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Pharmacology & Systems Physiology 

Photo of Nelson Horseman, PhD

Nelson Horseman, PhD

Professor Emeritus, Department of Molecular and Cellular Physiology, UC College of Medicine

Contact Information

Peer Reviewed Publications

Horseman, Nelson D; Gregerson, Karen A 2014. Prolactin actions. Journal of molecular endocrinology, 52 1, R95-106

Horseman, Nelson D; Hernandez, Laura L 2014. New concepts of breast cell communication to bone. Trends in endocrinology and metabolism: TEM, 25 1, 34-41

Christensen, Heather R; Murawsky, Michael K; Horseman, Nelson D; Willson, Tara A; Gregerson, Karen A 2013. Completely humanizing prolactin rescues infertility in prolactin knockout mice and leads to human prolactin expression in extrapituitary mouse tissues. Endocrinology, 154 12, 4777-89

Lemay, Danielle G; Ballard, Olivia A; Hughes, Maria A; Morrow, Ardythe L; Horseman, Nelson D; Nommsen-Rivers, Laurie A 2013. RNA sequencing of the human milk fat layer transcriptome reveals distinct gene expression profiles at three stages of lactation. PloS one, 8 7, e67531

Marshall, Aaron M; Hernandez, Laura L; Horseman, Nelson D 2013. Serotonin and Serotonin Transport in the Regulation of Lactation. Journal of mammary gland biology and neoplasia, ,

Chen, Chien-Chung; Stairs, Douglas B; Boxer, Robert B; Belka, George K; Horseman, Nelson D; Alvarez, James V; Chodosh, Lewis A 2012. Autocrine prolactin induced by the Pten-Akt pathway is required for lactation initiation and provides a direct link between the Akt and Stat5 pathways. Genes & development, 26 19, 2154-68

Collier, R J; Hernandez, L L; Horseman, N D 2012. Serotonin as a homeostatic regulator of lactation. Domestic animal endocrinology, 43 2, 161-70

Hernandez, Laura L; Grayson, Bernadette E; Yadav, Ekta; Seeley, Randy J; Horseman, Nelson D 2012. High fat diet alters lactation outcomes: possible involvement of inflammatory and serotonergic pathways. PloS one, 7 3, e32598

Hernandez, Laura L; Gregerson, Karen A; Horseman, Nelson D 2012. Mammary gland serotonin regulates parathyroid hormone-related protein and other bone-related signals. American journal of physiology. Endocrinology and metabolism, 302 8, E1009-15

Neville, Margaret C; Anderson, Steven M; McManaman, James L; Badger, Thomas M; Bunik, Maya; Contractor, Nikhat; Crume, Tessa; Dabelea, Dana; Donovan, Sharon M; Forman, Nicole; Frank, Daniel N; Friedman, Jacob E; German, J Bruce; Goldman, Armond; Hadsell, Darryl; Hambidge, Michael; Hinde, Katie; Horseman, Nelson D; Hovey, Russell C; Janoff, Edward; Krebs, Nancy F; Lebrilla, Carlito B; Lemay, Danielle G; MacLean, Paul S; Meier, Paula; Morrow, Ardythe L; Neu, Josef; Nommsen-Rivers, Laurie A; Raiten, Daniel J; Rijnkels, Monique; Seewaldt, Victoria; Shur, Barry D; VanHouten, Joshua; Williamson, Peter 2012. Lactation and neonatal nutrition: defining and refining the critical questions. Journal of mammary gland biology and neoplasia, 17 2, 167-88

Hernandez, L L; Collier, J L; Vomachka, A J; Collier, R J; Horseman, N D 2011. Suppression of lactation and acceleration of involution in the bovine mammary gland by a selective serotonin reuptake inhibitor. The Journal of endocrinology, 209 1, 45-54

Pai, Vaibhav P; Horseman, Nelson D 2011. Multiple cellular responses to serotonin contribute to epithelial homeostasis. PloS one, 6 2, e17028

Marshall, Aaron M; Nommsen-Rivers, Laurie A; Hernandez, Laura L; Dewey, Kathryn G; Chantry, Caroline J; Gregerson, Karen A; Horseman, Nelson D 2010. Serotonin transport and metabolism in the mammary gland modulates secretory activation and involution. The Journal of clinical endocrinology and metabolism, 95 2, 837-46

Hernandez, Laura L; Limesand, Sean W; Collier, Jayne L; Horseman, Nelson D; Collier, Robert J 2009. The bovine mammary gland expresses multiple functional isoforms of serotonin receptors. The Journal of endocrinology, 203 1, 123-31

Marshall, Aaron M; Pai, Vaibhav P; Sartor, Maureen A; Horseman, Nelson D 2009. In vitro multipotent differentiation and barrier function of a human mammary epithelium. Cell and tissue research, 335 2, 383-95

Pai, Vaibhav P; Marshall, Aaron M; Hernandez, Laura L; Buckley, Arthur R; Horseman, Nelson D 2009. Altered serotonin physiology in human breast cancers favors paradoxical growth and cell survival. Breast cancer research : BCR, 11 6, R81

Pai, Vaibhav P; Horseman, Nelson D 2008. Biphasic regulation of mammary epithelial resistance by serotonin through activation of multiple pathways. The Journal of biological chemistry, 283 45, 30901-10

Bao, Lei; Tessier, Christian; Prigent-Tessier, Anne; Li, Feixue; Buzzio, Oscar L; Callegari, Eduardo A; Horseman, Nelson D; Gibori, Geula 2007. Decidual prolactin silences the expression of genes detrimental to pregnancy. Endocrinology, 148 5, 2326-34

Dugan, Amy L; Schwemberger, Sandy; Noel, Greg J; Babcock, George F; Ogle, Cora K; Buckley, Donna J; Horseman, Nelson D; Gregerson, Karen A 2007. Stress and prolactin effects on bone marrow myeloid cells, serum chemokine and serum glucocorticoid levels in mice. Neuroimmunomodulation, 14 6, 287-96

Dugan, Amy L; Schwemberger, Sandy; Noel, Greg J; Babcock, George; Ogle, Cora K; Horseman, Nelson D 2007. Psychogenic stress prior to burn injury has differential effects on bone marrow and cytokine responses. Experimental biology and medicine (Maywood, N.J.), 232 2, 253-61

John, S K P; Jones, O M; Horseman, N; Thomas, P; Howell, R D; Fozard, J B J 2007. Inter general practice variability in use of referral guidelines for colorectal cancer. Colorectal disease : the official journal of the Association of Coloproctology of Great Britain and Ireland, 9 8, 731-5

Jones, O M; John, S K P; Horseman, N; Lawrance, R J; Fozard, J B J 2007. Low anastomotic leak rate after colorectal surgery: a single-centre study. Colorectal disease : the official journal of the Association of Coloproctology of Great Britain and Ireland, 9 8, 740-4

Jones, O M; John, S K P; Horseman, N; Lawrance, R J; Fozard, J B J 2007. Cause and place of death in patients dying with colorectal cancer. Colorectal disease : the official journal of the Association of Coloproctology of Great Britain and Ireland, 9 3, 253-7

Soares, Michael J; Alam, S M Khorshed; Duckworth, Mary Lynn; Horseman, Nelson D; Konno, Toshihiro; Linzer, Daniel I H; Maltais, Lois J; Nilsen-Hamilton, Marit; Shiota, Kunio; Smith, Jennifer R; Wallis, Michael 2007. A standardized nomenclature for the mouse and rat prolactin superfamilies. Mammalian genome : official journal of the International Mammalian Genome Society, 18 3, 154-6

Stull, Malinda A; Pai, Vaibhav; Vomachka, Archie J; Marshall, Aaron M; Jacob, George A; Horseman, Nelson D 2007. Mammary gland homeostasis employs serotonergic regulation of epithelial tight junctions. Proceedings of the National Academy of Sciences of the United States of America, 104 42, 16708-13

LaPensee, Christopher R; Horseman, Nelson D; Tso, Patrick; Brandebourg, Terry D; Hugo, Eric R; Ben-Jonathan, Nira 2006. The prolactin-deficient mouse has an unaltered metabolic phenotype. Endocrinology, 147 10, 4638-45

Obál, Ferenc; Garcia-Garcia, Fabio; Kacsóh, Balint; Taishi, Ping; Bohnet, Stewart; Horseman, Nelson D; Krueger, James M 2005. Rapid eye movement sleep is reduced in prolactin-deficient mice. The Journal of neuroscience : the official journal of the Society for Neuroscience, 25 44, 10282-9

Bailey, Jason P; Nieport, Kathryn M; Herbst, Matthew P; Srivastava, Sunil; Serra, Rosa A; Horseman, Nelson D 2004. Prolactin and transforming growth factor-beta signaling exert opposing effects on mammary gland morphogenesis, involution, and the Akt-forkhead pathway. Molecular endocrinology (Baltimore, Md.), 18 5, 1171-84

Dugan, Amy L; Schwemberger, Sandy; Babcock, George F; Buckley, Donna; Buckley, Arthur R; Ogle, Cora K; Horseman, Nelson D 2004. Effects of prolactin level on burn-induced aberrations in myelopoiesis. Shock (Augusta, Ga.), 21 2, 151-9

Matsuda, Manabu; Imaoka, Tatsuhiko; Vomachka, Archie J; Gudelsky, Gary A; Hou, Zhaoyuan; Mistry, Meenakshi; Bailey, Jason P; Nieport, Kathryn M; Walther, Diego J; Bader, Michael; Horseman, Nelson D 2004. Serotonin regulates mammary gland development via an autocrine-paracrine loop. Developmental cell, 6 2, 193-203

Dorshkind, Kenneth; Welniak, Lisbeth; Gault, Ruth A; Hixon, Julie; Montecino-Rodriguez, Encarnacion; Horseman, Nelson D; Gertner, Joseph M; Murphy, William J 2003. Effects of housing on the thymic deficiency in dwarf mice and its reversal by growth hormone administration. Clinical immunology (Orlando, Fla.), 109 2, 197-202

Hou, Zhaoyuan; Srivastava, Sunil; Mistry, Meenakshi J; Herbst, Matthew P; Bailey, Jason P; Horseman, Nelson D 2003. Two tandemly linked interferon-gamma-activated sequence elements in the promoter of glycosylation-dependent cell adhesion molecule 1 gene synergistically respond to prolactin in mouse mammary epithelial cells. Molecular endocrinology (Baltimore, Md.), 17 10, 1910-20

Krishnan, Nithya; Thellin, Olivier; Buckley, Donna J; Horseman, Nelson D; Buckley, Arthur R 2003. Prolactin suppresses glucocorticoid-induced thymocyte apoptosis in vivo. Endocrinology, 144 5, 2102-10

Ormandy, Christopher J; Naylor, Matthew; Harris, Jessica; Robertson, Fiona; Horseman, Nelson D; Lindeman, Geoffrey J; Visvader, Jane; Kelly, Paul A 2003. Investigation of the transcriptional changes underlying functional defects in the mammary glands of prolactin receptor knockout mice. Recent progress in hormone research, 58 , 297-323

Srivastava, Sunil; Matsuda, Manabu; Hou, Zhaoyuan; Bailey, Jason P; Kitazawa, Riko; Herbst, Matthew P; Horseman, Nelson D 2003. Receptor activator of NF-kappaB ligand induction via Jak2 and Stat5a in mammary epithelial cells. The Journal of biological chemistry, 278 46, 46171-8

Tessier, Christian; Prigent-Tessier, Anne; Bao, Lei; Telleria, Carlos M; Ferguson-Gottschall, Susan; Gibori, Gil B; Gu, Yan; Bowen-Shauver, Jennifer M; Horseman, Nelson D; Gibori, Geula 2003. Decidual activin: its role in the apoptotic process and its regulation by prolactin. Biology of reproduction, 68 5, 1687-94

Cruz-Soto, Martha E; Scheiber, Michael D; Gregerson, Karen A; Boivin, Gregory P; Horseman, Nelson D 2002. Pituitary tumorigenesis in prolactin gene-disrupted mice. Endocrinology, 143 11, 4429-36

Dugan, Amy L; Thellin, Olivier; Buckley, Donna J; Buckley, Arthur R; Ogle, Cora K; Horseman, Nelson D 2002. Effects of prolactin deficiency on myelopoiesis and splenic T lymphocyte proliferation in thermally injured mice. Endocrinology, 143 10, 4147-51

Horseman, Nelson D 2002. Prolactin receptor diversity in humans: novel isoforms suggest general principles. Trends in endocrinology and metabolism: TEM, 13 2, 47-8

Imaoka, Tatsuhiko; Horseman, Nelson D; Lockefeer, Jason A; Mori, Takao; Matsuda, Manabu 2002. Cortactin-binding protein 90 (CBP90) expression in the mouse mammary glands during prolactin-induced lobuloalveolar development. Zoological science, 19 4, 443-8

Dorshkind, K; Horseman, N D 2001. Anterior pituitary hormones, stress, and immune system homeostasis. BioEssays : news and reviews in molecular, cellular and developmental biology, 23 3, 288-94

Thellin, O; Noel, G; Khurana, S; Ogle, C K; Horseman, N D 2001. Stress hormone secretion and gut signal transducer (STAT) proteins after burn injury in rats. Shock (Augusta, Ga.), 16 5, 393-7

Dorshkind, K; Horseman, N D 2000. The roles of prolactin, growth hormone, insulin-like growth factor-I, and thyroid hormones in lymphocyte development and function: insights from genetic models of hormone and hormone receptor deficiency. Endocrine reviews, 21 3, 292-312

Foster, M P; Jensen, E R; Montecino-Rodriguez, E; Leathers, H; Horseman, N; Dorshkind, K 2000. Humoral and cell-mediated immunity in mice with genetic deficiencies of prolactin, growth hormone, insulin-like growth factor-I, and thyroid hormone. Clinical immunology (Orlando, Fla.), 96 2, 140-9

Hou, Z; Bailey, J P; Vomachka, A J; Matsuda, M; Lockefeer, J A; Horseman, N D 2000. Glycosylation-dependent cell adhesion molecule 1 (GlyCAM 1) is induced by prolactin and suppressed by progesterone in mammary epithelium. Endocrinology, 141 11, 4278-83

Ogle, C K; Kong, F; Guo, X; Wells, D A; Aosasa, S; Noel, G; Horseman, N 2000. The effect of burn injury on suppressors of cytokine signalling. Shock (Augusta, Ga.), 14 3, 392-8; discussion 39

Phelps, C J; Horseman, N D 2000. Prolactin gene disruption does not compromise differentiation of tuberoinfundibular dopaminergic neurons. Neuroendocrinology, 72 1, 2-10

Pratt, S L; Ogle, C K; Mao, J X; Zhao, W; Lovell, G; Horseman, N D 2000. Interleukin-6 signal transduction in human intestinal epithelial cells. Shock (Augusta, Ga.), 13 6, 435-40

Vomachka, A J; Pratt, S L; Lockefeer, J A; Horseman, N D 2000. Prolactin gene-disruption arrests mammary gland development and retards T-antigen-induced tumor growth. Oncogene, 19 8, 1077-84

Gao, J; Horseman, N D 1999. Prolactin-independent modulation of the beta-casein response element by Erk2 MAP kinase. Cellular signalling, 11 3, 205-10

Horseman, N D 1999. Citation for the 1999 Gerald D. Aurbach Lecture Award of the Endocrine Society to Dr. James E. Darnell. Endocrine reviews, 20 4, 591-2

Horseman, N D 1999. Prolactin and mammary gland development. Journal of mammary gland biology and neoplasia, 4 1, 79-88

Pratt, S L; Horseman, N D 1998. Identification of two Y-box binding proteins that interact with the promoters of columbid annexin I genes. Gene, 214 1-2, 147-56

Steger, R W; Chandrashekar, V; Zhao, W; Bartke, A; Horseman, N D 1998. Neuroendocrine and reproductive functions in male mice with targeted disruption of the prolactin gene. Endocrinology, 139 9, 3691-5

Horseman, N D; Zhao, W; Montecino-Rodriguez, E; Tanaka, M; Nakashima, K; Engle, S J; Smith, F; Markoff, E; Dorshkind, K 1997. Defective mammopoiesis, but normal hematopoiesis, in mice with a targeted disruption of the prolactin gene. The EMBO journal, 16 23, 6926-35

Schuler, L A; Nagel, R J; Gao, J; Horseman, N D; Kessler, M A 1997. Prolactin receptor heterogeneity in bovine fetal and maternal tissues. Endocrinology, 138 8, 3187-94

Gao, J; Hughes, J P; Auperin, B; Buteau, H; Edery, M; Zhuang, H; Wojchowski, D M; Horseman, N D 1996. Interactions among Janus kinases and the prolactin (PRL) receptor in the regulation of a PRL response element. Molecular endocrinology (Baltimore, Md.), 10 7, 847-56

Gunteski-Hamblin, A M; Song, G; Walsh, R A; Frenzke, M; Boivin, G P; Dorn, G W; Kaetzel, M A; Horseman, N D; Dedman, J R 1996. Annexin VI overexpression targeted to heart alters cardiomyocyte function in transgenic mice. The American journal of physiology, 270 3 Pt 2, H1091-100

Horseman, N D 1995. Prolactin, proliferation, and protooncogenes. Endocrinology, 136 12, 5249-51

Horseman, N D; Buntin, J D 1995. Regulation of pigeon cropmilk secretion and parental behaviors by prolactin. Annual review of nutrition, 15 , 213-38

Lai, C F; Ripperger, J; Morella, K K; Wang, Y; Gearing, D P; Horseman, N D; Campos, S P; Fey, G H; Baumann, H 1995. STAT3 and STAT5B are targets of two different signal pathways activated by hematopoietin receptors and control transcription via separate cytokine response elements. The Journal of biological chemistry, 270 40, 23254-7

Chen, X; Horseman, N D 1994. Cloning, expression, and mutational analysis of the pigeon prolactin receptor. Endocrinology, 135 1, 269-76

Gao, Y; Horseman, N D 1994. Structural and functional divergences of the columbid annexin I-encoding cp37 and cp35 genes. Gene, 143 2, 179-86

Horseman, N D 1994. Famine to feast--growth hormone and prolactin signal transducers. Endocrinology, 135 4, 1289-91

Horseman, N D; Yu-Lee, L Y 1994. Transcriptional regulation by the helix bundle peptide hormones: growth hormone, prolactin, and hematopoietic cytokines. Endocrine reviews, 15 5, 627-49

Kaetzel, M A; Pula, G; Campos, B; Uhrin, P; Horseman, N; Dedman, J R 1994. Annexin VI isoforms are differentially expressed in mammalian tissues. Biochimica et biophysica acta, 1223 3, 368-74

Sidis, Y; Horseman, N D 1994. Prolactin induces rapid p95/p70 tyrosine phosphorylation, and protein binding to GAS-like sites in the anx Icp35 and c-fos genes. Endocrinology, 134 4, 1979-85

Sidis, Y; Horseman, N D 1993. The hinge region of chicken annexin I contains no site for tyrosine phosphorylation. FEBS letters, 329 3, 296-300

Uhrin, P; Horseman, N D 1993. Regulation of the annexin Icp35 gene in transfected mammary gland cell lines. Biochemistry and molecular biology international, 30 2, 305-10

Haigler, H T; Mangili, J A; Gao, Y; Jones, J; Horseman, N D 1992. Identification and characterization of columbid annexin Icp37. Insights into the evolution of annexin I phosphorylation sites. The Journal of biological chemistry, 267 27, 19123-9

Horseman, N D; Chen, X; Liu, L; Poyet, P; Hitti, Y 1992. Cell and species distribution of prolactin-inducible annexin I mRNA. General and comparative endocrinology, 85 3, 405-14

Xu, Y H; Horseman, N D 1992. Nuclear proteins and prolactin-induced annexin Icp35 gene transcription. Molecular endocrinology (Baltimore, Md.), 6 3, 375-83

Hitti, Y S; Horseman, N D 1991. Structure of the gene encoding columbid annexin Icp35. Gene, 103 2, 185-92

Horseman, N D 1990. Hormonal regulation of an avian annexin I gene. Biochemical Society transactions, 18 6, 1113-5

Young, K H; Buhi, W C; Horseman, N; Davis, J; Kraeling, R; Linzer, D; Bazer, F W 1990. Biological activities of glycosylated and nonglycosylated porcine prolactin. Molecular and cellular endocrinology, 71 2, 155-62

Horseman, N D 1989. A prolactin-inducible gene product which is a member of the calpactin/lipocortin family. Molecular endocrinology (Baltimore, Md.), 3 5, 773-9

Horseman, N D; Hankovich, S 1989. A chamber and method which simplify fixing ultrathin sequencing gels. BioTechniques, 7 4, 340, 342, 344

Pauly, J R; Horseman, N D 1988. Autoradiographic localization of alpha-bungarotoxin binding sites in the suprachiasmatic region of rat brain. Brain research, 452 1-2, 105-12

Pauly, J R; Horseman, N D 1988. Perturbations of locomotor activity rhythms following suprachiasmatic bungarotoxin infusion. Physiology & behavior, 43 6, 859-65

Pukac, L A; Horseman, N D 1987. Regulation of cloned prolactin-inducible genes in pigeon crop. Molecular endocrinology (Baltimore, Md.), 1 2, 188-94

Horseman, N D; Nollin, L J 1985. The mitogenic, but not differentiative, response of crop tissue to prolactin is circadian phase dependent. Endocrinology, 116 5, 2085-9

Pauly, J R; Horseman, N D 1985. Anticholinergic agents do not block light-induced circadian phase shifts. Brain research, 348 1, 163-7

Pukac, L A; Horseman, N D 1984. Regulation of pigeon crop gene expression by prolactin. Endocrinology, 114 5, 1718-24

Horseman, N D; Ehret, C F 1982. Glucocorticosteroid injection is a circadian zeitgeber in the laboratory rat. The American journal of physiology, 243 3, R373-8

Horseman, N D; Meier, A H 1979. Circadian phase-dependent prolactin mechanisms in hepatic lipogenesis of a teleost. The Journal of endocrinology, 82 3, 367-72

Horseman, N D; Meier, A H 1979. Circadian-dependent prolactin effects: hepatic RNA metabolism and prostaglandin mediation. General and comparative endocrinology, 38 3, 269-74

Horseman, N D; Meier, A H 1978. Prostaglandin the osmoregulatory role of prolactin in a teleost. Life sciences, 22 17, 1485-90

Ottenweller, J E; Meier, A H; Ferrell, B R; Horseman, N D; Proctor, A 1978. Extrapituitary regulation of the circadian rhythm of plasma corticosteroid concentration in rats. Endocrinology, 103 5, 1875-9

Horseman, N D; Meier, A H; Culley, D D 1976. Daily variations in the effects of disturbance on growth, fattening, and metamorphosis in the bullfrog (Rana catesbeiana) tadpole. The Journal of experimental zoology, 198 3, 353-8

Dugan, A L; Malarkey, W B; Schwemberger, S; Jauch, E C; Ogle, C K; Horseman, N D . Serum levels of prolactin, growth hormone, and cortisol in burn patients: correlations with severity of burn, serum cytokine levels, and fatality. The Journal of burn care & rehabilitation, 25 3, 306-13

Dugan, Amy L; Gregerson, Karen A; Neely, Alice; Gardner, Jason; Noel, Greg J; Babcock, George F; Horseman, Nelson D . Mice treated with a benzodiazepine had an improved survival rate following Pseudomonas aeruginosa infection. Journal of burn care & research : official publication of the American Burn Association, 31 1, 1-12

Horseman, N D; Engle, S J; Ralescu, A . The logic of signaling from the cell surface to the nucleus. Trends in endocrinology and metabolism: TEM, 8 4, 123-9

Matsuda, Manabu; Lockefeer, Jason A; Horseman, Nelson D . Aldolase C/zebrin gene regulation by prolactin during pregnancy and lactation. Endocrine, 20 1-2, 91-100

Naylor, Mathew J; Lockefeer, Jason A; Horseman, Nelson D; Ormandy, Christopher J . Prolactin regulates mammary epithelial cell proliferation via autocrine/paracrine mechanism. Endocrine, 20 1-2, 111-4