Professor Paul Fowler

  Professor Paul Fowler The University of Aberdeen School of Medicine & Dentistry Professor Paul Fowler Chair in Translational Medical Sciences work +44 (0)1224 437528 work fax +44 (0) 1224 437465 pref

Institute of Medical Sciences Division of Applied Medicine University of Aberdeen Foresterhill Aberdeen AB25 2ZD Scotland

Chair in Translational Medical Sciences

BSc (Hons 1st) Zoology, Aberdeen 1982. PhD, Aberdeen 1986, FSB

Professor Paul Fowler

Contact Details

Telephone: +44 (0)1224 437528
Fax: +44 (0) 1224 437465


Institute of Medical Sciences Division of Applied Medicine University of Aberdeen Foresterhill Aberdeen AB25 2ZD Scotland


I am a signatory of the The 2013 Berlaymont Declaration on Endocrine Disrupters.

European Commission: Science in Society - How gender analysis contributes to research - REEF Project as a case study in Gendered Innovations:

Thanks to for awarding a summer studentship:

"Proteomic analysis of the effects of maternal smoking on the second trimester human fetal gonad "


Professional Society Membership:



I am a long-term member of Fauna & Flora International 

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Professor Fowler received his BSc Hons and PhD in Zoology at the University of Aberdeen. He later moved to Obstetrics & Gynaecology, University of Aberdeen, to work on ovarian hormones and antiprogesterones.  In 2000 he moved to the Institute of Medical Sciences, University of Aberdeen as a Professor of Translational Medical Sciences and is currently lead for the Cell, Developmental & Cancer Biology Research programme at Aberdeen.

Professor Fowler has spent much of his career working on elucidating mechanisms in the regulation of reproduction and has over 100 peer-reviewed publications, book chapters and editorials. At each stage he has made contributions in the use of technologies to answer reproductive questions, including: remote telemetry of body-testis temperature differentials, MRI analysis of body composition and mammary gland, phage library and protein purification techniques to investigate reproductive proteins and proteomic and microarray techniques to study fetal development and endometriosis. Since the turn of the century, his research has focused on the effects of environmental exposures and endocrine disruption on fetal development and subsequent health in both human and animal models. His group is one of the few to work on the normal second trimester human fetus and uses maternal smoking as a model to understand how adverse in-utero environment disturbs fetal development in our own species.

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Research Interests

Since the breakthrough epidemiological studies of the 1980s it has been clear that adult human health is dependent upon fetal development and fetal programming (‘early life programming’). Fetal growth restriction, for example, can lead to hypertension, type 2 diabetes, obesity and cardiovascular disease in the adult. Dysfunctional changes in the fetus can arise through other mechanisms, such as maternal cigarette smoking or environmental pollutant exposure and the health trajectory of an individual is likely to depend upon the pollutant burden carried at birth. Fetal pollutant exposure has clear adverse effects on development but we currently have very little knowledge about levels of fetal exposure to “real-life” complex pollutant cocktails and developmental consequences. This fundamental lack of knowledge is a major impasse to developing strategies for reducing fetal pollutant burdens and predicting likely health outcomes. Available data is limited, and often indirect (i.e. animal models or surrogates of exposure), but all studies suggest that fetal pollutant exposure affects lifelong health outcomes and, thereby, impacts on the general economy.  

Fetal reproductive development and the environment

(a) Human fetal development and its disruption

Better understanding of events during key stages in reproductive development in the human are key to determining the risks posed by exogenous chemicals, diet and lifestyle on reproductive health in humans, from breast cancer to infertility, testicular cancer to endometriosis, sperm counts to developmental defects. This kind of data is also key to translating the applicability and importance of findings in animal models to the human. Cigarette smoke contains well over 4,000 pollutant chemicals, including potentially toxic elements (PTEs, e.g. cadmium) persistent organic pollutants (POPs, e.g. dioxin) and, of course, nicotine. Many of these chemicals are endocrine-disrupting compounds (EDCs, e.g. nicotine reduces steroidogenesis). Thus, cigarette smoking provides a unique “real-life” model to study endocrine disruption in humans. Since up to 30% of women are smokers and few give up the habit, even when pregnant, it remains a major health challenge. Furthermore, fetal exposure to cigarette smoke chemicals constitutes one of the few models we can use to determine what effects environmental chemicals might have on our own species. This is one of the major models used in my research.

(i) Characterising normal events during human fetal development.

1st gene array study of the human fetal gonad : Gene expression analysis of human fetal ovarian primordial follicle formation. Fowler PA, Flannigan S, Mathers A, Gillanders K, Lea RG, Wood MJ, Maheshwari A, Bhattacharya S, Collie-Duguid ES, Baker PJ, Monteiro A, O'Shaughnessy PJ. J Clin Endocrinol Metab. 2009 Apr;94(4):1427-35.

(ii) Investigating how these events are affected by exposure to cigarette smoke chemicals.

1st  fetal testis gene reduced by maternal cigarette smoking: Maternal smoking during pregnancy specifically reduces human fetal desert hedgehog gene expression during testis development. Fowler PA, Cassie S, Rhind SM, Brewer MJ, Collinson JM, Lea RG, Baker PJ, Bhattacharya S, O'Shaughnessy PJ. J Clin Endocrinol Metab. 2008 Feb;93(2):619-26.

(b) Effects of environmental chemicals: using animal models to understand risks

Domestic animals, like humans and all other animals, are exposed to environmental pollutants through exposure to contaminated food, water, air and soil. Environmental pollutants include a range of (mainly) anthropogenic chemicals which have the capacity to interfere, subtly, with hormonal systems and therefore with their capacity to reproduce, rear offspring and fight disease); these chemicals, which included heavy metals and organic compounds of many different classes are described as endocrine disrupting compounds (EDCs). The physiological insult associated with exposure to EDCs involves many different chemical types, each acting on different physiological systems. Furthermore, while environmental concentrations of most individual chemicals are very low and below the No Observable Effect Level (NOEL), in many cases they can act additively to induce a physiological change, even when concentrations of individual chemicals appear harmless.

Empirical evidence in support of this has been provided by our previous work involving exposure of sheep to low (environmental) levels of EDCs through grazing pastures fertilised with sewage sludge (contains high concentrations of multiple EDCs). This work has shown that maternal exposure to sewage sludge treated pasture was associated with perturbation of the fetal hypothalamus-pituitary and ovary, despite the fact that fetal and adult tissue concentrations of individual EDCs at the time of slaughter were minimally increased by such exposure and were at concentrations below the NOEL. Work funded by the Wellcome Trust and the European Union is addressing the effects of exposure during specific windows of time, before and during gestation, on patterns of tissue accumulation and associated physiological changes. 

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Current Research

Current Team: Dr Pan Filis, Ms Samantha Flannigan, Ms Margaret Fraser.

RECENT HUMAN FETAL PUBLICATION  In utero exposure to cigarette smoke dysregulates human fetal ovarian developmental signalling. Fowler PA, Childs AJ, Courant F, MacKenzie A, Rhind SM, Antignac JP, Le Bizec B, Filis P, Evans F, Flannigan S, Maheshwari A, Bhattacharya S, Monteiro A, Anderson RA, O'Shaughnessy PJ. Hum Reprod. 2014 July 29(7):1471-82.

RECENT ANIMAL MODEL PUBLICATION   Exposure to chemical cocktails before or after conception--- the effect of timing on ovarian development. Bellingham M, Amezaga MR, Mandon-Pepin B, Speers CJ, Kyle CE, Evans NP, Sharpe RM, Cotinot C, Rhind SM, Fowler PA. Mol Cell Endocrinol. 2013 Aug 25;376(1-2):156-72.


Recent plenary and invited presentations:

  • Regulation and dysregulation of primordial follicle formation and activation. 3rd World Congress of Reproductive Biology, 2-4 September 2014, Edinburgh, UK.
  • Environmental chemicals and fetal development. 2ieme Journee des Jeunes Chercheurs de L’IRSET, February 4th 2014, University of Rennes, France.
  • On women and sheep: making and breaking the fetal ovary. INRA, 13 December 2013, Jouy-en-Josas, France.
  • The evidence for EDC involvement in reproductive dysfunction in women. International Workshop: Endocrine Disrupting Chemicals and Female Reproduction. 5-6 November 2013, Uppsala, Sweden.
  • Environment and endocrine disruptors in fetal reproductive development.  IfLS Conference: Reproductive Biology – from gametes to systems, and between generations. 17 September 2013, Southampton, UK.
  • Effects of chemicals on reproductive development. Subspecialist Trainees in Reproductive Medicine and Surgery (STIRMAS). 27 September 2013, Cambridge, UK.
  • Human fetal reproductive development and endocrinology: effects of maternal smoking. 7th Copenhagen Workshop of Endocrine Disrupters, 28-31 May 2013, Copenhagen, Denmark.
  • Impact of endocrine-disrupting compounds (EDCs) on female reproductive health. Fertility 2013, Liverpool, January 3-5 2013.
  • Effects of exposure to environmental chemicals during pregnancy on the development of the male and female reproductive access“. 17th International Congress on Animal Reproduction (ICAR 29012) in Vancouver, British Columbia, Canada, July 29 – August 2, 2012.
  • Environmental exposure in reproductive development and function of the female. World Congress on Reproductive Biology, Cairns Australia 9-11 October 2011.
  • Environmental effects on fetal reproductive development. Australia Society of Reproductive Biology, Cairns Australia 7-9 October 2011.
  • Effects of EDCs on the human foetus: lessons from smoking. 6th Copenhagen Workshop of Endocrine Disrupters, 25-30 April 2011, Copenhagen, Denmark.
  • Maternal smoking and sewage sludge: consequences for fetal gonadal development. Centre for Reproductive Biology Special Symposium: “Endocrine disruptors and reproductive health: separating fact from fiction.” 7 March 2011, University of Edinburgh.
  • Mixtures of environmental chemicals: concentrations and effects in the fetus. Covance Environmental Risk Assessment (ERA) Symposium, 2-3 March 2011, Brussels.
  • In-utero exposures to environmental chemicals and consequences for ovarian development. Gordon Research Conference: “Environmental Endocrine Disrupters", 30 May-4 June 2010, Les Diablerets, Switzerland.
  • Environmental endocrine disruption of the ovary. 37th Nordic Congress of Obstetrics & Gynecology, 15-18 June 2010, Copenhagen, Denmark.
  • Endocrine disruptors and gonadal development.  Symposium: Environmental factors and nosology of the endocrine system, 2-4 October 2009, Athens, Greece.
  • In-utero exposure to environmental chemicals and consequences ovarian development. 5th Copenhagen Workshop of Endocrine Disrupters, 20-22 May 2009, Copenhagen, Denmark.
  • The fetal ovary as a target for disruption. Danish Society of Obstetrics & Gynaecology, 17 April 2009, Copenhagen, Denmark
  • Is the fetus under chemical attack? St George’s University London, 26 March 2009.
  • Gender bending chemicals and the mammalian fetal gonad. XXth International Congress of Zoology, Paris 26-29 August 2008.

Some press releases:

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  • Grateful thanks to Dr Stewart Rhind (James Hutton Institute, Aberdeen, Scotland) for the years of friendship and collaboration before his untimely death.

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Research Grants

  • Fowler PA, Douglas A, Hay D, Iredale J, Fairley S, O'Shaughnessy PJ, Huestis M, The human fetal liver: development and response to maternal drug use. MRC, £588,601.
  • Fowler PA, Filis P, O'Shaughnessy PJ, Does exposure to a complex cocktail of environmental chemicals during pregnancy and lactation alter metabolic and steroid hormone systems in the adult liver? Society for Reproduction & Fertility, Academic Schorlarship, £8,900
  • Bellingham M, Fowler PA, O'Shaughnessy PJ, Prenatal programming of adult disease by maternal smoking: the adrenal gland as a key player? Wellcome Trust Institutional Strategic Support Fund (Catalyst), £18,230.
  • Fowler PA, Bhattacharya S, O'Shaughnessy PJ, Gatekeepers to the cell: which ones operate customs posts in the fetal liver and does maternal smoking allow drugs to slip through? NHS Grampian Endowments, £9,356.
  • Fowler PA, Collinson JM, O’Shaughnessy PJ, Bioincubator system to model toxic effects on the human fetal testis, Chief Scientist Office, £62,078.
  • Fowler PA, Harkany T, O’Shaughnessy PJ, Fetal human gonadal cannabinoid receptors: expression and effects of maternal smoking., NHS Grampian Endowments, £6,713.
  • Fowler PA, Cotinot C, O’Shaughnessy PJ, Inter-species comparison of the onset ovarian follicle formation, Wellcome Trust summer studentship, £5,760
  • Fowler PA, Cotinot C, Fischer B, Lea RG, Pocar P, Sinclair K, Rhind SM, Reproductive effects of environmental chemicals in females (REEF), European Commission, Framework 7, £2,322,380. REEF
  • Fowler PA, Bhattacharya S, Rhind SM, O'Shaughnessy PJ, Does maternal smoking disrupt key developmental sex steroid hormones in the fetus? NHS Grampian Endowments, £10,834.
  • Fowler PA, Sharpe R, Cotinot C, Evans N, Rhind S, Effect of in-utero exposure to environmental chemicals via maternal pasture ingestion on fetal development., Wellcome Trust, £425,786.
  • Fowler PA, Maheshwari A, Bhattacharya S, Follicular assembly in the human fetal ovary, IMS Microarray Core Facility grant, UoA, £6,000.
  • Fowler PA, Wood M, Human fetal gonad: 2 summer research studentships, SRF, £2,640.
  • Skaare JU et al, including Fowler PA, The mechanisms behind toxic effects of POPS, NFR, £170,377.
  • Fowler PA, Bhattacharya S, Hamilton MPR, al-Gubory K, Identification of implantation-related proteins, NHS R&D, £18,500.
  • Fowler PA, Bhattacharya S, Cash P, Circulating biomarkers for endometriosis, University of Aberdeen Commercialisation Fund, £19,095.
  • Shanbhag S, Cruickshank M, Millier I, Murray G, Fowler PA, Molecular basis of granulosa cell tumours, NHS Grampian Endowments, £11,343.
  • Fowler PA, Bhattacharya S, Cash P, Establishing a relationship between endometrial proteins and endometriosis., CSO, £18,856.
  • Fowler PA, Bhattacharya S, Optimisation of an in-vitro model to investigate the human fetal testis, CSO, £15,571.
  • Huhtaniemi I, Fowler PA, Cash P, Charlton H, Novel biological actions of luteinizing hormone and its receptor, Wellcome Trust, £1,408,031.
  • Fowler PA and colleagues, Translational medicine funding: >£700,000.

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Teaching Responsibilities

  • BM4004: Advanced Molecules, membranes and cells
  • BM3803: Epithelial physiology
  • DB3502: Reproductive biology - I coordinate this third year course.
  • SX3003: 6th Century course: An appetite for food and health
  • Phase I MBChB, ME1504: Endocrine and reproductive system

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External Responsibilities

  • Member, Editorial Board of the journal Molecular & Cellular Endocrinology, 2014-
  • Member, Food ingredients and packaging (CEF) panel, EFSA, 2014-2017
  • Member, Editorial Board of the journal Endocrinology, 2014-
  • 2009-2013: Chair, Society for Reproduction & Fertility
  • I have served/am serving on committees and council for the Society for Reproduction & Fertility and the British Fertility Society.
  • I am presently on the Editorial Boards/Advisory Boards for the journals Teratology Studies, Reproductive Biology & Endocrinology and Current Medicinal Chemistry.
  • I am/have been an External Examiner for the Universities of Edinburgh and Nottingham and St George's Hospital Medical School, London.
  • Member, AERES research institute evaluation committee, 2013
  • Member, Expert review Panel for CRACK IT NC3Rs, 2012
  • Member, Expert Working Group on BPA, EFSA 2012-2014
  • Member, Gendered Innovations, FP7 case study panel, 2012
  • Member, Standing Evaluation Panel, Environmental Pollution, Swedish Research Council Formas, 2010

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Admin Responsibilities

  • Lead, Cell, Developmental & Cancer Biology Research Programme, 2011-
  • Member, European Advisory Committee
  • Member, Division of Applied Medicine Executive
  • Member, Schools of Medicine & Dentistry and Medical Sciences Joint Research Committee
  • Member, Athena Swann CLSM assessment team

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I work to foster research at Aberdeen via my responsibility for the management, monitoring and generic training of laboratory postgraduate researchers in the School of Medicine under the aegis of the Graduate School, College of Medicine & Life Sciences.

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Contributions to Journals


  • Anderson, RA., McIlwain, L., Coutts, S., Kinnell, HL., Fowler, PA. & Childs, AJ. (2014). 'Activation of the aryl hydrocarbon receptor by a component of cigarette smoke reduces germ cell proliferation in the human fetal ovary'. Molecular Human Reproduction, vol 20, no. 1, pp. 42-48.
    [Online] DOI: 10.1093/molehr/gat059
    [Online] AURA: Anderson_et_al_MHR14_AHR_activation_T1_human_fetal_ovary_ger...
  • Al-Gubory, KH., Arianmanesh, M., Garrel, C., Bhattacharya, S., Cash, P. & Fowler, PA. (2014). 'Proteomic analysis of the sheep caruncular and intercaruncular endometrium reveals changes in functional proteins crucial for the establishment of pregnancy'. Reproduction, vol 147, no. 5, pp. 599-614.
    [Online] DOI: 10.1530/REP-13-0600
    [Link] N2 -
  • O'Shaughnessy, PJ., Monteiro, A., Fowler, PA. & Morris, ID. (2014). 'Identification of Leydig cell-specific mRNA transcripts in the adult rat testis'. Reproduction, vol 147, pp. 671-682.
    [Online] DOI: 10.1530/REP-13-0603
  • Krogenæs, AK., Ropstad, E., Gutleb, AC., Hårdnes, N., Berg, V., Dahl, E. & Fowler, PA. (2014). 'In Utero Exposure to Environmentally Relevant Concentrations of PCB 153 and PCB 118 Disrupts Fetal Testis Development in Sheep'. Journal of Toxicology and Environmental Health. Part A, vol 77, no. 9-11, pp. 628-649.
    [Online] DOI: 10.1080/15287394.2014.887426
  • Tremoen, NH., Fowler, PA., Ropstad, E., Verhaegen, S. & Krogenæs, A. (2014). 'Exposure to the Three Structurally Different PCB Congeners (PCB 118, 153, and 126) Results in Decreased Protein Expression and Altered Steroidogenesis in the Human Adrenocortical Carcinoma Cell Line H295R'. Journal of Toxicology and Environmental Health. Part A, vol 77, no. 9-11, pp. 516-534.
    [Online] DOI: 10.1080/15287394.2014.886985
  • Fowler, PA., Childs, AJ., Courant, F., MacKenzie, A., Rhind, SM., Antignac, J-P, Le Bizec, B., Filis, P., Evans, F., Flannigan, S., Maheshwari, A., Bhattacharya, S., Monteiro, A., Anderson, RA. & O'Shaughnessy, PJ. (2014). 'In utero exposure to cigarette smoke dysregulates human fetal ovarian developmental signalling'. Human Reproduction, vol 29, no. 7, pp. 1471-1489.
    [Online] DOI: 10.1093/humrep/deu117
  • O'Shaughnessy, PJ., Monteiro, A., Bhattacharya, S., Fraser, MJ. & Fowler, PA. (2013). 'Steroidogenic enzyme expression in the human fetal liver and potential role in the endocrinology of pregnancy'. Molecular Human Reproduction, vol 19, no. 3, pp. 177-187.
    [Online] DOI: 10.1093/molehr/gas059
  • Hombach-Klonisch, S., Danescu, A., Begum, F., Amezaga, MR., Rhind, SM., Sharpe, RM., Evans, NP., Bellingham, M., Cotinot, C., Mandon-Pepin, B., Fowler, PA. & Klonisch, T. (2013). 'Peri-conceptional changes in maternal exposure to sewage sludge chemicals disturbs fetal thyroid gland development in sheep'. Molecular and Cellular Endocrinology, vol 367, no. 1-2, pp. 98-108.
    [Online] DOI: 10.1016/j.mce.2012.12.022
  • Bellingham, M., Amezaga, MR., Mandon-Pepin, B., Speers, CJB., Kyle, CE., Evans, NP., Sharpe, RM., Cotinot, C., Rhind, SM. & Fowler, PA. (2013). 'Exposure to chemical cocktails before or after conception: The effect of timing on ovarian development'. Molecular and Cellular Endocrinology, vol 376, no. 1-2, pp. 156-172.
    [Online] DOI: 10.1016/j.mce.2013.06.016
    [Online] AURA: Bellingham_et_al_MCE13_fetal_sheep_ovary_sewage_sludge_prepo...
  • Knipe, A., Fowler, PA., Ramsay, S., Haydon, DT., McNeilly, AS., Thirgood, S. & Newey, S. (2013). 'The effects of population density on the breeding performance of mountain hare Lepus timidus'. Wildlife Biology, vol 19, no. 4, pp. 473-482.
    [Online] DOI: 10.2981/12-109
  • Fowler, PAF., Bellingham, M., Sinclair, KD., Evans, NP., Pocar, P., Fischer, B., Schaedlich, K., Schmidt, S-J, Amezaga, MR., Bhattacharya, S., Rhind, SM. & O'Shaughnessy, PJ. (2012). 'Impact of endocrine-disrupting compounds (EDCs) on female reproductive health'. Molecular and Cellular Endocrinology, vol 355, no. 2, pp. 231-239.
    [Online] DOI: 10.1016/j.mce.2011.10.021
  • Kucerova, R., Dorà, N., Mort, RL., Wallace, K., Leiper, LJ., Lowes, C., Neves, C., Walczysko, P., Bruce, F., Fowler, PA., Rajnicek, AM., McCaig, CD., Zhao, M., West, JD. & Collinson, JM. (2012). 'Interaction between hedgehog signalling and PAX6 dosage mediates maintenance and regeneration of the corneal epithelium'. Molecular vision, vol 18, pp. 139-150.
  • Bellingham, M., McKinnell, C., Fowler, PA., Amezaga, MR., Zhang, Z., Rhind, SM., Cotinot, C., Mandon-Pepin, B., Evans, NP. & Sharpe, RM. (2012). 'Foetal and post-natal exposure of sheep to sewage sludge chemicals disrupts sperm production in adulthood in a subset of animals'. International journal of andrology, vol 35, no. 3, pp. 317-329.
    [Online] DOI: 10.1111/j.1365-2605.2011.01234.x
  • McLaggan, D., Amezaga, MR., Petra, E., Frost, A., Duff, EI., Rhind, SM., Fowler, PA., Glover, LA. & Lagido, C. (2012). 'Impact of sublethal levels of environmental pollutants found in sewage sludge on a novel Caenorhabditis elegans model biosensor'. PLoS ONE, vol 7, no. 10, e46503.
    [Online] DOI: 10.1371/journal.pone.0046503
    [Online] AURA: Impact_of_sublethal.pdf
  • Willing, C., Peich, M., Danescu, A., Kehlen, A., Fowler, PAF. & Hombach-Klonisch, S. (2011). 'Estrogen-independent actions of environmentally relevant AhR-agonists in human endometrial epithelial cells'. Molecular Human Reproduction, vol 17, no. 2, pp. 115-126.
    [Online] DOI: 10.1093/molehr/gaq081
  • Arianmanesh, M., McIntosh, RH., Lea, RG., Fowler, PA. & Al-Gubory, KH. (2011). 'Ovine corpus luteum proteins, with functions including oxidative stress and lipid metabolism, show complex alterations during implantation'. Journal of Endocrinology, vol 210, pp. 47-58.
    [Online] DOI: 10.1530/JOE-10-0336
  • O'Shaughnessy, PJ., Monteiro, A. & Fowler, PAF. (2011). 'Identification of stable endogenous reference genes for real-time PCR in the human fetal gonad using an external standard technique'. Molecular Human Reproduction, vol 17, no. 10, pp. 620-625.
    [Online] DOI: 10.1093/molehr/gar032
    [Online] AURA: Fowler_Identification_of_stable_endogenous_reference_genes.p...
  • O'Shaughnessy, PJ., Monteiro, A., Bhattacharya, S. & Fowler, PA. (2011). 'Maternal Smoking and Fetal Sex Significantly Affect Metabolic Enzyme Expression in the Human Fetal Liver'. Journal of Clinical Endocrinology and Metabolism, vol 96, no. 9, pp. 2851-2860.
    [Online] DOI: 10.1210/jc.2011-1437
  • Fowler, PA., Bhattacharya, S., Flannigan, S., Drake, AJ. & O'Shaughnessy, PJ. (2011). 'Maternal cigarette smoking and effects on androgen action in male offspring: unexpected effects on second-trimester anogenital distance'. Journal of Clinical Endocrinology and Metabolism, vol 96, no. 9, pp. E1502-E1506.
    [Online] DOI: 10.1210/jc.2011-1100
  • Rhind, SM., Evans, NP., Bellingham, M., Sharpe, RM., Cotinot, C., Mandon-Pepin, B., Loup, B., Sinclair, KD., Lea, RG., Pocar, P., Fischer, B., van der Zalm, E., Hart, K., Schmidt, J-S, Amezaga, MR. & Fowler, PA. (2010). 'Effects of environmental pollutants on the reproduction and welfare of ruminants'. Animal, vol 4, no. 7, pp. 1227-1239.
    [Online] DOI: 10.1017/S1751731110000595
    [Online] AURA: Rhind_et_al_Animal10_review_Effects_of_environmental_polluta...
  • Bellingham, M., Fowler, PA., Amezaga, MR., Whitelaw, CM., Rhind, SM., Cotinot, C., Mandon-Pepin, B., Sharpe, RM. & Evans, NP. (2010). 'Foetal hypothalamic and pituitary expression of gonadotrophin-releasing hormone and galanin systems is disturbed by exposure to sewage sludge chemicals via maternal ingestion'. Journal of Neuroendocrinology, vol 22, no. 6, pp. 527-533.
    [Online] DOI: 10.1111/j.1365-2826.2010.01974.x
  • Garrel, C., Fowler, PAF. & Al-Gubory, KH. (2010). 'Developmental changes in antioxidant enzymatic defences against oxidative stress in sheep placentomes'. Journal of Endocrinology, vol 205, no. 1, pp. 107-116.
    [Online] DOI: 10.1677/JOE-09-0362
  • Rhind, SM., Kyle, CE., MacKie, CC., McDonald, L., Zhang, Z., Duff, EI., Bellingham, M., Amezaga, MR., Mandon-Pepin, B., Loup, B., Cotinot, C., Evans, NP., Sharpe, RM. & Fowler, PAF. (2010). 'Maternal and fetal tissue accumulation of selected endocrine disrupting compounds (EDCs) following exposure to sewage sludge-treated pastures before or after conception'. Journal of Environmental Monitoring, vol 12, no. 8, pp. 1582-1593.
    [Online] DOI: 10.1039/c0em00009d
  • Bellingham, M., Fowler, PA., Amezaga, MR., Rhind, SM., Cotinot, C., Mandon-Pepin, B., Sharpe, RM. & Evans, NP. (2009). 'Exposure to a complex cocktail of environmental endocrine-disrupting compounds disturbs the Kisspeptin/GPR54 system in ovine hypothalamus and pituitary gland'. Environmental Health Perspectives, vol 117, no. 10, pp. 1556-1562.
  • Fowler, PA., Flannigan, S., Mathers, A., Gillanders, K., Lea, RG., Wood, MJ., Maheshwari, A., Bhattacharya, S., Collie-Duguid, ESR., Baker, PJ., Monteiro, A. & O'Shaughnessy, PJ. (2009). 'Gene expression analysis of human fetal ovarian primordial follicle formation'. Journal of Clinical Endocrinology and Metabolism, vol 94, no. 4, pp. 1427-1435.
    [Online] DOI: 10.1210/jc.2008-2619
  • Fowler, PA., Bhattacharya, S., Gromoll, J., Monteiro, A. & O'Shaughnessy, PJ. (2009). 'Maternal smoking and developmental changes in luteinizing hormone (LH) and the LH receptor in the fetal testis'. Journal of Clinical Endocrinology and Metabolism, vol 94, no. 12, pp. 4688-4695.
    [Online] DOI: 10.1210/jc.2009-0994
  • Fowler, PA., Dorà, NJ., McFerran, H., Amezaga, MR., Miller, DW., Lea, RG., Cash, P., McNeilly, AS., Evans, NP., Cotinot, C., Sharpe, RM. & Rhind, SM. (2008). 'In utero exposure to low doses of environmental pollutants disrupts fetal ovarian development in sheep'. Molecular Human Reproduction, vol 14, no. 5, pp. 269-280.
    [Online] DOI: 10.1093/molehr/gan020
  • Fowler, PA., Cassie, S., Rhind, SM., Brewer, MJ., Collinson, JM., Lea, RG., Baker, PJ., Bhattacharya, S. & O'Shaughnessy, PJ. (2008). 'Maternal smoking during pregnancy specifically reduces human fetal desert hedgehog gene expression during testis development'. Journal of Clinical Endocrinology and Metabolism, vol 93, no. 2, pp. 619-626.
    [Online] DOI: 10.1210/JC.2007-1860
  • Maheshwari, A. & Fowler, PA. (2008). 'Primordial follicular assembly in humans: revisited'. Zygote, vol 16, no. 4, pp. 285-296.
    [Online] DOI: 10.1017/S0967199408004802
    [Online] AURA: Maheshwari_Fowler_Zygote08_rev_prim_foll_form.pdf
  • Fowler, PAF., Tattum, J., Bhattacharya, S., Klonisch, T., Hombach-Klonisch, S., Gazvani, R., Lea, RG., Miller, ID., Simpson, W. & Cash, P. (2007). 'An investigation of the effects of endometriosis on the proteome of human eutopic endometrium: A heterogeneous tissue with a complex disease'. Proteomics, vol 7, no. 1, pp. 130-142.
    [Online] DOI: 10.1002/pmic.200600469
  • O'Shaughnessy, PJ., Baker, PJ., Montiero, A., Cassie, S., Bhattacharya, S. & Fowler, PAF. (2007). 'Developmental Changes in Human Fetal Testicular Cell Numbers and Messenger Ribonucleic Acid Levels during the Second Trimester'. Journal of Clinical Endocrinology and Metabolism, vol 92, no. 12, pp. 4792-4801.
    [Online] DOI: 10.1210/JC.2007-1690
  • Fowler, PAF., Dora, NJ., McFerran, HL., Sharpe, RM., McNeilly, AS., Lea, R., Miller, K., Cash, P. & Rhind, SM. (2007). 'Maternal dietary exposure to environmental chemicals disturbs fetal ovary development'. Human Reproduction, vol 22, no. Suppl. 1, O-237, pp. i95-i96.
    [Online] DOI: 10.1093/humrep/dem1059
  • Fowler, PAF., Abramovich, D., Haites, NE., Cash, P., Groome, NP., Al-Qahtani, A., Murray, TJ. & Lea, R. (2007). 'Human fetal testis Leydig cell disruption by exposure to the pesticide dieldrin at low concentrations'. Human Reproduction, vol 22, no. 11, pp. 2919-2927.
    [Online] DOI: 10.1093/humrep/dem256
  • Maheshwari, A., Fowler, PAF. & Bhattacharya, S. (2006). 'Assessment of ovarian reserve - should we perform tests of ovarian reserve'. Human Reproduction, vol 21, no. 11, pp. 2729-2735.
    [Online] DOI: 10.1093/humrep/del188
  • Lea, RG., Hannah, LT., Redmer, DA., Aitken, RP., Milne, J., Fowler, PAF., Murray, J. & Wallace, JM. (2005). 'Developmental indices of nutritionally-induced placental growth restriction in the adolescent sheep'. Pediatric Research, vol 57, no. 4, pp. 599-604.
    [Online] DOI: 10.1203/01.PDR.0000155949.08547.66
  • Hombach-Klonisch, S., Kehlen, A., Fowler, PAF., Huppertz, B., Jugert, JF., Bischoff, G., Schlüter, E., Buchmann, J. & Klonisch, T. (2005). 'Regulation of functional steroid receptors and ligand-induced responses in telomerase-immortalized human endometrial epithelial cells'. Journal of Molecular Endocrinology, vol 34, pp. 517-534.
    [Online] DOI: 10.1677/jme.1.01550
  • Sorsa-Leslie, TK., Mason, HD., Harris, WJ. & Fowler, PAF. (2005). 'Selection of gonadotrophin surge attenuating factor phage antibodies by bioassay'. Reproductive Biology and Endocrinology, vol 3, pp. 49-66.
    [Online] DOI: 10.1186/1477-7827-3-49
    [Online] AURA: Sorsa-Leslie_Selection_of_gonadotrophin_surge_attenuating_fa...
  • Fowler, PAF. & Spears, N. (2004). 'The cultured rodent follicle as a model for investigations of gonadotrophin surge-attenuating factor (GnSAF) production'. Reproduction, vol 127, no. 6, pp. 679-688.
    [Online] DOI: 10.1530/rep.1.00141
  • Fowler, PAF., Groome, NP. & Al-Gubory, KH. (2003). 'Demonstration of a non-steroidal, non-inhibin factor in the ovine corpus luteum of pregnancy that reduces pituitary responsiveness to GnRH-induced LH secretion in vitro'. Reproduction, vol 126, no. 1, pp. 35-42.
    [Online] DOI: 10.1530/rep.0.1260035
  • Gazvani, R., Smith, LM. & Fowler, PAF. (2002). 'Effect of interleukin-8 (IL-8), anti-IL-8, and IL-12 on endometrial cell survival in combined endometrial gland and stromal cell cultures derived from women with and without endometriosis'. Fertility and Sterility, vol 77, no. 1, pp. 62-67.
    [Online] DOI: 10.1016/S0015-0282(01)02954-5
  • Fowler, PAF., Sorsa-Leslie, TK., Cash, P., Dunbar, B., Melvin, WT., Wilson, Y., Mason, HD. & Harris, WJ. (2002). 'A 60-66 kDa protein with gonadotrophin surge attenuating factor bioactivity is produced by human ovarian granulosa cells'. Molecular Human Reproduction, vol 8, no. 9, pp. 823-832.
    [Online] DOI: 10.1093/molehr/8.9.823
  • Rae, MT., Rhind, SM., Fowler, PAF., Miller, DW., Kyle, CE. & Brooks, AN. (2002). 'Effect of maternal undernutrition on fetal testicular steroidogenesis during the CNS androgen-responsive period in male sheep fetuses'. Reproduction, vol 124, pp. 33-39.
    [Online] DOI: 10.1530/rep.0.1240033
  • Hawkins, CE., Dallas, JF., Fowler, PA.F., Woodroffe, R. & Racey, PA. (2002). 'Transient Masculinization in the Fossa, Cryptoprocta ferox (Carnivora, Viverridae)'. Biology of Reproduction, vol 66, no. 3, pp. 610-615.
    [Online] DOI: 10.1095/biolreprod66.3.610
  • Gazvani, R., Pakarinen, P., Fowler, PAF., Logan, S. & Huhtaniemi, I. (2002). 'Lack of association of the common immunologically anomalous LH with endometriosis'. Human Reproduction, vol 17, no. 6, pp. 1532-1534.
    [Online] DOI: 10.1093/humrep/17.6.1532
  • Gazvani, MR., Smith, LM., Haggarty, P., Fowler, PAF. & Templeton, AA. (2001). 'High omega-3:omega-6 fatty acid ratios in culture medium reduce endometrial-cell survival in combined endometrial gland and stromal cell cultures from women with and without endometriosis'. Fertility and Sterility, vol 76, no. 4, pp. 717-722.
    [Online] DOI: 10.1016/S0015-0282(01)01991-4
  • Fowler, PAF., Sorsa, T., Harris, WJ., Knight, PG. & Mason, HD. (2001). 'Relationship between follicle size and gonadotrophin surge attenuating factor (GnSAF) bioactivity during spontaneous cycles in women'. Human Reproduction, vol 16, no. 7, pp. 1353-1358.
    [Online] DOI: 10.1093/humrep/16.7.1353
  • Murray, TJ., Fowler, PAF., Haites, NE., Abramovich, D. & Lea, R. (1999). 'Proliferation and apoptosis in the developing human fetal testis: possible effects of endocrine disrupting chemicals (EDCs)'. Molecular and Cellular Endocrinology, vol 155, no. 1-2, pp. 187.
  • Fowler, PAF., Mason, HD. & Wilson, Y. (1999). 'Conclusive evidence for the production of a 60-66 kDa form of gonadotrophin surge-attenuating factor by human ovarian granulosa cells'. Human Reproduction, vol 14, pp. 17.
  • Knight, PG., Feist, SA., Tannetta, DS., Bleach, ECL., Fowler, PAF., O'Brien, M. & Groome, NP. (1998). 'Measurement of inhibin-A (alpha beta A dimer) during the oestrous cycle, after manipulation of ovarian activity and during pregnancy in ewes'. Journal of Reproduction and Fertility, vol 113, no. 1, pp. 159-166.
  • Fowler, PAF. & McNeilly, AS. (1997). 'Maternal pituitary gonadotroph function in relation to GnRH receptor and LH beta mRNA content during pregnancy in ewes'. Journal of Reproduction and Fertility, vol 110, no. 2, pp. 267-278.
  • Hardie, L., Trayhurn, P., Abramovich, D. & Fowler, PAF. (1997). 'Circulating leptin in women: a longitudinal study in the menstrual cycle and during pregnancy'. Clinical Endocrinology, vol 47, no. 1, pp. 101-106.
    [Online] DOI: 10.1046/j.1365-2265.1997.2441017.x
  • Fowler, PAF. & Price, AH. (1997). 'Follicle-stimulating hormone stimulates circulating gonadotropin surge-attenuating/inhibiting factor bioactivity in cows'. Biology of Reproduction, vol 57, no. 2, pp. 278-285.
  • Kazem, R., Messinis, LE., Fowler, PAF., Groome, NP., Knight, PG. & Templeton, A. (1996). 'Effect of mifepristone (RU486) on the pituitary response to gonadotrophin releasing hormone in women'. Human Reproduction, vol 11, no. 12, pp. 2585-2590.
  • Byrne, B., Fowler, PAF. & Templeton, A. (1996). 'The effects of steroidal and non-steroidal ovarian hormones on pituitary responsiveness to gonadotrophin surge-attenuating factor'. Journal of Endocrinology, vol 150, no. 3, pp. 413-422.
  • Muttukrishna, S., Fowler, PAF., George, L., Groome, NP. & Knight, GP. (1996). 'Changes in peripheral serum levels of total activin A during the human menstrual cycle and pregnancy'. Journal of Clinical Endocrinology and Metabolism, vol 81, no. 9, pp. 3328-3334.
  • Fowler, PAF., Danforth, D. & Templeton, A. (1996). 'Immunopurified human gonadotropin surge-attenuating factor (GnSAF/IF) bioactivity has a molecular mass of 60-70 kDa'. Biology of Reproduction, vol 54, pp. 144.
  • Mroueh, JM., Arbogast, LK., Fowler, PAF., Templeton, A., Friedman, CI. & Danforth, DR. (1996). 'Identification of gonadotrophin surge-inhibiting factor (GnSIF)/attenuin in human follicular fluid'. Human Reproduction, vol 11, no. 3, pp. 490-496.
  • Byrne, B., Fowler, PAF. & Templeton, A. (1996). 'Role of progesterone and nonsteroidal ovarian factors in regulating gonadotropin-releasing hormone self-priming in vitro'. Journal of Clinical Endocrinology and Metabolism, vol 81, no. 4, pp. 1454-1459.
  • Fowler, PAF., KNIGHT., P. & Templeton, A. (1995). 'Evidence for Gonadotropin Surge-Attenuating Factor (GNSAF) in Serum from Pregnant-Women'. Biology of Reproduction, vol 52, pp. 78.
  • BYRNE, B., Fowler, PAF. & Templeton, A. (1995). 'Gonadotropin Surge Attenuating Factor(GNSAF) Reduces Progesterone Augmentation of Unprimed GNRH-induced LH-secretion'. Biology of Reproduction, vol 52, no. Suppl. 1, pp. 78.
  • MUTTUKRISHNA, S., GEORGE, L., Fowler, PAF., GROOME., NP. & Knight, G. (1995). 'Measurement of Serum Concentrations og Inhibin-A (ALPHA-BETA(A) DIMER) During Human-Pregnancy'. Clinical Endocrinology, vol 42, no. 4, pp. 391-397.
  • BYRNE, B., Fowler, PAF., Fraser, M., CULLER, MD. & Templeton, A. (1995). 'Gonadotropin Surge-Attenuating Factor Bioactivity in Serum from Superovulated Women is not Blocked by Inhibin Antibody'. Biology of Reproduction, vol 52, no. 1, pp. 88-95.
    [Online] DOI: 10.1095/biolreprod52.1.88
  • Fowler, PAF., Fahy, U., Culler, MD., Knight, PG., Wardle, PG., McLaughlin, EA., Cunningham, P., Fraser, M., Hull, MGR. & Templeton, A. (1995). 'Gonadotropin surge-attenuating factor bioactivity is present in follicular-fluid from naturally cycling women'. Human Reproduction, vol 10, no. 1, pp. 68-74.
    [Online] DOI: 10.1093/humrep/10.1.68
  • Fowler, PAF., Fraser, M., CUNNINGHAM, P., Knight, PG., BYRNE., B., McLaughlin, EA., WARDLE., PG., HULL., MGR. & Templeton, A. (1994). 'Higher Gonadotropin Surge-Attenuating Factor Bioactivity is Found in Small Follicles from Superovulated Women'. Journal of Endocrinology, vol 143, no. 1, pp. 33-44.
  • Fowler, PAF., CUNNINGHAM., P., Fraser, M., MACGREGOR, F., BYRNE, B., PAPPAS, A., MESSINIS, IE. & Templeton, A. (1994). 'Circulating Gonadotropin Surge-Attenuating Factor From Superovulated Women Suppressing In-Vitro Gonadotropin-Releasing-Hormone Self-Priming'. Journal of Endocrinology, vol 143, no. 1, pp. 45-54.
  • MUTTUKRISHNA, S., Fowler, PAF., GROOME., NP., Mitchell, GG., Robertson, WR. & Knight, PG. (1994). 'Serum concentrations of dimeric inhibin during the spontaneous human menstrual cycle and after treatment with exogenous gonadotrophin'. Human Reproduction, vol 9, no. 9, pp. 1634-1642.
  • Fowler, PAF., MESSINIS., IE., CUNNINGHAM., P., Fraser, M. & Templeton, A. (1993). 'Effects of gonadotrophin surge-attenuating factor on the two pools of gonadotrophin-releasing hormone-induced luteinizing hormone secretion in vitro'. Human Reproduction, vol 8, no. 6, pp. 822-828.
  • Fowler, PAF., TOWNSEND., C., MESSINIS, IE., CUNNINGHAM., P. & Templeton, A. (1992). 'Gonadotropin Surge-Attenuating Factor Attenuates Invitro LH-Secretion Induced by Gonadotropin-Releasing-Hormone from Cultured Ovine Pituitary-Cells Only During the Breeding-Season'. Journal of Endocrinology, vol 135, no. 2, pp. 221-227.
  • GLASBEY, CA. & Fowler, PAF. (1992). 'Regression-models fitted using conditional independence to estimate pig fatness from magnetic-resonance images'. Journal of the Royal Statistical Society. Series D (The Statistician), vol 41, no. 2, pp. 179-184.
  • Fowler, PA., Fuller, MF., Glasbey, CA., Cameron, GG. & Foster, MA. (1992). 'Validation of the in vivo measurement of adipose tissue by magnetic resonance imaging of lean and obese pigs'. The American Journal of Clinical Nutrition, vol 56, no. 1, pp. 7-13.
  • Fowler, PAF., FULLER., MF., GLASBEY., CA., Foster, A., Cameron, G., McNeill, G. & Maughan, RJ. (1991). 'Total and Subcutaneous Adipose-Tissue in Women: the measurement of distribution and accurate prediction of quantity by using magnetic resonance imaging'. The American Journal of Clinical Nutrition, vol 54, no. 1, pp. 18-25.
  • McNeill, G., Fowler, PAF., Maughan, RJ., MCGAW., BA., FULLER., MF., GVOZDANOVIC., D. & GVOZDANOVIC, S. (1991). 'Body fat in lean and overweight women estimated by six methods'. British Journal of Nutrition, vol 65, no. 2, pp. 95-103.
  • Fowler, PAF., Knight, C. & Foster, MA. (1991). 'In-vivo magnetic resonance imaging studies of mammogenesis in non-pregnant goats treated with exogenous steroids'. Journal of Dairy Research, vol 58, no. 2, pp. 151-157.
    [Online] DOI: 10.1017/S0022029900029691
  • Fowler, PAF., KNIGHT., CH. & FOSTER, MA. (1991). 'Omitting the dry period between lactations does not reduce subsequent milk production in goats'. Journal of Dairy Research, vol 58, no. 1, pp. 13-19.
    [Online] DOI: 10.1017/S002202990003346X
  • FULLER, MF., Fowler, PAF., McNeill, G. & FOSTER, MA. (1990). 'Body composition: the precision and accuracy of new methods and their suitablity for longitudinal studies'. Proceedings of the Nutrition Society, vol 49, no. 3, pp. 423-436., 10.1079/PNS19900050
    [Online] DOI: 10.1079/PNS19900050
  • Fowler, PAF., MESSINIS., IE. & Templeton, A. (1990). 'Inhibition of LHRH-induced LH and FSH release by Gonadotropin Surge-Attenuating Factor(GNSAF) from Human Follicular-Fluid'. Journal of Reproduction and Fertility, vol 90, no. 2, pp. 587-594.
  • Knight, CJ., Fowler, PAF. & WILDE, CJ. (1990). 'Galactopoietic and Mammogenic Effects of Long-Term Treatment with Bovine Growth-Hormone and Thrice Daily Milking in Goats'. Journal of Endocrinology, vol 127, no. 1, pp. 129-138.
  • Fowler, PAF. & Racey, P. (1990). 'Daily and Seasonal Cycles of Body-Temperature and Aspects of Heterothermy in the Hedgehog Erinaceus-Europeas'. Journal of Comparative Physiology. B, Biochemical, Systemic, and Environmental Physiology, vol 160, no. 3, pp. 299-307.
  • Fowler, PAF., CASEY., CE., CAMERON., GG., FOSTER., MA. & KNIGHT, CH. (1990). 'Cyclic Changes in Composition and Volume of the Breast During the Menstrual-Cycle, Measured by Magnetic-Resonance-Imaging'. British Journal of Obstetrics and Gynaecology, vol 97, no. 7, pp. 595-602.
  • Fowler, PAF. & Racey, P. (1990). 'Effect of Melatonin Administration and Long Day-Length on Endocrine Cycles in the Hedgehog Erinaceus-Europaeus'. Journal of Pineal Research, vol 8, no. 3, pp. 193-204.
    [Online] DOI: 10.1111/j.1600-079X.1990.tb00679.x
  • Fowler, PAF., KNIGHT., CH., CAMERON., GG. & FOSTER, MA. (1990). 'Use of Magnetic-Resonance-Imaging in the Study of Goat Mammary-Glands Invivo'. Journal of Reproduction and Fertility, vol 89, no. 1, pp. 359-366.
  • Fowler, PAF., KNIGHT., CH., CAMERON., GG. & FOSTER, MA. (1990). 'Invivo Studies of Mammary Development in the Goat Using Magnetic-Resonance-Imaging (MRI)'. Journal of Reproduction and Fertility, vol 89, no. 1, pp. 367-375.
  • Fowler, PAF. & Racey, P. (1990). 'Effect of melatonin administration and long day-length on endocrine cycles in the hedgehog Erinaceus europaeus'. Journal of Pineal Research, vol 8, no. 1, pp. 67-78.
    [Online] DOI: 10.1111/j.1600-079X.1990.tb00808.x
  • Foster, MA., Fowler, PAF., Fuller, MF. & Knight, CH. (1988). 'Non-invasive methods for assessment of body-composition'. Proceedings of the Nutrition Society, vol 47, no. 3, pp. 375-385.
    [Online] DOI: 10.1079/PNS19880056
  • Fowler, PAF. (1988). 'Seasonal Endocrine Cycles in the European Hedgehog, Erinaceus-Europaeus'. Journal of Reproduction and Fertility, vol 84, no. 1, pp. 259-272.
  • BOAG, B. & Fowler, PAF. (1988). 'The Prevalence of Helminth-Parasites from the Hedgehog Erinaceus-Europaeus in Great-Britain'. Journal of Zoology, vol 215, no. 2, pp. 379-382.
  • Fowler, PAF. & Racey, P. (1988). 'Overwintering Strategies of the Badger, Meles-Meles, at 57-Degrees-N'. Journal of Zoology, vol 214, pp. 635-651.
  • Fowler, PAF. (1988). 'Thermoregulation in the Female Hedgehog, Erinaceus-Europaeus, During the Breeding-Season'. Journal of Reproduction and Fertility, vol 82, no. 1, pp. 285-292.
  • Fowler, PAF. & Racey, P. (1987). 'Relationship between body and testis temperatures in the European Hedgehog, Erinaceus-Europaeus, during hibernation and sexual reactivation'. Journal of Reproduction and Fertility, vol 81, no. 2, pp. 567-573.


  • Fowler, PA., O'Shaughnessy, PJ., Rhind, SM. & Ayres, J. (2009). 'PAH Exposure'. Environmental Health Perspectives, vol 117, no. 4, pp. A140; author reply A140-1.
    [Online] DOI: 10.1289/ehp.0800445

Comments and Debates

  • Fowler, PA. & O'Shaughnessy, PJ. (2013). 'The goldilocks principle and developmental androgens in males, what is "just right"?'. Endocrinology, vol 154, no. 5, pp. 1669-1671.
    [Online] DOI: 10.1210/en.2013-1279
  • Bergman, A., Andersson, A-M, Becher, G., van den Berg, M., Blumberg, B., Bjerregaard, P., Bornehag, C-G, Bornman, R., Brandt, I., Brian, JV., Casey, SC., Fowler, PA., Frouin, H., Giudice, LC., Iguchi, T., Hass, U., Jobling, S., Juul, A., Kidd, KA., Kortenkamp, A., Lind, M., Martin, OV., Muir, D., Ochieng, R., Olea, N., Norrgren, L., Ropstad, E., Ross, PS., Rudén, C., Scheringer, M., Skakkebaek, NE., Söder, O., Sonnenschein, C., Soto, A., Swan, S., Toppari, J., Tyler, CR., Vandenberg, LN., Vinggaard, AM., Wiberg, K. & Zoeller, RT. (2013). 'Science and policy on endocrine disrupters must not be mixed: a reply to a "common sense" intervention by toxicology journal editors'. Environmental Health, vol 12, no. 1, 69.
    [Online] DOI: 10.1186/1476-069X-12-69
    [Online] AURA: Bergman_et_al_EnvironHlth13_commentary_science_and_policy_vs...
  • Fowler, PAF. & Templeton, A. (1991). 'Ovarian Response to Gonadotropins: Effects of Growth-Hormone'. Clinical Endocrinology, vol 35, no. 2, pp. 117-118., 10.1111/j.1365-2265.1991.tb03507.x
    [Online] DOI: 10.1111/j.1365-2265.1991.tb03507.x

Scientific Reviews

  • O'Shaughnessy, PJ. & Fowler, PA. (2014). 'Development of the human fetal testis'. Annales d'Endocrinologie, vol 75, no. 2, pp. 48-53.
    [Online] DOI: 10.1016/j.ando.2014.03.009
  • Evans, NP., Bellingham, M., Sharpe, RM., Cotinot, C., Rhind, SM., Kyle, C., Erhard, H., Hombach-Klonisch, S., Lind, PM. & Fowler, PA. (2014). 'Does grazing on biosolids treated pasture pose a pathophysiological risk associated with increased exposure to endocrine disrupting compounds?'. Journal of Animal Science, vol 92, no. 8, pp. 3185-3198.
    [Online] DOI: 10.2527/jas.2014-7763
  • Stefansdottir, A., Fowler, PA., Powles-Glover, N., Anderson, RA. & Spears, N. (2014). 'Use of ovary culture techniques in reproductive toxicology'. Reproductive toxicology (Elmsford, N.Y.).
    [Online] DOI: 10.1016/j.reprotox.2014.08.001
  • Bellingham, M., Fiandanese, N., Byers, A., Cotinot, C., Evans, NP., Pocar, P., Amezaga, MR., Lea, RG., Sinclair, KD., Rhind, SM., Fowler, PA. & Fowler, PA.F (2012). 'Effects of exposure to environmental chemicals during pregnancy on the development of the male and female reproductive axes'. Reproduction in Domestic Animals, vol 47, no. Suppl. 4, pp. 15-22.
    [Online] DOI: 10.1111/j.1439-0531.2012.02050.x
  • O'Shaughnessy, P. & Fowler, PAF. (2011). 'Endocrinology of the mammalian fetal testis'. Reproduction, vol 141, pp. 37-46.
    [Online] DOI: 10.1530/REP-10-0365
  • Al-Gubory, KH., Fowler, PA. & Garrel, C. (2010). 'The roles of cellular reactive oxygen species, oxidative stress and antioxidants in pregnancy outcomes'. International Journal of Biochemistry & Cell Biology, vol 42, no. 10, pp. 1634-1650.
    [Online] DOI: 10.1016/j.biocel.2010.06.001
  • Klonisch, T., Fowler, PA. & Hombach-Klonisch, S. (2004). 'Molecular and genetic regulation of testis descent and external genitalia development'. Developmental Biology, vol 270, no. 1, pp. 1-18.
    [Online] DOI: 10.1016/j.ydbio.2004.02.018
  • Fowler, PAF., Sorsa-Leslie, TK., Harris, WJ. & Mason, HD. (2003). 'Ovarian gonadotrophin surge-attenuating factor (GnSAF): where are we after 20 years of research?'. Reproduction, vol 126, no. 6, pp. 689-699.
    [Online] DOI: 10.1530/rep.0.1260689
  • Fowler, PAF. & Huhtaniemi, IT. (2002). 'The ovarian gonadotropin receptors in health and disease'. Reviews in Endocrine and Metabolic Disorders, vol 3, no. 1, pp. 55-63.
    [Online] DOI: 10.1023/A:1012752819058
  • Fowler, PAF., Murray, TJ., Abramovich, D., Haites, NE. & Lea, RG. (2002). 'Environmental chemical effects on testicular function'. Reproductive Medicine Review, vol 10, no. 2, pp. 77-100.
    [Online] DOI: 10.1017/S0962279902000212
  • Murray, TJ., Lea, RG., Haites, NE., Abramovich, D. & Fowler, PAF. (2001). 'Endocrine disrupting chemicals'. Early Pregnancy, vol 5, no. 2, pp. 80-112.
  • Fowler, PAF. & Templeton, A. (1996). 'The nature and function of putative gonadotropin surge-attenuating inhibiting factor (GnSAF/IF)'. Reviews in Endocrine and Metabolic Disorders, vol 17, no. 2, pp. 103-120.
    [Online] DOI: 10.1210/edrv-17-2-103
  • FULLER, MF., Fowler, PAF., McNeill, G. & Foster, A. (1994). 'Imaging Techniques for the Assessment of Body-Composition'. The journal of nutrition, vol 124, no. 8, pp. S1546-S1550.
  • Fowler, PAF., Templeton, A. & MESSINIS, IE. (1993). 'The Ovarian Modulation of Gonadotropin-Releasing Hormone-Induced Luteinizing-Hormone Secretion in Women'. Human Reproduction, vol 8, no. Suppl. 2, pp. 112-116.

Chapters in Books, Reports and Conference Proceedings


  • Lea, RG., Byres, AS., Bellingham, M., Cotinot, C., Evans, N., Mandon-Pepin, B., Sinclair, KD. & Fowler, PA. (2014). 'Endocrine disruptors and ovine reproductive development'. in J Juengel, A Miyamoto, C Price, L Reynolds, M Smith & B Webb (eds), Reproduction in Domestic Ruminants VIII. vol. 8, Reproduction in Domestic Rumminants, Context Products Ltd, Leicestershire, UK, pp. 209-228.

Conference Proceedings

  • Templeton, A., Fowler, PAF. & Messinis, IE. (1992). 'Nonsteroidal control of gonadotropin secretion: the role of gonadotropin surge-attenuating factor'. in Hormones in gynecological endocrinology. PARTHENON PUBLISHING GROUP LTD, Lancaster, United Kingdom, pp. 125-133.
  • Foster, A., Fowler, PAF., Cameron, G., FULLER., M. & Knight, C. (1989). 'NMR Imaging Studies of Live Animals'. in E Kallweit & M Henning (eds), Application of NMR Techniques on the Body Composition of Live Animals. ELSEVIER APPL SCI PUBL LTD, Barking, Essex.

Peer-Reviewed Chapters

  • McKinlay, R., O'Shaughnessy, PJ., Sharpe, RM. & Fowler, PAF. (2012). 'In utero exposure to environmental chemicals: lessons from maternal cigarette smoking and its effects on gonad development and puberty'. in E Diamanti-Kandarakis & A Gore (eds), Endocrine Disruptors and Puberty: Contemporary Endocrinology. Humana Press, pp. 11.
    [Online] DOI: 10.1007/978-1-60761-561-3

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