Carol A. Bagnell

Carol A. Bagnell.

Foran Hall 126
Phone: 848-932-6334


Studies in my laboratory focus on the physiology of the hormone relaxin in the control of female reproductive tract development and remodeling. Relaxin, a member of the insulin-like family of hormones, has long been regarded as a hormone of pregnancy known for its trophic and connective tissue remodeling actions in the female reproductive tract that facilitate delivery at term and prepare the mammary gland for lactation. However, studies show that relaxin is not only a hormone of pregnancy but has multiple roles in non-pregnant animals and targets both reproductive and non-reproductive tissues. In fact, recent studies show that a developmentally-regulated, estrogen-sensitive relaxin receptor system is present in the porcine uterus and cervix from birth. In addition, porcine milk contains relaxin that is transmitted into the peripheral circulation of neonates only if they are allowed to nurse. These studies indicate that a window of opportunity for transmission of milk-borne relaxin from mother to neonate is open during the first few days of life. Thus, factors, such as relaxin, with the potential to control reproductive tract development during early postnatal life are communicated to the neonate in milk by way of a maternally driven lactocrine mechanism. Currently, specific areas of research include:

  • determining how milk-borne relaxin influences the developmental program and trajectory of uterine and cervical tissues.
  • studying the interaction between relaxin and estrogen signaling pathways in regulating reproductive tissue growth.
  • identifying molecular markers and mediators that control reproductive development (e.g. Hoxa, Wnt, estrogen and relaxin receptor family genes) and how they respond to hormonal and environmental perturbations.
  • studying the effects of relaxin on non-reproductive targets including cardiac and immune systems.


University of Hawaii
Title Location Year
Chair Dept. of Animal Sciences, Rutgers University 2008 - 2014
Professor Dept. of Animal Sciences, Rutgers University 2002
Visiting Scientist Merck & Co., Inc.
Dept. of Atherosclerosis & Endocrinology
2001 - 2002
Associate Professor Dept. of Animal Sciences, Rutgers University 1994
Associate Chair Dept. of Animal Sciences, Rutgers University 1995
Assistant Professor Dept. of Animal Sciences, Rutgers University 1988
Associate Researcher Pacific Biomedical Research Center (PBRC)
Dept of Anatomy and Reproductive Biology
Assistant Researcher Pacific Biomedical Research Center (PBRC)
Dept of Anatomy and Reproductive Biology


Degree Location Year
Postdoc University of Hawaii 1984
Ph.D. Medical College of Georgia 1983
M.S. West Virginia University 1976
B.S. Glassboro State College, NJ 1974

Professional Societies

American Association for the Advancement of Science
Endocrine Society
Sigma Xi
Society for the Study of Reproduction
Women in Endocrinology

Editorial Positions

Editorial Board, Reproductive Biology and Endocrinology, 2002 - present
Editorial Board, Domestic Animal Endocrinology, 2002 - 2005
Editorial Board, Biology of Reproduction, 1999 - 2004

Current Research Associates and Graduate Students

Ashley George Ph.D. student Maternal Programming of the Neonatal Uterus
Teh-Yuan Ho Research Associate

Former Research Associates and Graduate Students

Students Degree Title Location
Meredith Camp PhD, 2015 Clinical Research Associate Kindred Biological Sciences, Inc.
Burlingame, CA
Henry Bignell MS, 2015 County Extension Agent University of Florida
Gainesville, FL
Kathleen Rahman PhD, 2014 Instructor and Undergraduate Program Director Department of Animal Sciences
Rutgers University
Amy-Lynn Frankshun PhD, 2011 Staff Fellow Food & Drug Administration
Rockville, MD
Joseph Chen PhD, 2010 Senior Scientist Siemens Healthcare Diagnostics
Emily Lamprecht PhD, 2009 Development Manager Cargill Animal Nutrition
Elk River, MN
Wenbo Yan Ph.D, 2005 Assistant Professor Nyack University
New York, NY
Karen Santora M.S., 2005 Regulatory Affairs Associate Sandoz Inc
Denver, CO
Jennifer Neumann M.S., 2002 Physician Charlotte, NC
Judy Lenhart Ph.D., 1999 Director, Global Communications Celgene Corporation
Berkeley Heights, NJ
Kathleen Ohleth Ph.D.,1997 President and CEO Precise Publications
Bridgewater, NJ
Qi Zhang Ph.D., 1994 Pathologist Columbia University
New York, NY

Research Associates

Frank Petersen Ph.D. Research Scientist NJ Dept of Agriculture
Trenton, NJ
Peter Ryan Ph.D. Professor Missisippi State University


11:067:327 Animal Reproduction
11:067:430 Animal Microtechniques and Tissue Culture
16:067:502 Physiology of Reproduction
16:067:701 Graduate Research in Animal Science

Representative Publications

Rahman KM, Camp ME, Prasad N, McNeel AK, Levy SE, Bartol FF, Bagnell CA. 2015. Age and nursing affect the neonatal porcine uterine transcriptome. Biol Reprod, in press. PubMed

Harvey C, Chen J, Bagnell CA, Uzumcu M. 2015. Methoxychlor and its metabolite HPTE inhibit cAMP production and expression of estrogen receptors alpha and beta in rat granulosa cell in vitro. Reproductive Toxicology 51:72-78. PubMed

Camp ME, Wiley AA, Boulos MB, Rahman KM, Bartol FF, Bagnell CA. 2014. Effects of age, nursing, and oral IGF1 supplementation on neonatal porcine cervical development. Reproduction 148: 441-451. PubMed

Rahman KM, Lovich JE, Lam C, Camp ME, Wiley AA, Bartol FF, Bagnell CA. 2014. Nursing supports neonatal porcine testicular development. Domestic Animal Endocrinology 48: 84-92. PubMed

Bartol FF, Wiley AA, Miller DJ, Silva AJ, Roberts KE, Davolt MLP, Chen JC, Frankshun A-L, Camp ME, Rahman KM, Vallet JL, and Bagnell CA. 2013. Lactation Biology Symposium: Lactocrine signaling and developmental programming. Journal of Animal Science 91:696-705. PubMed

Miller DJ, Wiley AA, Chen JC, Bagnell CA, Bartol FF. 2013. Nursing for 48 h from birth supports uterine gland development and endometrial cell- compartment specific gene expression. Biol Reprod 88 (1):4, 1-10. PubMed

Frankshun A-L, Chen JC, Barron L, Ho T-Y, Miller DJ, Rahman K, Bartol FF, Bagnell CA. 2012. Nursing during the first two days of life is essential for the expression of proteins important for growth and remodeling of the neonatal porcine cervix. Endocrinology 153(9):4511-21. PubMed

Bartol FF and Bagnell CA. 2012. Lactocrine programming of female reproductive tract development: Environmental connections to the reproductive continuum. Mol Cell Endocrinol 354:16-21.PubMed

Chen JC, Frankshun A-L, Wiley AA, Miller DJ, Welch KA, Ho T-Y, Bartol FF, Bagnell CA. 2011. Relaxin and milk-borne lactocrine-acting factors affect morphoregulatory gene and protein expression patterns in the neonatal porcine uterus. Reproduction 141:675-683.PubMed

Ho T-Y, Santora K, Frankshun A-L, Bagnell CA. 2011. Effects of relaxin and estrogens on bone remodeling markers, receptor activator of NF-kB ligand (RANKL) and osteoprotegerin (OPG), in rat adjuvant-induced arthritis. Bone 48:1346-1353.PubMed

Frankshun A-L, Ho T-Y, Reimer DC, Chen J, Lasano S, Steinetz BG, Bartol FF, Bagnell CA. 2011. Characterization and biological activity of relaxin in porcine milk. Reproduction 141: 373-380.PubMed

Chen JC, Wiley AA, Ho T-Y, Frankshun A-L, Hord KM, Bartol FF, Bagnell CA. 2010.Transient estrogen exposure from birth affects uterine expression of developmental markers in neonatal gilts with lasting consequences in pregnant adults. Reproduction 139: 623-630.PubMed

Bagnell CA, Steinetz BG, Bartol FF. 2009. Milk-borne relaxin and the lactocrine hypothesis for maternal programming of neonatal tissues. Ann N Y Acad Sci 1160:152-157. PubMed

Bartol FF, Wiley AA, Bagnell CA. 2009. Relaxin and maternal lactocrine programming of neonatal uterine development. Ann N Y Acad Sci 1160:158-163.PubMed

Chen J, Wiley AA, Kauffold H, Wahner M, Bartol FF, Bagnell CA. 2009. Perinatal zearalenone exposure affects RXFP1, RXFP2 and morphoregulatory gene expression in the neonatal porcine uterus. Ann N Y Acad Sci 1160: 188-189. PubMed

Ho TY, Dilts MA, Bartol FF, Bagnell CA. 2009. Relaxin promotes matrix metalloproteinase-2 and decreases Wnt/b-catenin expression in the neonatal porcine heart. Ann N Y Acad Sci 1160:287-288. PubMed

Bartol FF, Wiley AA, Bagnell CA. 2008. Epigenetic programming of porcine endometrial function and the lactocrine hypothesis. Reprod Dom Anim 43(2):281-287. PubMed

Yan W, Chen J, Wiley AA, Bartol FF, Bagnell CA. 2008. Relaxin and estrogen affect estrogen receptor ?, vascular endothelial growth factor and relaxin receptor expression in the neonatal porcine uterus and cervix. Reproduction 135:705-712. PubMed

Santora K, Rasa C, Visco D, Steinetz BG, Bagnell CA. 2007. Antiarthritic effects of relaxin, in combination with estrogen, in rat adjuvant-induced arthritis. J Pharmacol & Experimental Therapeutics Aug 322(2): 887-893. PubMed

Ho TY, Yan W, Bagnell CA. 2007. Relaxin-induced matrix metalloproteinase-9 expression is associated with the activation of the NF-kB pathway in human THP-1 cells. J Leukocyte Biology 81:1303-1310. PubMed

Masters RA, Crean BD, Yan W, Moss A, Ryan PL, Wiley AA, Bagnell CA, Bartol FF. 2007. Neonatal porcine endometrial development and epithelial proliferation affected by age and exposure to estrogen and relaxin. Domestic Anim Endocrinol 33: 335-346. PubMed

Lamprecht E D, Bagnell CA, Williams CA. 2007. Inflammatory responses to three modes of intense exercise in Standardbred mares. Equine Sci. Soc. Proc. 20: 76-77.

Yan W, Wiley AA, Bathgate RAD, Frankshun AL, Lasano S, Crean BD, Steinetz BG, Bagnell CA, Bartol FF. 2006. Expression of LGR7 and LGR8 by neonatal porcine uterine tissues and transmission of milk-borne relaxin into the neonatal circulation by suckling. Endocrinology 147:4303-10. PubMed

Yan W, Ryan PL, Bartol FF, Bagnell CA. 2006. Uterotrophic effects of relaxin related to age and estrogen receptor activation in neonatal pigs. Reproduction 131:943-950.PubMed

Neumann J, Lazaris A, Huang YJ, Karatzas C, Ryan R, Bagnell CA. 2006. Production and characterization of recombinant equine prorelaxin. Domestic Anim Endocrinol 31:173-185. PubMed

Bartol FF, Wiley AA, BagnellCA. 2006. Uterine development and endometrial programming. Reproduction Suppl 62:113-130. PubMed

Yan W, Wiley, AA, Bartol FF and Bagnell CA. 2005. Tissue-specific effects of relaxin on the reproductive tract of neonatal gilts. Ann N Y Acad Sci. 1041:132-5.PubMed

Bagnell CA, Yan W, Wiley AA, Bartol FF. 2005. Effects of relaxin on neonatal porcine uterine growth and development. Ann N Y Acad Sci. 1041:248-55.PubMed

Ho TY andBagnell CA. 2005. Relaxin induces matrix metalloproteinase-9 through activation of nuclear factor kappa B in human THP-1 cells. Ann N Y Acad Sci. 1041:314-6.PubMed

Santora K, Rasa C, Ronan J, Visco D, Steinetz B, Bagnell CA. 2005. Effects of relaxin in a model of rat adjuvant-induced arthritis. Ann N Y Acad Sci. 1041:481-5.PubMed

Lenhart JA, Ryan PL, Ohleth KA, Palmer SS, Bagnell CA. 2002. Relaxin increases secretion of tissue inhibitor of matrix metalloproteinase-1 and -2 during uterine and cervical growth and remodeling in the pig. Endocrinology 143:91-98.PubMed

Ryan PL, Baum DL, Lenhart JA, Ohleth KM, Bagnell CA. 2001. Expression of uterine and cervical epithelial cadherin expression during relaxin-induced growth in pigs. Reproduction 122:929-937.PubMed

Lenhart JA, Ryan PL, Ohleth KM, Palmer SS, Bagnell CA. 2001. Relaxin increases secretion of matrix metalloproteinase-2 and -9 during uterine and cervical growth and remodeling in the pig. Endocrinology 142:3941-3949.PubMed

Ryan PL, Bennett-Wimbush K Vaala WE, Bagnell CA. 2001. Systemic relaxin in pregnant pony mares grazed on endophyte infected fescue: Effects of fluphenazine treatment. Theriogenology 56:471-483.PubMed

Kirkup K, Mallin A, Bagnell CA. 2000. Inhibition of pig granulosa cell adhesion and growth in vitro by immunoneutralization of epithelial cadherin. J Reprod & Fertility 120:275-281.PubMed

Ryan PL, Bennet-Wimbush K, Vaala W, Bagnell CA. 1999. Relaxin as a biochemical marker of placental insufficiency in the horse: a review. Pferdeheilkunde (Equine Health)15:622-626.
Lenhart JA, Ryan PL, Ohleth KM, Bagnell CA. 1999. Expression of connexin-26,-32 and -43 gap junction proteins in the porcine cervix and uterus during pregnancy and relaxin-induced growth. Biol Reprod 61:1452-1459.PubMed

Lenhart JA, Ohleth KM, Ryan PL, Palmer SS, Bagnell CA.1999. Effect of relaxin on tissue inhibitor of metalloproteinase-1 and -2 in the porcine uterus and cervix. In: RA Greenwald, S Zuker, LM Golub (eds). Inhibition of Matrix Metalloproteinases: Therapeutic Applications, Annals of the NY Acad of Sciences Vol 878: 565-566.PubMed

Ohleth KM, Zhang Q, Lenhart JA, Ryan PL, Bagnell CA. 1999. Trophic effects of relaxin on reproductive tissue: Role of the IGF system. Steroids 64: 634-639.PubMed

Ohleth KM and Bagnell CA.1999. Relaxin secretion and gene expression in porcine granulosa and theca cells are stimulated during in vitro luteinization. Biol Reprod 60: 499-507.PubMed

Lenhart JA, Downey BR, Bagnell CA. 1998. Connexin 43 gap junction protein expression during follicular development in the porcine ovary. Biol Reprod 58:583-590.PubMed

Wang-Lee JL, Lenhart JA, Ohleth KM, Ryan PL, Bagnell CA. 1998. Regulation of urokinase and tissue-type plasminogen activator by relaxin in the uterus and cervix of the prepubertal gilt. J Reprod Fertility 114: 119-125.PubMed

Ohleth KM, Zhang Q, Bagnell CA. 1998. Relaxin protein and gene expression in ovarian follicles of immature pigs. J Molecular Endocrinology 21:179-187.PubMed

Ryan P, Vaala W, Bagnell CA.1998. Evidence the equine relaxin in a good indicator of placental insufficiency in the mare. Proceedings American Assoc Equine Practitioners 44: 62-63.

Ohleth KM, Lenhart JA, Ryan PL, Radecki SV, Bagnell CA. 1997. Relaxin increases insulin-like growth factors and IGF binding proteins of the pig uterus in vivo. Endocrinology 138:3652-3258.PubMed

Ryan PL, Valentine AF and Bagnell CA1996. Expression of E-cadherin in the developing and adult pig ovary. Biol Reprod 55:1091-1097. PubMed