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Mammary Development
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Mammary Gland Development
Study Questions & References

Mammary Gland
Study Questions

Cross section of a beef heifer's udder.

Study Questions

1. The image below is a histological section of mamamry tissue from an ovariectomized gilt that had been treated with genistein (a phytoestrogen) for 10 days to induce estrogenic responses. What are the structures indicated by the 1, 2, 3, 4, and 5 on the image, respectively?

Histological slide of glit mammary tissue.

2. If you were to inhibit relaxin secretion during late pregnancy in the gilt what effect would it have on mammary growth? If you then let those gilts suckle piglets after birth, how would postpartum mammary growth be affected? How would milk yield and piglet growth be affected?

3. The image below is a cross-section through the udder of a heifer at 2 months of gestation. If you were able to excise the areas of the udder circled in red at this point, what would happen to further mammary development? Why?

Cross-section of a heifer's udder at 2 months pregnacy.


General references on mammary growth and develoment:

Anderson, R.R., 1978, Embryonic and fetal development of the mammary apparatus. In: Lactation: A Comprehensive Treatise, Vol. IV, Ed. B.L. Larson, Academic Press, New York, pp. 3-40.

Cunha, G.R. and Y.K. Hom., 1996, Role of mesenchymal-epithelial interactions in mammary gland development. J. Mammary Gland Biol. Neoplasia 1:21-35.

Das, R. and B.K. Vonderhaar. 1997. Prolactin as a mitogen in mammary cells. J. Mam. Gland Biol. Neoplasia 2:29-40.

Fendrick, J.L., A. M. Raafat, and S. Z. Haslam. 1998. Mammary gland growth and development from the postnatal period to postmenopause: ovarian steroid receptor ontogeny and regulation in the mouse. J. Mam. Gland Biol. Neoplasia 3:7-22.

Forsyth, IA 1986 Variation among species in the endocrine control of mammary growth and function: the roles of prolactin, growth hormone, and placental lactogen. J. Dairy Sci. 69:886-903.

Hovey, RC, McFadden TB, Akers RM 1999 Regulation of mammary gland growth and morphogenesis by the mammary fat pad: a species comparison. J. Mammary Gland Biol. Neoplasia 4:53-68

Imagawa W, Yang J, Guzman R, Nandi S 1994 Control of mammary gland development. Ch. 56 in The Physiology of Lactation, 2nd edition, Eds. Knobil, K, Neill, J., et al., Raven Press, NY, p. 1033.

Kleinberg, D.L. 1997. Early mammary development: growth hormone and IGF-1. J. Mam. Gland Biol. Neoplasia 2:49-58.

Knight, CH, Peaker, M 1982 Development of the mammary gland. J. Reprod. Fert. 65:521-536.

Raynaud, A., 1961, Morphogenesis of the mammary gland. In: Milk: The Mammary Gland and Its Secretion, Vol. 1., Eds. S.K. Kon and A.T Cowie, Academic Press, New York, pp.3-46.

Raynaud, A. 1971, Foetal development of the mammary gland and hormonal effects on its morphogenesis. In: Lactation, Ed. I.R. Falconer, Penn. St. Univ. Press, University Park, pp. 3-29.

Sakakura, T., 1987, Mammary embryogenesis. In: The Mammary Gland: Development, Regulation, and Function, Eds. C.W. Neville and M.C. Daniel, Plenum Press, New York, pp. 37-66.

Sinha, Y.N. and H.A. Tucker. 1969. Mammary development and pituitary prolactin levels of heifers from birth through puberty and during the estrous cycle. J. Dairy Sci. 52:507-512.

Tucker, H.A., 1987, Quantitative estimates of mammary growth during various physiological states: a review. J. Dairy Sci. 70:1958-1966.

Turner, C.W., 1952, The Mammary Gland. Lucas Bros. Columbia, MO.

Turner, C.W., 1939, The Comparative Anatomy of the Mammary Glands. Univ. Coop. Store, Columbia, MO.

Woodward, T.L., J.W. Xie, and S.Z. Haslam. 1998. The role of mammary stroma in modulating the proliferative response to ovarian hormones in the normal mammary gland. J. Mam. Gland Biol. Neoplasia 3:117-132.

Influence of energy intake on mammary development:

Buskirk, D.D., D.B. Faulkner, W.L. Hurley, D.J. Kesler, F.A. Ireland, T.G. Nash, J.C. Castree,and J.L. Vicini. 1996. Growth, reproductive performance, mammary development, and milk production of beef heifers as influenced by prepubertal dietary energy and administration of bovine somatotropin. J. Anim. Sci. 74:2649-2662

Carstens, G.E., D.E. Glaser, F.M. Byers, L.W. Greene, and D.K. Lunt. 1997. Effects of bovine somatotropin treatment and intermittent growth pattern on mammary gland development in heifers. J. Anim. Sci. 75:2378-2388

Grings, E.E., D.M. deAvila, R.G. Eggert, and J.J. Reeves. 1990. Conception rate, growth, and lactation of dairy heifers treated with recombinant somatotropin. J. Dairy Sci. 73:73-77.

Harrison, R.D., I.P. Reynolds, and W. Little. 1983. A quantiative analysis of mammary glands of dairy heifers reared at different rates of live weight gain. J. Dairy Res. 50:405-412.

Lacasse, P., E. Block, and D. Petitclerc. 1994. Effect of plane of nutrition before and during gestation on the concentration of hormones in dairy heifers. J. Dairy Sci. 77:439-445.

McFadden, T.B., T.E. Daniel, and R.M. Akers. 1990. Effects f plane of nutrition, growth hormone and unsaturated fat on mammary growth in prepubertal lambs. J. Anim. Sci. 68:3171-3179.

Park, C.S., G.M. Erickson, Y.J. Choi, and G.D. Marx. 1987. Effect of compensatory grwoth on regulation of growth and lactation: Response of dairy heifers to a stair-step growth pattern. J. Anim. Sci. 64:1751-1758

Petitclerc, D. R.D. Kineman, S.A. Zinn, and H.A. Tucker. 1985. Mammary growth response of Holstein heifers to photoperiod. J. Dairy Sci. 68:86-90.

Ringuet, H. D. Petitclerc, M.T. Sorensen, P. Gaudreau, G. Pelletier, J. Morisset, Y. Couture,a nd P. Brazeau. 1989. Effect of human somatotropin-releasing factor and photoperiods on carcass parameters and mammary gland development of dairy heifers. J. Dairy Sci. 72:2928-2935.

Sejrsen, K. 1978. Mammary development and milk yield in relationto growth rate in dairy and dual-purpose heifers. Acta Agric. Scand. 28:41-46.

Sejrsen, K., J. Foldager, M.T. Sorenson, R.M. Akers, and D.E. Bauman. 1986. Effect of exogenous bovine somatotropin on pubertal mammary development in heifers. J. Dairy Sci. 69:1528

Sejrsen, K., J.T. Huber, H.A. Tucker, and R.M. Akers. 1982. Influence of nutrition on mammary development in pre- and postpubertal hiefers. J. Dairy Sci. 65:793

Sejrsen, K., J.T. Huber, and H.A. Tucker. 1983. Influence of amount fed on hormone concentrations and their relationship to mammary growth in heifers. J. Dairy Sci. 66:845.

Waldo, D.R., A.V. Capuco, and C.E. Rexroad, Jr. 1989. Replacement heifer growth rates affects milk producing ability. Feedstuffs, November 27, 1989, pp. 15-17.

Stelwagen, K. and D.G. Grieve. 1990. Effect of plane of nutrition on growth and mammary gland development in Holstein heifers. J. Dairy Sci. 73:2333-2341.

Growth facotrs and mammary development

Daniel, C.W., S. Robinson, and G.B. Silberstein, 1996, The role of TGF-ß in patterning and growth of the mammary ductal tree. J. Mam. Gland Biol. Neoplasia 1:331-342.

DiAugustine, R.P., R. G. Richards, adn J. Sebastian. 1997. EGF-Related peptides and their receptors in mammary gland development. J. Mam. Gland Biol. Neoplasia 2:109-118.

Plaut, K., 1993, Role of epidermal growth factor and transforming growth factors in mammary development and lactation. J. Dairy Sci. 76:1526.

Smith, G.H. 1996, TGF-ß and functional differentiation. J. Mam. Gland Biol. Neoplasia 1:343-352.

Hormonal control of mammary growth during pregnancy

Hayden, T. J., C. R. Thomas, S. V. Smith, and I. A. Forsyth. 1980. Placental lactogen in the goat in relation to stage of gestation, number of fetuses, metabolites, progesterone and time of day. J. Endocrinol. 86:279-290

Hurley, W.L.; Doane, R.M.; O’Day-Bowman, M.B.; Winn, R.J.; Mojonnier, L.E.; Sherwood, O.D. (1991) Effect of relaxin on mammary development in ovariectomized pregnant gilts. Endocrinology 128:1285-1290.

O’Day, M.B.; Winn, R.J.; Easter, R.A.; Dziuk, P.J.; Sherwood, O.D. (1989) Hormonal control of the cervix in pregnant gilts. II. Relaxin promotes changes in the physical properties of the cervix in ovariectomized hormone-treated pregnant gilts. Endocrinology 125:3004-3010.

Thomford, P. J. H. K. L. Sander, J. Z. Kendall, O. D. Sherwood, and P. J. Dziuk. 1984. Maintenance of pregnancy and levels of progesterone and relaxin in the serum of gilts following a stepwise reduction in the number of corpora lutea. Biol. Reprod. 31:494-498.

Winn, R.J.; Baker, M.D.; Merle, C.A.; Sherwood, O.D. (1994) Individual and combined effects of relaxin, estrogen, and progesterone in ovariectomized gilts. II. Effects on mammary development. Endocrinology 135, 1250-1255.

Mammary growth during lactation

Anderson, R. R., M. H. Lu, J. J. Trojanor, and J. L. Clark. 1974. Milk production, wet weight, dry weight, potassium, and nucleic acid measurements of cow udders. J. Dairy Sci. 57:1350.

Anderson, R. R., J. R. Harness, A. F. Snead, and M. F. Salah. 1981. Mammary growth patterns in goats during pregnancy and lactation. J. Dairy Sci. 64:427.

Anderson, R. R. M. S. Salah, J. R. Harness, and A. F. Snead. 1982. Mammary growth patterns in guinea pigs during puberty, pregnancy and lactation. Biol. Reprod. 26:620.

Anderson, R. R. and I. M. Wahab. 1990. Changes in parenchyma and stroma of goat udders during pregnancy, lactation, and involution. Small Ruminant Res. 3:605.

Kim, S. W., W. L. Hurley, I. K. Han, and R. A. Easter. 1999. Changes in tissue composition associated with mammary gland growth during lactation in sows. J. Anim. Sci. 77:2510-2516.

Munford, R. E. 1963. Changes in the mammary glands of rats and mice during pregnancy, lactation and involution. 2. Levels of deoxyribonucleic acid, and alkaline and acid phosphatases. J. Endocrinol. 28:17.

Swanson, E. W., and J. I. Poffenberger. 1979. Mammary gland development of dairy heifers during their first gestation. J. Dairy Sci. 62:702.

Tucker, H. A. 1966. Regulation of mammary nucleic acid content by various suckling intensities. Am. J. Physiol. 210:1209.

Tucker, H. A. 1969. Factors affecting mammary gland cell numbers. J. Dairy Sci. 52:720.

Zaleski, H.M.; Winn, R.J.; Jennings, R.L.; Sherwood, O.D. (1996) Effects of relaxin on lactational performance in ovariectomized gilts. Biology of Reproduction 55:671-675.

Cystic Ovaries

Gaverick HA 1997 Ovarian follicular cysts in dairy cows. J Dairy Sci 80:995-1004

Hamilton SA, Garverick HA, Keisler DH, Xu ZZ, Loos K, Youngquist RS, Salfen BE 1995 Characterization o ovarian follicular cysts and associated endocrine profiles in dairy cows. Biol Reprod 53:890-898

Kesler DJ, Garverick HA 1982 Ovarian cysts in dairy cattle: A review. J Anim Sci 55:1147-1159

Kesler DJ, Garverick HA, Caudle AB, Bierschwal CJ, Elmore RG, Youngquist RS 1978 Clinical and endocrine responses of dairy cows with ovarian cysts to GnRH and PGF2 alpha. J anim Sci 46:719-725

Kesler DJ, Garverick HA, Caudle AB, Elmore RG, Bierschwal CJ 1980 Reproductive hormone and ovarian changes in cows with ovarian cysts. J Dairy Sci 63:166-170

Kinder JE, Kojima FN, Bergfeld EGM, Wehrman ME, Fike KE 1996 Progestin and estrogen regulation of pulsatile LH release and development of persistent ovarian follicles in cattle. J Anim Sci 74:1424-1440

Peter AT 2004 An update on cystic ovarian degeneration in cattle. Reprod Dom Anim 39:1-7

Stevenson JS, Call EP 1988 Reproductive disorders in the periparturient dairy cow. J Dairy Sci 71:2572-2583

Ovarian tumors in cattle

Durba CR, de Briones EA, long SE 1972 A case of granulose-cell tumor in a heifer.. Vet Rec 91:9-11

Leder RR, Lane Vm, Barrett DP 1988 Ovariectomy as treatment for granulose cell tumor in a heifer. J Am Vet Med Assoc 192:1299-1300

Roberts SJ 1953 An ovarian tumor in a heifer. .Cornell Vet 43:531-536

Short RV, Shorter DR, Linzell JL 1963 Granulosa cell tumor of the ovary in a virgin heifer. J Endocrinol 27:327-332

Vestweber JG, Cole DE, Quadri SK, Schmidt MK 1986 Premature mammary development in a heifer with abdominal mesothelioma. J Am Vet Med Assoc 189:804-805

Whitacre MD, Van Camp SD, Maclachlan NJ, Umstead JA 1988 Premature lactation in a heifer with a sex cord-stromal tumor. J Am Vet Med Assoc 193:946-948

Pituitary tumors

Colao A, Loche S, Cappa M, Di Sarno A, Landi ML, Sarnacchiaro F, Facciolli G, Lombardi, G 1998 Prolactinomas in children and adolescents, clinical presentation and long-term follow-up. J Clin Endocrinol Metab 83:2777-2780

Colao A, Di Sarno A, Sarnacchiaro F, Ferone D, Di Renzo G, Merola B, Annunziato L, Lombardi, G 1997 Prolactinomas resistant to standard dopmanie agonists respond to chronic cabergoline treatment. J Clin Endocrinol Metab 82:876-883

Hutchinson J, Murphy M, Harries R, Skinner C 2000 Galactorrhoea and hyperprolactinaemia associated with protease-inhibitors. The Lancet 356:1003-004

Phytoestrogens and related topics

Adams NR   (1977)   Morphological changes in the organs of ewes grazing oestrogenic subterranean clover. Res Vet Sci 22:216-221

Adams NR (1995) Detection of the effects of phytoestrogens on sheep and cattle. J Anim Sci 73:1509-1515

Akiyama, T., J. Ishida, S. Nakagawa, H. Ogawara, S. Watanabe, N. Itoh, M. Shibuya, and Y. Fukami.   1987.   Genistein, a specific inhibitor of tyrosine-specific kinases.   J. Biol. Chem. 5592-5595.

Brown NM, Lamartiniere CA (1995)   Xenoestrogens alter mammary gland differentiation and cell proliferation in the rat.   Environ Health Perspect 103:708-713

Brown NM, Wang J, Cotroneo MS, Zhoa Y-X, Lamartiniere CA 1998 Prepubertal genistein treatment modulates TGF- a , EGF and EGF-receptor mRNAs and proteins in the rat mammary gland. Mol Cell Endorcinol 144:149-165

Chemopreventative Branch and Agent Development Committee, NCI, DCPC (1997)   Clinical development plan:   genistein.   J Cell Biochem 265:114-126

Drane HM, Wrathall AE, Patterson, DSP, Hebert CN   (1981) Possible oestrogenic effects of feeding soymeal to prepubertal gilts. Br Vet J 137:283-288

Eldridge-White R, Easter RA, Heaton DM, O'Day MB, Petersen GC, Shanks RD, Tarbell MK, Sherwood OD (19898) Hormonal control of the cervix in pregnant gilts. I. Changes in the physical properties of the cervix correlate temporally with elevated serum levels of estrogen and relaxin. Endocrinology 125:2996-3003

Fan, G. and J. A. Rillema.   1992.   Effect of tyrosine kinase inhibitor, genistein, on the actions of prolactin in cultured mouse mammary tissues.   Mol. Cell. Endocrinol. 83:51-55.

Kaldas RS, Hughes, Jr, CL (1989) Reproductive and general metabolic effects of phytoestrogens in mammals. Reprod Tox 3:81-89

Lamartiniere CA, Moore JB, Brown NM, Thompson R, Hardin MJ, Barnes S (1995)   Genistein suppresses mammary cancer in rats. Carcinogenesis 16:2833-2840

Lindzey J, Korach KS (1998) Estrogen action on the female reproductive tract. In Encyclopedia of Reproduction , Vol 2, ed E Knobil and JD Neill, Academic Press, NY, pp 79-86

Liu K-S (1997) Soybeans: Chemistry, Technology, and Utilization . Chapman and Hall, New York, NY

Lundh T (1995) Metabolism of estrogenic isoflavones in domestic animals. Proc Soc Exp Biol Med 208:33-339

Murrill WB, Brown NM, Zhang J-X, Manzolillo PA, Barnes S, Lamartiniere CA (1996)   Prepubertal genistein exposure suppresses mammary cancer and enhances gland differentiation in rats.   Carcinogenesis 17:1451-1457

Nwannenna AI, Lundh TJ, Madej A, Fredriksson G, Bjornhag G. (1995) Clinical changes in ovariectomized ewes exposed to phytoestrogens and 17 beta-estradiol implants. Proc Soc Exp Biol Med 208:92-7

Santell RC, Chang YC, Nair MG, Helferich WG. (1997) Dietary genistein exerts estrogenic effects upon the uterus, mammary gland and the hypothalamic/pituitary axis in rats. J Nutr 127:263-269

Suttner A, Danilovich NA, Banz WJ, Winters TA (1998) The effects of the phytoestrogen daidzein on in situ apoptosis in primary porcine granulosa cells. Biol Reprod 58(Suppl 1):74

Verdeal, K., R. B. Brown, T. Richardson, and D. S. Ryan.   Affinity of phytoestrogens for estradiol-binding proteins and effect of coumestrol on growth of 7,12-dimethylbenz[a]anthracene-induced rat mammary tumors.   J. Natl. Cancer Inst. 64:285-290.

Wang W, Tanaka Y, Han Z, Higuchi CM 1995 Proliferative response of mammary glandular tissue to formononetin.   Nutr Cancer 23:131-140

Whitten PL, Lewis C, Naftolin F (1993) A phytoestrogen induces the premature anovulatory syndrome in lactationally exposed female rats. Biol Reprod 49:1117-1121

Whitten PL, Russell E, Naftolin F (1992) Effects of a normal, human-concentration, phytoestrogen diet on rat uterine growth. Steroids 57:98-106

Winn RJ, Baker MD, Sherwood OD (1994a) Individual and combined effects of relaxin, estrogen, and progesterone in ovariectomized gilts. I. Effects on the growth, softening and histological properties of the cervix. Endocrinology 135:1241-1249

Winn RJ, Baker MD, Merle CA, Sherwood OD (1994b) Individual and combined effects of relaxin, estrogen, and progesterone in ovariectomized gilts. II. Effects on mammary development. Endocrinology 135:1250-1255


Tucker HA (1994) Lactation and its hormonal control. Ch. 57, In: The Physiology of Reproduction, 2nd edition, Eds. E. Knobil, J. Neill, et al. Raven Press, Ltd., New York. p. 1065.

Collier, R.J., D.E. Bauman, and R.L. Hay. 1977. Effects of reserpine on milk production and serum prolactin of cows hormonally induced into lactation. J. Dairy Sci. 60: 896.

Sheeham, D. M. 1998. Herbal medicines, phytoestrogens, and toxicity: risk:benefit considerations. Proc. Soc. Exp. Biol. Med. 217: 379-385

Zava, Dollbaum, and Blen. 1998. Estrogen and progestin bioactivity of foods, herbs, and spices. Proc. Soc. Exp. Biol. Med. 217: 369.

Bacon CW 1995 Toxic endophyte-infected fescue and range grasses: historic perspectives. J Anim Sci 73:861-870

Cheeke PR 1995 Endogenous toxins and mycotoxins in forage grasses and their effects on livestock. J Anim Sci 73:909-918

Porter JK 1995 Analysis of endophyte toxins: fescue and other grasses toxic to livestock. J Anim Sci 73:871-880

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