Dr Alison Forhead

Senior Lecturer in Physiology and Medical Science

School of Biological and Medical Sciences

Teaching and supervision

Courses

Modules taught

Module leader in:

Human Structure and Function

Co-teacher in:

Scientific Skills
Professional and Experimental Skills
Research Methods
Integrated Physiology
Project

Research

My research interests are in mammalian endocrinology and the endocrine regulation of physiology and development in the fetus. Currently, my research work is focussed on two main areas:

Development and regulation of endocrine systems in the fetus
These studies have examined the bioavailability of hormones in the fetus, in particular, the glucocorticoids, thyroid hormones, leptin, insulin-like growth factors and the renin-angiotensin system. Our analyses of hormone systems in utero have included measurements of circulating concentrations, tissue metabolism and metabolic enzymes, cellular uptake mechanisms and receptor expression. These studies have demonstrated complex interactions between endocrine systems before birth and have elucidated a variety of cellular and molecular mechanisms of developmental control.

Endocrine control of fetal growth, development and maturation
My research has also investigated the role of hormones in the regulation of normal fetal development. These studies have an integrative approach to systems animal biology by examining a wide variety of fetal tissues and organs, and aspects of fetal physiology, including growth, cardiovascular and renal function, nutrition and metabolism. In particular, several of these studies have established the importance of endocrine signals in fetal maturation near to delivery and in the successful transition from the intrauterine to extrauterine environment at birth. We have investigated the mechanisms of glucocorticoid action in several physiological systems, and demonstrated the important roles of other hormones, such as thyroid hormones and angiotensin II, in mediating many of the maturational effects of glucocorticoids.

Overall, my research has an integrative approach to the study of mammalian endocrinology and systems animal physiology. The research findings have important implications for the understanding of normal fetal growth and development, the consequences of prematurity and fetal endocrine disorders, and the mechanisms underlying the intrauterine programming of adult (patho)physiology.

Research grants and awards

Main sources of funding: BBSRC, Isaac Newton Trust, The Royal Society

Groups

Fetal growth and development

Publications

Davies KL, Miles J, Camm EJ, Smith Dj, Barker P, Taylor K, Forhead AJ, Fowden AL, Prenatal cortisol exposure impairs adrenal function but not glucose metabolism in adult sheep
Journal of Endocrinology 260 (3) (2024) ISSN: 0022-0795 eISSN: 1479-6805 Prenatal cortisol exposure impairs adrenal function but not glucose metabolism in adult sheep Open Access version on RADAR
Young R, Lewandowska D, Long E, Wooding FBP, De Blasio MJ, Davies KL, Camm EJ, Sangild PT, Fowden AL, Forhead AJ, Hypothyroidism impairs development of the gastrointestinal tract in the ovine fetus
Frontiers in Physiology 14 (2023) ISSN: 1664-042X eISSN: 1664-042X Hypothyroidism impairs development of the gastrointestinal tract in the ovine fetus Open Access version on RADAR
Davies KL, Camm EJ, Smith DJ, Miles J, Forhead AJ, Murray AJ, Fowden AL, Developmental programming of mitochondrial substrate metabolism in skeletal muscle of adult sheep by cortisol exposure before birth
Journal of Developmental Origins of Health and Disease 14 (1) (2022) pp.77-87 ISSN: 2040-1744 eISSN: 2040-1752
Davies KL, Camm EJ, Smith DJ, Miles J, Forhead AJ, Murray AJ, Fowden AL, Developmental programming of mitochondrial substrate metabolism in skeletal muscle of adult sheep by cortisol exposure before birth
Journal of Developmental Origins of Health and Disease 14 (1) (2022) pp.77-87 ISSN: 2040-1744 eISSN: 2040-1752 Developmental programming of mitochondrial substrate metabolism in skeletal muscle of adult sheep by cortisol exposure before birth Open Access version on RADAR
Davies KL, Smith DJ, El-Bacha T, Wooding FBP, Forhead AJ, Murray AJ, Fowden AL, Camm EJ , Cortisol regulates cerebral mitochondrial oxidative phosphorylation and morphology of the brain in a region-specific manner in the ovine fetus
Biomolecules 12 (6) (2022) ISSN: 2218-273X eISSN: 2218-273X Cortisol regulates cerebral mitochondrial oxidative phosphorylation and morphology of the brain in a region-specific manner in the ovine fetus Open Access version on RADAR
Wooding FBP, Forhead AJ, Wilsher S, Allen WR, Roberts RM, Green JA, Beckers JF, Melo de Sousa N, Charpigny G , Asymmetric expression of proteins in the granules of the placentomal binucleate cells in Giraffa Camelopardalis
Biology of Reproduction 106 (4) (2022) pp.814-822 ISSN: 0006-3363 eISSN: 1529-7268 Asymmetric expression of proteins in the granules of the placentomal binucleate cells in Giraffa Camelopardalis Open Access version on RADAR
Lanham Stuart A, Blache Dominique, Oreffo Richard OC, Fowden Abigail L, Forhead Alison J, Pancreas deficiency modifies bone development in the ovine fetus near term
Journal of Endocrinology 252 (1) (2021) pp.71-80 ISSN: 0022-0795 eISSN: 1479-6805 Pancreas deficiency modifies bone development in the ovine fetus near term Open Access version on RADAR
Davies KL, Camm EJ, Smith DJ, Vaughan OR, Forhead AJ, Murray AJ, Fowden AL, Glucocorticoid maturation of mitochondrial respiratory capacity in skeletal muscle before birth
Journal of Endocrinology 251 (1) (2021) pp.53-68 ISSN: 0022-0795 eISSN: 1479-6805 Glucocorticoid maturation of mitochondrial respiratory capacity in skeletal muscle before birth Open Access version on RADAR
Davies Katie L., Smith Danielle J., El‐Bacha Tatiana, Stewart Max E., Easwaran Akshay, Wooding Peter F. P., Forhead Alison J., Murray Andrew J., Fowden Abigail L., Camm Emily J., Development of cerebral mitochondrial respiratory function is impaired by thyroid hormone deficiency before birth in a region‐specific manner
The FASEB Journal 35 (5) (2021) ISSN: 0892-6638 eISSN: 1530-6860 Development of cerebral mitochondrial respiratory function is impaired by thyroid hormone deficiency before birth in a region‐specific manner Open Access version on RADAR
Camm EJ, Inzani I, De Blasio MJ, Davies KL, Lloyd IR, Wooding FBP, Blache D, Fowden AL, Forhead AJ, Thyroid hormone deficiency suppresses fetal pituitary-adrenal function near term: implications for the control of fetal maturation and parturition
Thyroid 31 (6) (2020) pp.861-869 ISSN: 1050-7256 eISSN: 1557-9077 Thyroid hormone deficiency suppresses fetal pituitary-adrenal function near term: implications for the control of fetal maturation and parturition Open Access version on RADAR
Valenzuela OA, Jellyman JK, Allen V, Niu Y, Holdstock NB, Forhead AJ, Giussani DA, Fowden AL, Herrera EA, Neonatal glucocorticoid overexposure alters cardiovascular function in young adult horses in a sex-linked manner
Journal of Developmental Origins of Health and Disease 12 (2) (2020) pp.309-318 ISSN: 2040-1744 eISSN: 2040-1752 Neonatal glucocorticoid overexposure alters cardiovascular function in young adult horses in a sex-linked manner Open Access version on RADAR
Davies KL, Camm JE, Atkinson E, Lopez T, Forhead AJ, Murray AJ, Fowden AL, Development and thyroid hormone dependence of skeletal muscle mitochondrial function towards birth
The Journal of Physiology 598 (12) (2020) pp.2453-2468 ISSN: 0022-3751 eISSN: 1469-7793 Development and thyroid hormone dependence of skeletal muscle mitochondrial function towards birth Open Access version on RADAR
Harris SE, Blasio MJ, Zhao X, Ma M, Davies KL, Wooding FBP, Hamilton RS, Blache D, Meredith D, Murray AJ, Fowden AL, Forhead AJ, Thyroid deficiency before birth alters the adipose transcriptome to promote overgrowth of white adipose tissue and impair thermogenic capacity
Thyroid 30 (6) (2020) pp.794-805 ISSN: 1050-7256 eISSN: 1557-9077) Thyroid deficiency before birth alters the adipose transcriptome to promote overgrowth of white adipose tissue and impair thermogenic capacity Open Access version on RADAR
De Blasio MJ, Lanham SA, Blache D, Oreffo ROC, Fowden AL, Forhead AJ, Sex- and bone-specific responses in bone structure to exogenous leptin and leptin receptor antagonism in the ovine fetus
American Journal of Physiology - Regulatory, Integrative and Comparative Physiology 314 (6) (2018) pp.R781-R790 ISSN: 0363-6119 eISSN: 1522-1490 Sex- and bone-specific responses in bone structure to exogenous leptin and leptin receptor antagonism in the ovine fetus Open Access version on RADAR
Franko KL, Forhead AJ, Fowden AL, Effects of stress during pregnancy on hepatic glucogenic capacity in rat dams and their fetuses
Physiological Reports 5 (11) (2017) ISSN: 2051-817X eISSN: 2051-817X Effects of stress during pregnancy on hepatic glucogenic capacity in rat dams and their fetuses Open Access version on RADAR
Harris SE, De Blasio MJ, Davis MA, Kelly A, Davenport HM, Wooding FBP, Blache D, Meredith D, Anderson M, Fowden AL, Limesand SW, Forhead AJ, Hypothyroidism in utero stimulates pancreatic beta cell proliferation and hyperinsulinaemia in the ovine fetus during late gestation
The Journal of Physiology 595 (11) (2017) pp.3331-3343 ISSN: 0022-3751 eISSN: 1469-7793 Hypothyroidism in utero stimulates pancreatic beta cell proliferation and hyperinsulinaemia in the ovine fetus during late gestation Open Access version on RADAR
Valenzuela OA, Jellyman JK, Allen VL, Holdstock NB, Forhead AJ, Fowden AL, Effects of birth weight, sex and neonatal glucocorticoid overexposure on glucose-insulin dynamics in young adult horses
Journal of Developmental Origins of Health and Disease 8 (2) (2017) pp.206-215 ISSN: 2040-1744 Effects of birth weight, sex and neonatal glucocorticoid overexposure on glucose-insulin dynamics in young adult horses Open Access version on RADAR
de Blasio M, Boije M, Kempster SL, Smith GCS, Charnock-Jones S, Denyer A, Hughes A, Wooding FBP, Blache D, Fowden AL, Forhead A, Leptin matures aspects of lung structure and function in the ovine fetus
Endocrinology 157 (1) (2016) pp.395-404 ISSN: 0013-7227 eISSN: 1945-7170 Leptin matures aspects of lung structure and function in the ovine fetus Open Access version on RADAR
Forhead A, Jellyman J, De Blasio M, Johnson E, Giussani D, Pipkin F, Fowden A, Maternal dexamethasone treatment alters tissue and circulating components of the renin-angiotensin system in the pregnant ewe and fetus
Endocrinology 156 (8) (2015) pp.3038-3046 ISSN: 0013-7227 eISSN: 1945-7170 Maternal dexamethasone treatment alters tissue and circulating components of the renin-angiotensin system in the pregnant ewe and fetus Open Access version on RADAR
De Blasio M, Boije M, Vaughan OR, Bernstein BS, Davies KL, Plein A, Kempster SL, Smith GCS, Charnock-Jones DS, Blache D, Wooding F, Giussani DA, Fowden AL, Forhead AJ, Developmental expression and glucocorticoid control of the leptin receptor in fetal ovine lung
PLoS ONE 10 (8) (2015) ISSN: 1932-6203
Fowden AL, Forhead AJ, Sferruzzi-Perri AN, Burton GJ, Vaughan OR, Endocrine regulation of placental phenotype
Placenta 36 (Supplement 1) (2015) pp.s50-s59 ISSN: 0143-4004 eISSN: 1532-3102
Jellyman J, Valenzuela O, Allen V, Forhead A, Holdstock N., Fowden A., Neonatal glucocorticoid overexposure programs pituitary-adrenal function in ponies
Domestic Animal Endocrinology 50 (2015) pp.45-49 ISSN: 0739-7240
Wooding FBP, Kimura J, Forhead AJ, Functional immunocytochemistry of Tragulus placenta: Implications for ruminant evolution
Placenta 35 (5) (2014) pp.305-310 ISSN: 0143-4004 eISSN: 1532-3102
Forhead A, Fowden A, Thyroid hormones in fetal growth and prepartum maturation
Journal of Endocrinology 221 (3) (2014) pp.R87-R103 ISSN: 0022-0795
Sferruzzi-Perri A, Vaughan O, Forhead A, Fowden A, Hormonal and nutritional drivers of intrauterine growth
Current Opinion in Clinical Nutrition and Metabolic Care 16 (3) (2013) pp.298-309 ISSN: 1363-1950 eISSN: 1473-6519
Fowden A, Jellyman J, Valenzuela O, Forhead A, Nutritional Programming of Intrauterine Development: A Concept Applicable to the Horse?
Journal of Equine Veterinary Science 33 (5) (2013) pp.295-304 ISSN: 0737-0806
Fowden A, Forhead A, Ousey J, Endocrine adaptations in the foal over the perinatal period
Equine Veterinary Journal 44 (s41) (2012) pp.130-139 ISSN: 0425-1644 eISSN: 2042-3306
Jellyman J, Allen V, Forhead A, Holdstock N, Fowden A, Hypothalamic-pituitary-adrenal axis function in pony foals after neonatal ACTH-induced glucocorticoid overexposure
Equine Veterinary Journal 44 (s41) (2012) pp.38-42 ISSN: 0425-1644 eISSN: 2042-3306
Fowden A, Forhead A, Insulin deficiency alters the metabolic and endocrine responses to undernutrition in fetal sheep near term
Endocrinology 153 (8) (2012) pp.4008-4018 ISSN: 0013-7227 eISSN: 1945-7170
Belteki G, Kempster S, Forhead A, Giussani D, Fowden A, Curley A, Charnock-Jones D, Smith G, Paraoxonase-3, a Putative Circulating Antioxidant, Is Systemically Up-Regulated in Late Gestation in the Fetal Rat, Sheep, and Human
The Journal of Clinical Endocrinology & Metabolism 95 (8) (2011) pp.3798-3805 ISSN: 0021-972X eISSN: 1945-7197
Kempster S, Belteki G, Forhead A, Fowden A, Catalano R, Lam B, McFarlane I, Charnock-Jones D, Developmental control of the Nlrp6 inflammasome and a substrate, IL-18, in mammalian intestine
American Journal of Physiology - Gastrointestinal and Liver Physiology 300 (2) (2011) pp.G253-G263 ISSN: 0193-1857 eISSN: 1522-1547
Fowden A, Forhead A, Adrenal glands are essential for activation of glucogenesis during undernutrition in fetal sheep near term
Endocrinology and Metabolism 300 (1) (2011) pp.E94-E102 ISSN: 0193-1849
Lanham S, Fowden A, Roberts C, Cooper C, Oreffo R, Forhead A, Effects of hypothyroidism on the structure and mechanical properties of bone in the ovine fetus
Journal of Endocrinology 210 (2011) pp.189-198 ISSN: 0022-0795 eISSN: 1479-6805
Forhead A, Jellyman J, Gillham K, Ward J, Blache D, Fowden A, Renal growth retardation following angiotensin II type 1 (AT 1) receptor antagonism is associated with increased at 2 receptor protein in fetal sheep
Journal of Endocrinology 208 (2) (2011) pp.137-145 ISSN: 0022-0795 eISSN: 1479-6805
Fowden A, Ward J, Wooding F, Forhead A, Developmental programming of the ovine placenta.
Society of Reproduction and Fertility Supplement 67 (2010) pp.41-57 ISSN: 1747-3403
Franko K, Forhead A, Fowden A, Differential effects of prenatal stress and glucocorticoid administration on postnatal growth and glucose metabolism in rats
Journal of Endocrinology 204 (3) (2010) pp.319-329 ISSN: 0022-0795
Wooding F, Ramirez-Pinilla M, Forhead A, Functional studies of the placenta of the lizard Mabuya sp. (Scincidae) using immunocytochemistry
Placenta 31 (8) (2010) pp.675-685 ISSN: 0143-4004 eISSN: 1532-3102
Forhead A, Cutts S, Matthews P, Fowden A, Role of thyroid hormones in the developmental control of tissue glycogen in fetal sheep near term
Experimental Physiology 94 (10) (2009) pp.1079-1087 ISSN: 0958-0670
Franko K, Forhead A, Fowden A, Effects of maternal dietary manipulation during different periods of pregnancy on hepatic glucogenic capacity in fetal and pregnant rats near term
Nutrition, Metabolism & Cardiovascular Diseases 19 (8) (2009) pp.555-562 ISSN: 0939-4753
Fowden A, Forhead A, Endocrine Regulation of Feto-Placental Growth
Hormone Research 72 (5) (2009) pp.257-265 ISSN: 0301-0163 eISSN: 1423-0046
Fowden A, Forhead A, Hormones as epigenetic signals in developmental programming
Experimental Physiology 94 (6) (2009) pp.607-625 ISSN: 0958-0670
Forhead A, Fowden A, The hungry fetus? Role of leptin as a nutritional signal before birth
The Journal of Physiology 587 (6) (2009) pp.1145-1152 ISSN: 0022-3751
Holmes K, Charnock Jones S, Forhead A, Giussani D, Fowden A, Licence D, Kempster S, Smith G, Localization and control of expression of VEGF-A and the VEGFR-2 receptor in fetal sheep intestines
Pediatric Research 63 (2) (2008) pp.143-148 ISSN: 0031-3998 eISSN: 1530-0447
Fowden A, Forhead A, Ousey J, The endocrinology of equine parturition
Experimental and Clinical Endocrinology & Diabetes 116 (7) (2008) pp.393-403 ISSN: 0947-7349 eISSN: 1439-3646
Fowden A, Forhead A, Coan P, Burton G, The placenta and intrauterine programming
Journal of Neuroendocrinology 20 (4) (2008) pp.439-450 ISSN: 0953-8194
Wooding F, Kimura J, Fukuta K, Forhead A, A Light and Electron Microscopical Study of the Tragulid (Mouse Deer) Placenta
Placenta 28 (10) (2007) pp.1039-1048 ISSN: 0143-4004 eISSN: 1532-3102
O'Connor D, Blache D, Hoggard N, Brookes E, Wooding F, Fowden A, Forhead A, Developmental control of plasma leptin and adipose leptin messenger ribonucleic acid in the ovine fetus during late gestation: Role of glucocorticoids and thyroid hormones
Endocrinology 148 (8) (2007) pp.3750-3757 ISSN: 0013-7227 eISSN: 1945-7170
Forhead A, Jellyman J, Gardner D, Giussani D, Kaptein E, Visser T, Fowden A, Differential effects of maternal dexamethasone treatment on circulating thyroid hormone concentrations and tissue deiodinase activity in the pregnant ewe and fetus
Endocrinology 148 (2) (2007) pp.800-805 ISSN: 0013-7227 eISSN: 1945-7170
Franko K, Giussani D, Forhead A, Fowden A, Effects of dexamethasone on the glucogenic capacity of fetal, pregnant, and non-pregnant adult sheep
Journal of Endocrinology 192 (1) (2007) pp.67-73 ISSN: 0022-0795
Fowden A, Forhead A, Effects of pituitary hormone deficiency on growth and glucose metabolism of the sheep fetus
Endocrinology 148 (10) (2007) pp.4812-4820 ISSN: 0013-7227 eISSN: 1945-7170
Wooding F, Dantzer V, Klisch K, Jones C, Forhead A, Glucose Transporter 1 Localisation Throughout Pregnancy in the Carnivore Placenta: Light and Electron Microscope Studies
Placenta 28 (5-6) (2007) pp.453-464 ISSN: 0143-4004 eISSN: 1532-3102
Forhead A., Curtis K., Kaptein E., Visser T., Fowden A., Developmental control of iodothyronine deiodinases by cortisol in the ovine fetus and placenta near term
Endocrinology 147 (12) (2006) pp.5988-5994 ISSN: 0013-7227 eISSN: 1945-7170
Ward J., Forhead A., Wooding F., Fowden A., Functional significance and cortisol dependence of the gross morphology of ovine placentomes during late gestation
Biology of Reproduction 74 (1) (2006) pp.137-145 ISSN: 0006-3363 eISSN: 1529-7268
Fowden A., Giussani D., Forhead A., Intrauterine programming of physiological systems: Causes and consequences
Physiology 21 (1) (2006) pp.29-37 ISSN: 1548-9213
Gnanalingham M., Giussani D., Sivathondan P., Forhead A., Stephenson T., Symonds M., Gardner D., Chronic umbilical cord compression results in accelerated maturation of lung and brown adipose tissue in the sheep fetus during late gestation
Endocrinology and Metabolism 289 (2005) ISSN: 0193-1849
Giussani D., Forhead A., Fowden A., Development of cardiovascular function in the horse fetus
The Journal of Physiology 565 (3) (2005) pp.1019-1030 ISSN: 0022-3751
Fletcher A., Forhead A., Fowden A., Ford W., Nathanielsz P., Giussani D., Effects of gestational age and cortisol treatment on ovine fetal heart function in a novel biventricular Langendorff preparation
The Journal of Physiology 562 (2) (2005) pp.493-505 ISSN: 0022-3751
Fowden A., Giussani D., Forhead A., Endocrine and metabolic programming during intrauterine development
Early Human Development 81 (9) (2005) pp.723-734 ISSN: 0378-3782
Gnanalingham M., Mostyn A., Forhead A., Fowden A., Symonds M., Stephenson T., Increased uncoupling protein-2 mRNA abundance and glucocorticoid action in adipose tissue in the sheep fetus during late gestation is dependent on plasma cortisol and triiodothyronine
The Journal of Physiology 567 (1) (2005) pp.283-292 ISSN: 0022-3751
Fowden A., Gardner D., Ousey J., Giussani D., Forhead A., Maturation of pancreatic β-cell function in the fetal horse during late gestation
Journal of Endocrinology 186 (3) (2005) pp.467-473 ISSN: 0022-0795
Fahmi A., Forhead A., Fowden A., Vandenberg J., Cortisol influences the ontogeny of both α- and β-subunits of the cardiac sodium channel in fetal sheep
Journal of Endocrinology 180 (3) (2004) pp.449-455 ISSN: 0022-0795 eISSN: 1479-6805
Fowden A., Forhead A., Endocrine mechanisms of intrauterine programming
Reproduction 127 (5) (2004) pp.515-526 ISSN: 1470-1626 eISSN: 1741-7899
Forhead A., Ousey J., Allen W., Fowden A., Postnatal insulin secretion and sensitivity after manipulation of fetal growth by embryo transfer in the horse
Journal of Endocrinology 181 (3) (2004) pp.459-467 ISSN: 0022-0795 eISSN: 1479-6805
Forhead A., Fowden A., Role of angiotensin II in the pressor response to cortisol in fetal sheep during late gestation
Experimental Physiology 89 (3) (2004) pp.323-329 ISSN: 0958-0670
Forhead A, Poore K, Mapstone J, Fowden A, Developmental regulation of hepatic and renal gluconeogenic enzymes by thyroid hormones in fetal sheep during late gestation
The Journal of Physiology 548 (3) (2003) pp.941-947 ISSN: 0022-3751
Zimmermann H., Gardner D., Jellyman J., Fowden A., Giussani D., Forhead A., Effect of dexamethasone on pulmonary and renal angiotensin-converting enzyme concentration in fetal sheep during late gestation
American Journal of Obstetrics & Gynecology 189 (5) (2003) pp.1467-1471 ISSN: 0002-9378 eISSN: 1097-6868
Lisle S., Lewis R., Petry C., Ozanne S., Hales C., Forhead A., Effect of maternal iron restriction during pregnancy on renal morphology in the adult rat offspring
British Journal of Nutrition 90 (1) (2003) pp.33-39 ISSN: 0007-1145
Mostyn A., Pearce S., Budge H., Elmes M., Forhead A., Fowden A., Stephenson T., Symonds M., Influence of cortisol on adipose tissue development in the fetal sheep during late gestation
Journal of Endocrinology 176 (1) (2003) pp.23-30 ISSN: 0022-0795 eISSN: 1479-6805
Ousey J., Forhead A., Rossdale P., Grainger L., Houghton E., Fowden A., Ontogeny of uteroplacental progestagen production in pregnant mares during the second half of gestation
Biology of Reproduction 69 (2) (2003) pp.540-548 ISSN: 0006-3363 eISSN: 1529-7268
Coomer R., Forhead A., Bathe A., Head M., Plasma angiotensin-converting enzyme (ACE) concentration in Thoroughbred racehorses
Equine Veterinary Journal 35 (1) (2003) pp.96-98 ISSN: 0425-1644 eISSN: 2042-3306
Giussani D., Forhead A., Gardner D., Fletcher A., Allen W., Fowden A., Postnatal cardiovascular function after manipulation of fetal growth by embryo transfer in the horse
The Journal of Physiology 547 (1) (2003) pp.67-76 ISSN: 0022-3751
Li J., Forhead A., Dauncey M., Gilmour R., Fowden A., Control of growth hormone receptor and insulin-like growth factor-I expression by cortisol in ovine fetal skeletal muscle
The Journal of Physiology 541 (2) (2002) pp.581-589 ISSN: 0022-3751
O'Connor S., Fowden A., Holdstock N., Giussani D., Forhead A., Development changes in pulmonary and renal angiotensin-converting enzyme concentration in fetal and neonatal horses
Reproduction, Fertility and Development 14 (2002) pp.413-417 ISSN: 1031-3613 eISSN: 1448-5990
Forhead A., Fowden A., Effects of thyroid hormones on pulmonary and renal angiotensin-converting enzyme concentrations in fetal sheep near term
Journal of Endocrinology 173 (1) (2002) pp.143-150 ISSN: 0022-0795 eISSN: 1479-6805
Lewis R., Forhead A., Petry C., Ozanne S., Nicolas Hales C., Long-term programming of blood pressure by maternal dietary iron restriction in the rat
British Journal of Nutrition 88 (3) (2002) pp.283-290 ISSN: 0007-1145
Forhead A., Thomas L., Crabtree J., Hoggard N., Gardner D., Giussani D., Fowden A., Plasma leptin concentration in fetal sheep during late gestation: Ontogeny and effect of glucocorticoids
Endocrinology 143 (4) (2002) pp.1166-1173 ISSN: 0013-7227 eISSN: 1945-7170
Clarke K., Ward J., Forhead A., Giussani D., Fowden A., Regulation of 11β-hydroxysteroid dehydrogenase type 2 activity in ovine placenta by fetal cortisol
Journal of Endocrinology 172 (3) (2002) pp.527-534 ISSN: 0022-0795 eISSN: 1479-6805
Poore K, Forhead A, Gardner D, Giussani D, Fowden A, The effects of birth weight on basal cardiovascular function in pigs at 3 months of age
The Journal of Physiology 539 (3) (2002) pp.969-978 ISSN: 0022-3751
Forhead A., Li J., Gilmour R., Dauncey M., Fowden A., Thyroid hormones and the mRNA of the GH receptor and IGFs in skeletal muscle of fetal sheep
Endocrinology and Metabolism 282 (2002) ISSN: 0193-1849
Fowden A., Ousey J., Forhead A., Rossdale P., Grainger L., Houghton E., Uteroplacental production of 5α-pregnane-3,20-dione (5αDHP) in pregnant mares
Theriogenology 58 (2002) pp.821-824 ISSN: 0093-691X eISSN: 1879-3231
Fowden A., Ousey J., Forhead A., Comparative aspects of prepartum maturation: Provision of nutrients
Pferdeheilkunde 17 (6) (2001) pp.653-658 ISSN: 0177-7726
Forhead A, Gulati V, Poore K, Fowden A, Ontogeny of pulmonary and renal angiotensin-converting enzyme in pigs
Molecular and Cellular Endocrinology 185 (1-2) (2001) pp.127-133 ISSN: 0303-7207
Fowden A., Mapstone J., Forhead A., Regulation of glucogenesis by thyroid hormones in fetal sheep during late gestation
Journal of Endocrinology 170 (2) (2001) pp.461-469 ISSN: 0022-0795 eISSN: 1479-6805
Sangild P., Schmidt M., Jacobsen H., Fowden A., Forhead A., Avery B., Greve T., Blood chemistry, nutrient metabolism, and organ weights in fetal and newborn calves derived from in vitro-produced bovine embryos
Biology of Reproduction 62 (6) (2000) pp.1495-1504 ISSN: 0006-3363 eISSN: 1529-7268
Forhead A., Li J., Saunders J., Dauncey M., Gilmour R., Fowden A., Control of ovine hepatic growth hormone receptor and insulin-like growth factor I by thyroid hormones in utero
Endocrinology and Metabolism 278 (2000) ISSN: 0193-1849
Forhead A., Broughton Pipkin F., Taylor P., Baker K., Balouzet V., Giussani D., Fowden A., Developmental changes in blood pressure and the renin-angiotensin system in pony fetuses during the second half of gestation.
Journal of Reproduction and Fertility. Supplement (56) (2000) pp.693-703 ISSN: 0449-3087
Forhead A., Broughton Pipkin F., Fowden A., Effect of cortisol on blood pressure and the renin-angiotensin system in fetal sheep during late gestation
The Journal of Physiology 526 (1) (2000) pp.167-176 ISSN: 0022-3751
Fowden A., Forhead A., White K., Taylor P., Equine uteroplacental metabolism at mid- and late gestation
Experimental Physiology 85 (5) (2000) pp.539-545 ISSN: 0958-0670
Fletcher A., Goodfellow M., Forhead A., Gardner D., Mcgarrigle H., Fowden A., Giussani D., Low doses of dexamethasone suppress pituitary-adrenal function but augment the glycemic response to acute hypoxemia in fetal sheep during late gestation
Pediatric Research 47 (5) (2000) pp.684-691 ISSN: 0031-3998 eISSN: 1530-0447
Fowden A., Taylor P., White K., Forhead A., Ontogenic and nutritionally induced changes in fetal metabolism in the horse
The Journal of Physiology 528 (1) (2000) pp.209-219 ISSN: 0022-3751
Forhead A., Gillespie C., Fowden A., Role of cortisol in the ontogenic control of pulmonary and renal angiotensin-converting enzyme in fetal sheep near term
The Journal of Physiology 526 (2) (2000) pp.409-416 ISSN: 0022-3751
Fowden A., Forhead A., Bloomfield M., Taylor P., Silver M., Pancreatic α cell function in the fetal foal during late gestation
Experimental Physiology 84 (4) (1999) pp.697-705 ISSN: 0958-0670
Forhead A., Li J., Gilmour R., Fowden A., Control of hepatic insulin-like growth factor II gene expression by thyroid hormones in fetal sheep near term
Endocrinology and Metabolism 275 (1998) ISSN: 0193-1849
Forhead A., Melvin R., Balouzet V., Fowden A., Developmental changes in plasma angiotensin-converting enzyme concentration in fetal and neonatal lambs
Reproduction, Fertility and Development 10 (5) (1998) pp.393-398 ISSN: 1031-3613 eISSN: 1448-5990
Fowden A., Li J., Forhead A., Glucocorticoids and the preparation for life after birth: Are there long-term consequences of the life insurance?
Proceedings of the Nutrition Society 57 (1) (1998) pp.113-122 ISSN: 0029-6651
Fowden A., Li J., Forhead A., Sliver M., Hormones as nutritional signals during intrauterine development
Equine Veterinary Journal 30 (6) (1998) pp.464- ISSN: 0425-1644 eISSN: 2042-3306
Forhead A., Pipkin F., Sutherland M., Fowden A., Changes in the maternal and fetal renin-angiotensin systems in response to angiotensin II type 1 receptor blockade and angiotensin-converting enzyme inhibition in pregnant sheep during late gestation
Experimental Physiology 82 (4) (1997) pp.761-776 ISSN: 0958-0670
Fowden A., Forhead A., Silver M., Macdonald A., Glucose, lactate and oxygen metabolism in the fetal pig during late gestation
Experimental Physiology 82 (1) (1997) pp.171-182 ISSN: 0958-0670
Forhead A., Dobson H., Plasma glucose and cortisol responses to exogenous insulin in fasted donkeys
Research in Veterinary Science 62 (3) (1997) pp.265-269 ISSN: 0034-5288
Forhead A., Whybrew K., Hughes P., Broughton Pipkin F., Sutherland M., Fowden A., Comparison of angiotensin II type 1 receptor blockade and angiotensin-converting enzyme inhibition in pregnant sheep during late gestation
British Journal of Pharmacology 119 (2) (1996) pp.393-401 ISSN: 0007-1188
Fowden A., Szemere J., Hughes P., Gilmour R., Forhead A., The effects of cortisol on the growth rate of the sheep fetus during late gestation
Journal of Endocrinology 151 (1) (1996) pp.97-105 ISSN: 0022-0795
Forhead A., Fowden A., Silver M., Hughes P., Broughton-Pipkin F., Sutherland M., Haemodynamic responses to an angiotensin II receptor antagonist (GR117289) in maternal and fetal sheep
Experimental Physiology 80 (2) (1995) pp.285-298 ISSN: 0958-0670
Forhead A., Smart D., Smith R., Dobson H., Transport-induced stress responses in fed and fasted donkeys
Research in Veterinary Science 58 (2) (1995) pp.144-151 ISSN: 0034-5288
Forhead A., Fowden A., Silver M., Broughton-Pipkin F., Angiotensin II antagonism in sheep during late gestation
Early Human Development 38 (1) (1994) pp.63- ISSN: 0378-3782
Forhead A., Smart D., Smith R., Dobson H., Endocrine and metabolic responses to transportation in periparturient donkeys
Animal Reproduction Science 36 (1994) pp.315-327 ISSN: 0378-4320 eISSN: 1873-2232
Forhead A., French J., Ikin P., Fowler J., Dobson H., Relationship between plasma insulin and triglyceride concentrations in hypertriglyceridaemic donkeys
Research in Veterinary Science 56 (3) (1994) pp.389-392 ISSN: 0034-5288
Smart D., Singh I., Smith R., Forhead A., Dobson H., The hypothalamic-pituitary-adrenal axis in post-partum ewes
Animal Reproduction Science 35 (1994) pp.223-229 ISSN: 0378-4320 eISSN: 1873-2232
Smart D., Forhead A., Smith R., Dobson H., Transport stress delays the oestradiol-induced LH surge by a non-opioidergic mechanism in the early postpartum ewe
Journal of Endocrinology 142 (3) (1994) pp.447-451 ISSN: 0022-0795

Fowden A., Ward J., Forhead A., Control of fetal metabolism: Relevance to developmental origins of health and disease ,
Cambridge University Press (2006) ISBN: 9780521847438

Forhead AJ, Fowden AL, Formation and growth of the fetus
in Michael Skinner (ed.), Encyclopedia of Reproduction, Elsevier (2018) ISBN: 9780128118993
Fowden A, Forhead A, Endocrine interactions in the control of fetal growth
in Bhatia J, Bhutta ZA, Kalhan SC (ed.), Endocrine interactions in the control of fetal growth, S. Karger AG (2013) ISBN: 978-3-318-02387-9 eISBN: 978-3-318-02388-6
, The consequences for preterm infants of antenatal glucocorticoid treatment.
in Birth Rites and Rights, Hart Publishing (2011)
, Glucocorticoids and placental programming
in The Placenta and Human Developmental Programming, Cambridge University Press (2010)
Vaughan O, Forhead A, Fowden A, Glucocorticoids and placental programming
in Burton GJ, Barker DJP, Moffett A, Thornburg K (ed.), Glucocorticoids and placental programming, Cambridge University Press (2010) ISBN: 9780521199452 eISBN: 9780511933806
, The role of hormones in intrauterine development
in Fetal Origins of Cardiovascular and Lung Disease, NIH Monographs (2001)

Fowden AL, Vaughan OR, Murray AJ & Forhead AJ , Metabolic consequences of glucocorticoid exposure before birth ,
(2022) Metabolic consequences of glucocorticoid exposure before birth Open Access version on RADAR
Fowden AL, Forhead AJ, Endocrine regulation of fetal metabolism towards term: Hormone and fetal metabolism ,
(2021) Endocrine regulation of fetal metabolism towards term: Hormone and fetal metabolism Open Access version on RADAR
Fowden AL, Giussani DA, Forhead AJ, Physiological development of the equine fetus during late gestation ,
(2019) Physiological development of the equine fetus during late gestation Open Access version on RADAR
Forhead AJ, Breathing for two: maternal asthma and lung development in the fetus ,
(2019) Breathing for two: maternal asthma and lung development in the fetus Open Access version on RADAR
Forhead AJ, Legacy of excess: consequences of maternal obesity for the adult offspring ,
(2018) Legacy of excess: consequences of maternal obesity for the adult offspring Open Access version on RADAR