductive system: implications for inducible transgenics. Scientific Reports doi:10.1038/s41598-017-09016-4

Takov K, Wu J, Denvir MA, Smith LB, Hadoke PWF (2017) The role of Androgen Receptors in Atherosclerosis. Molecular and Cellular Endocrinology

Engels M, Span PN, Mitchell RT, Heuvel JJTM, Marijnissen-van Zanten MA, van Herwaarden AE, Hulsbergen-van de Kaa CA, Oosterwijk E, Stikkelbroeck NM, Smith LB, Sweep FCGJ, and Claahsen-van der Grinten HL (2017) GATA transcription factors in testicular adrenal rest tumours Endocrine Connections 6:866-875

Soffientini U, Rebourcet D, Abel MH, Lee S, Hamilton G, Fowler PA, Smith LB, O'Shaughnessy PJ (2017) Identification of Sertoli cell-specific transcripts in the mouse testis and the role of FSH and androgen in the control of Sertoli cell activity. BMC Genomics 18(1):15 Dec 2017

Darbey A and Smith LB (2017) Deliverable transgenics & gene therapy possibilities for the testes Molecular and Cellular Endocrinology

Rebourcet D, Darbey A, Monteiro A, Soffientini U, Tsai YT, Handel I, Pitetti JL, Nef S, Smith LB, O'Shaughnessy PJ (2017). Sertoli Cell Number Defines and Predicts Germ and Leydig Cell Population Sizes in the Adult Mouse Testis Endocrinology 158(9):2955-2969

O’Hara L and Smith LB. (2017) The Genetics of Androgen Receptor Signalling In Male Fertility. In The Genetics of Male and Female Infertility; Monographs in Human Genetics Series. 21:86-100Eds Vogt PH and Schmid M (Karger)

O’Hara L and Smith LB. (2017) Mouse Genetics: how does it inform male fertility research? In The Sperm Cell 2nd Edition; Eds de Jonge C and Barratt C. FRONT COVER IMAGE

Mitchell RT, O’Hara L and Smith LB. (2017) Gonadotropin and Steroid Hormone Control of Spermatogonial Differentiation. Pp 147-178 In The Biology of Mammalian Spermatogonia. Eds: Oatley J and Griswold M. 

Nolan P, Banks G, Lassi G, Tinarelli F, Simon M, Lawson T, Johnson S, Sweeting S, Chesham J, Barnard A, Horner N, Westerberg H, Smith LB, Hastings M, Hirst R, Tucci V, Hoerder-Suabedissen A, Wilcox A, Lau P, Molnár Z, and Rutman A. (2017) A missense mutation in Katnal1 underlies behavioural, neurological and ciliary anomalies.  Molecular Psychiatry (In Press)

Ribeiro CM, Ferreira LG, Thimoteo DS, Smith LB, Hinton BT, Avellar MC. (2017) Novel androgen-induced activity of an antimicrobial β-defensin: Regulation of Wolffian duct morphogenesis. Mol Cell Endocrinol. 15;442:142-152. 


Cobice D, Livingstone D, Mackay C, Goodwin R, Smith LB, Walker B, Andrew R. (2016) Spatial Localization and Quantitation of Androgens in Mouse Testis by Mass Spectrometry Imaging. Analytical Chemistry (In Press).

O’Hara L, Livigni A, Theo T, Boyer B, Angus T, Wright D, Chen S-H, Raza S, Barnett MW, Digard P, Smith LB, and Freeman TC. (2016) Modelling the structure and dynamics of biological pathways PLoS Biology 14(8):e1002530.

Rebourcet D, Wu J, Cruickshanks L, Smith SE, Milne L, Fernando A, Wallace RJ, Gray CD, Hadoke PWF, Mitchell RT, O’Shaughnessy PJ and Smith LB (2016) Sertoli cells modulate testicular vascular network development, structure and function to influence circulating testosterone concentrations in adult male mice. Endocrinology doi:10.1210/en.2016-1156

Stanton PG, Foo CFH, Rainczuk A, Stephens AN, Condina M, O’Donnell L, Weidner W, Ishikawa T, Cruickshanks L, Smith LB, McLachlan RI. (2016) Mapping The Testicular Interstitial Fluid Proteome From Normal Rats. Proteomics 16(17):2391-402.

Wu J, Hadoke PWF, Takov K, Denvir MA and Smith LB (2016) Influence of androgen receptor in vascular cells on reperfusion following hindlimb ischaemia. PLoS ONE 11(5): e0154987.

Zhu D, Hadoke PWF, Wu J, Vesey A, Lerman D, Dweck M, Newby D, Smith LB and MacRae V. (2016) Ablation of the androgen receptor from vascular smooth muscle cells demonstrates a role for testosterone in vascular calcification. Scientific Reports doi:10.1038/srep24807

Macdonald J*, Smith LB*, Anderson RA and Mitchell RT (2016) Spatiotemporal profiling of luteinising hormone/human choriogonadotropin receptor in the human fetal testis The Lancet 387:S74

Smith LB (2016) Non-Classical Testosterone Signalling: A New Pathway Controlling Spermatogenesis? Biology of Reproduction.  biolreprod.115.137950

O’Hara L and Smith LB. (2016) The Genetics of Androgen Receptor Signalling In Male Fertility. In The Genetics of Male and Female Infertility; Monographs in Human Genetics Series. Eds Vogt PH and Schmid M (Karger) (In Press).

Mitchell RT, O’Hara L and Smith LB. (2016) Gonadotropin and Steroid Hormone Control of Spermatogonial Differentiation. In The Biology of Mammalian Spermatogonia. Eds: Oatley J and Griswold M. (In Press)

O’Hara L and Smith LB. (2016) Development and Characterisation of Cell-Specific Androgen Receptor Knockout Mice. In Methods in Molecular Biology – The Nuclear Receptor Superfamily. Ed. McEwan IJ. 1443:219-48.


Smith LB, O’Shaughnessy PJ and Rebourcet D. Testicular cell ablation models: what have we learned? Invited Review: Andrology DOI: 10.1111/andr.12107 JOURNAL FRONT COVER

O’Hara L and Smith LB. Mouse Genetics: how does it inform male fertility research? In The Sperm Cell 2nd Edition; Eds de Jonge C and Barratt C (In Press).

Chambers T and Smith LB. Hormone Imbalances in Men. Nursing in Practice 85:74-76

Guillermet-Guibert J, Smith LB, Halet G, Pearce W, Whitehead MA, Rebourcet D, León K, Crépieux P, Nock G, Strömstedt M, Enerback M, Chelala C, Graupera M, Carroll J, Cosulich S, Saunders PTK, Huhtaniemi I, Vanhaesebroeck B. The p110β PI 3-kinase is a Sertoli cell-specific integrator of androgen receptor signalling in sperm development. PLoS Genetics. DOI: 10.1371/journal.pgen.1005304

Dixit KCS, Wu J, Smith LB, Hadoke PWF and Wu FCW: Endotext: Chapter 16: Androgens and Coronary Artery Disease.

Van den Driesche S, Macdonald J, Anderson RA, Johnston ZC, Chetty T, Smith LB, Mckinnell C, Dean A, Homer NZ, Jorgensen A, Camacho-Moll ME, Sharpe RM, and Mitchell RT (2015) Prolonged exposure to acetaminophen reduces testosterone production by the human fetal testis in a xenograft model. Science Translational Medicine. Vol. 7, Issue 288, p. 288ra80 (WIDESPREAD PRESS COVERAGE)

Willems A, Roesl C, Mitchell RT, Milne L, Jeffery N, Smith S, Verhoeven G, Brown P and Smith LB. (2015) Sertoli Cell Androgen Receptor Signalling in Adulthood is Essential for Post-Meiotic Germ Cell Development. Molecular Reproduction and Development DOI: 10.1002/mrd.22506. JOURNAL FRONT COVER

O’Hara L, Curley M, Tedim Ferreira M, Cruickshanks L, Milne L, and Smith LB. (2015) Pituitary Androgen Receptor Signalling Regulates Prolactin But Not Gonadotrophins In The Male Mouse. PLoS ONE DOI: 10.1371/journal.pone.0121657

O’Hara L and Smith LB. (2015) Androgen Receptor Roles in Spermatogenesis and Infertility.  Best Practice and Research Clinical Endocrinology and Metabolism  doi:10.1016/j.beem.2015.04.006

O’Hara L, McInnes K, Simitsidellis I, Morgan S, Atanassova N, Slowikowska-Hilczer J, Kula K, Szarras-Czapnik M, Milne L, Mitchell RT, and Smith LB. (2015) Autocrine Androgen Action is Essential for Leydig Cell Maturation and Function, and Protects Against Late-Onset Leydig Cell Apoptosis in Both Mice and Men. The FASEB Journal. Mar;29(3):894-910

Fijak M, Sauber LJ, Wenzel JP, Aslani F, Walecki M, Wahle E, Eisel F, Bhushan S, Hackstein H, Baal N, Schuler G, Konrad L, Rafiq A, O’Hara L, Smith LB and Meinhardt A. (2015) Influence of testosterone on inflammatory response in testicular cells and expression of transcription factor Foxp3 in T cells. American Journal of Reproductive Immunology Jan 19. doi: 10.1111/aji.12363

Mitchell RT, Mungall W, McKinnell C, Sharpe RM, Cruickshanks L, Milne L, and Smith LB. (2015) Anogenital distance (AGD) plasticity in adulthood: Implications for its use as a biomarker of fetal androgen action. Endocrinology. Jan; 156(1): 24–31.


Zimmermann C, Romero Y, Warnefors M, Bilican A, Borel C, Smith LB, Kotaja N, Kaessmann H, and Nef S. (2014) Germ Cell-Specific Targeting of DICER or DGCR8 reveals a novel role for endo-siRNAs in the progression of mammalian spermatogenesis and male fertility. PLoS ONE DOI: 10.1371/journal.pone.0107023

Rebourcet D, O’Shaughnessy PJ, Monteiro A, Milne L, Cruickshanks L, Jeffrey N, Guillou F, Freeman TC, Mitchell RT, and Smith LB. (2014) Sertoli cells maintain Leydig cell number and Peritubular Myoid cell activity in the adult mouse testis PLoS ONE 9(8): e105687. doi:10.1371/journal.pone.0105687

Smith LB, Walker WH and O’Donnell L. (2014) Hormonal Regulation of Spermatogenesis through Sertoli cells by Androgens and Estrogens. - In Sertoli Cell Biology, Griswold M, Ed.

Wu J, Hadoke PWF, Mair I, Lim WG, Miller E, Denvir MA, and Smith LB (2014) Modulation of Neointimal Lesion Formation by Endogenous Androgens is Independent of Vascular Androgen Receptor. Cardiovascular Research doi: 10.1093/cvr/cvu142

Hume D, Gow D, Sauter K, Pridans C, Moffat L, Sehgal A, Stutchfield B, Raza S, Beard P, Tsai YT, Bainbridge G, Boner P, Fici G, Garcia-Tapia D, Martin R, Oliphant T, Shelly J, Tiwari R, Wilson T, Smith LB, Forbes S, and Mabbott N. (2014) Characterisation of a novel Fc conjugate of Macrophage Colony-Stimulating Factor (CSF1). Molecular Therapy doi: 10.1038/mt.2014.112

Kilcoyne KR, Smith LB, Atanassova N, Macpherson S, McKinnell C, van den Driesche S, Jobling M, Chambers TJG, de Gendt K, Verhoeven G, OHara L, Platts S, Renato de Franca L, Lara NLM, Anderson RA, and Sharpe RM (2014) Fetal programming of adult Leydig cell function via androgenic effects on stem/progenitor cells. Proc Nat Aca Sci USA (In Press)

Rebourcet D, O’Shaughnessy PJ, Pitetti J-L, Monteiro A, O’Hara L, Milne L, Tsai Y-T, Cruickshanks L, Riethmacher D, Guillou F, Mitchell RT, van ‘t Hof R, Freeman TC, Nef S, and Smith LB. (2014) Sertoli cells control peritubular myoid cell fate and support adult Leydig cell development in the prepubertal testis. Development  (In Press)

Smith LB, and Walker WH. (2014) The Regulation of Spermatogenesis by Androgens. Seminars in Cell and Developmental Biology Mar 2. pii: S1084-9521(14)00024-X. doi: 10.1016/j.semcdb.2014.02.012.

Sauter K, Pridans C, Sehgal A, Tsai YT, Bradford B, Raza S, Moffat L, Gow D, Beard P, Mabbott N, Smith LB, and Hume D. (2014) Pleiotropic Effects Of Extended Blockade Of Csf1r Signalling In Adult Mice Journal of Leukocyte Biology (In Press)

O’Hara L, York JP, Zhang P, and Smith LB (2014) Targeting of GFP-Cre to the mouse Cyp11a1 locus both drives Cre Recombinase expression in steroidogenic cells and permits generation of Cyp11a1 knock out mice. PLoS ONE 9(1): e84541. doi:10.1371/journal.pone.0084541

Smith LB and Walker  WH. (2014) Hormone Signaling in the Testis. In Knobil and Neill’s Physiology of Reproduction (Fourth Edition) (In Press)

Mitchell RT, Camacho-Moll M, Macdonald J, Anderson RA, Kelnar CJH, O’Donnell, M, Sharpe RM, Smith LB, Grigor KM, Wallace WHB, Stoop H, Wolffenbuttel KP, Donat R, *Saunders PTK and *Looijenga LHJ. (2014) Intratubular germ cell neoplasia of the human testis: heterogeneous protein expression and relation to invasive potential Modern Pathology doi: 10.1038/modpathol.2013.246


Hall EA, Keighren M, Ford MJ, Davey T, Jarman AP, Smith LB, Jackson IJ and Mill P. (2013) Acute versus chronic loss of mammalian Azi1/Cep131 results in distinct ciliary phenotypes. PLoS Genetics PLoS Genet 9(12): e1003928. doi:10.1371/journal.pgen.1003928 (FACULTY 1000 Citation)

Smith LB, Mitchell RT and McEwan IJ. (2013) Testosterone: From Basic Research to Clinical Applications. Springer Briefs – Reproductive Biology Series

Mitchell R, Sharpe RM, Anderson RA, McKinnell C, Macpherson S, Smith LB, Wallace WHB, Kelner C, and van den Driesche S (2013) Diethylstilboestrol exposure does not reduce testosterone production in human fetal testis xenografts. PLoS ONE 8(4): e61726. doi:10.1371/journal.pone.0061726


Smith LB, Milne L, Nelson N, Eddie S, Brown P, Atanassova N, O’Bryan MK, O’Donnell L, Rhodes D, Wells S, Napper D, Nolan P, Lalanne Z, Cheeseman M and Peters J. (2012) KATNAL1 Regulation of Sertoli Cell Microtubule Dynamics is Essential for Spermiogenesis and Male Fertility. PLoS Genetics PLoS Genet 8(5): e1002697. doi:10.1371/journal.pgen.1002697

O’Donnell L, Rhodes D, O’Connor AE, Smith SJ, Clark B, Borg C, Whittle B, Merriner DJ, Smith LB, McNally F, de Kretser DM, Goodnow CC, Ormandy CJ, Jamsai D and O’Bryan MK. (2012) An essential role for p80 katanin and microtubule severing in male gamete production. PLoS Genetics PLoS Genet 8(5): e1002698. doi:10.1371/journal.pgen.1002698

Van den Driesche S, Walker M, McKinnell C, Scott HM, Eddie SL, Mitchell RT, Seckl JR, Drake AJ, Smith LB, Anderson RA and Sharpe RM (2012) Proposed Role for COUP-TFII in Regulating Fetal Leydig Cell Steroidogenesis, Perturbation of which leads to Masculinization Disorders in Rodents. PLoS ONE 7(5): e37064. doi:10.1371/journal.pone.0037064

McInnes KJ, Smith LB, Hunger NI, Saunders PTK, Andrew R and Walker BR. (2012) Deletion of the Androgen Receptor in adipose tissue in male mice elevates retinol binding protein 4 and reveals independent effects on visceral fat mass and on glucose homeostasis. Diabetes 2012 Mar 13. [Epub ahead of print]

O’Hara L and Smith LB. (2012) Androgen Receptor Signalling in Vascular Endothelial Cells is Dispensable for Spermatogenesis and Male Fertility. BMC Research Notes 5:16

Dean A, Smith LB, Macpherson S and Sharpe RM.  (2012) The effect of dihydrotestosterone exposure during or prior to the masculinization programming window on reproductive development in male and female rats. International journal of Andrology doi: 10.1111/j.1365-2605.2011.01236.x.

Welsh M, Moffat L, Belling K, Renato de França L, Segatelli T, Saunders PTK, Sharpe RM, and Smith LB (2012) Androgen Receptor Signalling in Peritubular Myoid Cells is Essential for Normal Differentiation and Function of Adult Leydig Cells. International Journal of Andrology 35: 25-40


Denison FC, Smith LB, Muckett PJ, O’Hara L,Carling D, Woods A. (2011) LKB1 is an Essential Regulator of Spermatozoa Release During Spermiation in the Mammalian Testis. PLoS ONE 6(12): e28306. doi:10.1371/journal.pone.0028306

Welsh M, Moffat L, McNeilly A, Brownstein D, Saunders PTK, Sharpe RM, Smith LB. (2011) Smooth muscle cell-specific knockout of androgen receptor: a new model for prostatic disease. Endocrinology. 152(9):3541-51.

Smith LB and Saunders PTK (2011) The Skeleton: the new controller of male fertility? Cell. 144(5) 642-643

O’Hara L, Welsh M, Saunders PTK and Smith LB (2011) Androgen Receptor Expression In The Caput Epididymal Epithelium Is Essential For Development Of The Initial Segment And Epididymal Spermatozoa Transit. Endocrinology 152 (2): 718-729 (Faculty of 1000 citation)

Smith LB (2011) Good Planning and Serendipity: Exploiting the Cre/Lox system in the Testis. Reproduction. 141(2):151-61

Orr B, Vanpoucke G, Grace OC, Smith LB, Anderson RA, Riddick ACP, Franco OE, Hayward SW and Thomson AA. (2011) Expression of Pleiotrophin in the Prostate is Androgen Regulated and it Functions as an Autocrine Regulator of Mesenchyme and Cancer Associated Fibroblasts and as a Paracrine Regulator of Epithelia. Prostate. 71(3):305-17


Smith LB, Hadoke PWF, Dyer E, Denvir M, Brownstein D, Miller E, Nelson N, Wells S, Cheeseman M, and Greenfield A. (2010) Haploinsufficiency of the murine Col3α1 locus causes aortic dissection: a novel model of the vascular type of Ehlers Danlos Syndrome. Cardiovascular Research. doi: 10.1093/cvr/cvq356

Welsh M, Sharpe RM, Moffat L, Atanassova N, Saunders PTK, Kilter S, Bergh A, Smith LB, (2010) Androgen action via testicular arteriole smooth muscle cells is important for Leydig cell function, vasomotion and testicular fluid dynamics. PLoS One 5(10): e13632.

Welsh M, Moffat L, Jack L, McNeilly A, Brownstein D, Saunders PT, Sharpe RM, Smith LB. (2010) Deletion of androgen receptor in the smooth muscle of the seminal vesicles impairs secretory function and alters its responsiveness to exogenous testosterone and estradiol. Endocrinology. 151:3374-85.

Willems A, De Gendt K, Allemeersch J, Smith LB, Welsh M, Swinnen JV and Verhoeven G (2010) Early effects of Sertoli cell-selective androgen receptor ablation on testicular gene expression. Int J. Androl 33(3):507-1


Welsh M, Saunders PT, Atanassova N, Sharpe RM, Smith LB. (2009) Androgen action via testicular peritubular myoid cells is essential for male fertility. FASEB J. Dec; 23(12):4218-30 (Faculty of 1000 citation)

Smith LB, Saunders PTK (2009) What makes a man? The effect of androgen receptors on male fertility. The Biochemist 31:12-15

Welsh M, MacLeod DJ, Walker M, Smith LB and Sharpe RM. (2009) Critical androgen-sensitive periods of rat penis and clitoris development. International Journal of Andrology 32, 1–8

Welsh M, Sharpe RM, Walker M, Smith LB, Saunders PT. (2009). New insights into the role of androgens in Wolffian duct stabilization in male and female rodents. Endocrinology. 150: 2472-2480

Pastorelli LM, Wells S, Fray M, Smith A, Hough T, Harfe BD, McManus MT, Smith LB, Woolf AS, Cheeseman M, Greenfield A. (2009) Genetic analyses reveal a requirement for Dicer1 in the mouse urogenital tract. Mamm Genome. Volume: 20 Issue: 3 Pages: 140-151


Welsh M, Saunders PT, Fisken M, Scott HM, Hutchison GR, Smith LB, and Sharpe RM (2008) Identification in rats of a programming window for reproductive tract masculinization, disruption of which leads to hypospadias and cryptorchidism. J Clin Invest 118: 1479-90

Smith LB, Willan J, Warr N, Brook FA, Cheeseman M, Sharpe R, Siggers P, Greenfield A. (2008). The Maestro (Mro) gene is dispensable for normal sexual development and fertility in mice. PLoS ONE. 3(12):e409