Endocrine changes during the peripartal period related to colostrogenesis in mammalian species.
- Review
- Journal Article
Summary
This research review examines the hormonal and functional changes across varying mammalian species that relate to the production of colostrum, the initial high-nutrient milk produced following birth. The species studied include ungulates (cattle, sheep, goats, pigs, horses), rodents (rats, mice), rabbits, carnivores (cats, dogs), and humans. It particularly focuses on a pattern of declining activity of progesterone towards the end of pregnancy, and how this process varies across species; colostrum production is pivotal for newborn survival where placental transfer of immunoglobulins does not occur or is partial.
Key Endocrine Changes during the Peripartal Period
The paper examines specific endocrine changes that occur towards the end of an animal’s pregnancy. This understanding is crucial for the management of parturition (the action of giving birth) and lactogenesis (the initiation of milk secretion) in diverse mammalian species:
- The declining activity of gestagens, specifically progesterone, is found to be critical in late pregnancy to initiate both parturition and lactation across species.
- However, the exact endocrine pathways and timing of gestagen withdrawal vary between different mammal types.
- The research emphasizes that in certain species where a corpus luteum (a hormone-secreting body in the uterus) is sustained throughout the entire pregnancy, parturition and lactogenesis are initiated by prostaglandin-induced luteolysis (the degeneration of the corpus luteum).
Differences in Gestagens Activity Reduction
Another observation from the review revolves around the processes through which different species achieve reduced levels of gestagens during pregnancy:
- The decline of gestagen activity at the end of pregnancy is a complex process in species where placental gestagen production takes over, compared to those with a sustained corpus luteum.
- In sheep, there is a redirection of hormone production away from progesterone towards estradiol to achieve lower gestagen activity.
- In humans and horses, the uterus eventually becomes insensitive to progesterone, leading to parturition despite high progesterone concentrations. However, lactogenesis is not completed until progesterone levels drop. In contrast to humans, horses need to begin lactogenesis before giving birth as horse newborns need immediate immunization via colostrum.
Implications of the Research
Understanding the endocrine changes and related pathways during pregnancy is essential as it allows researchers to control key events relating to colostrogenesis (formation of colostrum), parturition, and the start of lactation. This knowledge, however, remains incomplete in many species:
- There is a call for further research to clarify the processes that induce lactogenesis prior to parturition in horses, for instance.
- The research’s implications can also influence how neonatal intensive care is managed across species, particularly for those creatures where colostrum and immunoglobulin intake is critical for neonatal survival immediately after birth.
Cite This Article
Publication
Researcher Affiliations
- Veterinary Physiology, Vetsuisse Faculty, University of Bern, Bremgartenstrasse 109a, 3012 Bern, Switzerland.
- Veterinary Physiology, Vetsuisse Faculty, University of Bern, Bremgartenstrasse 109a, 3012 Bern, Switzerland.
- Department of Animal Science, Penn State University, University Park, PA 16802, USA.
- Veterinary Physiology, Vetsuisse Faculty, University of Bern, Bremgartenstrasse 109a, 3012 Bern, Switzerland.
MeSH Terms
- Pregnancy
- Female
- Swine
- Cattle
- Humans
- Rats
- Horses
- Animals
- Rabbits
- Sheep
- Dogs
- Mice
- Progestins
- Dinoprost
- Placenta / metabolism
- Parturition
- Colostrum / metabolism
- Progesterone / metabolism
- Rodentia / metabolism
Conflict of Interest Statement
References
- Akers RM. Lactation and the mammary gland. 1st ed.Ames, IA: Wiley-Blackwell.
- Alekseev NP. Physiology of human female lactation. 1st ed.Cham, CH: Springer.
- Aurich C, Aurich JE, Parvizi N. Opioidergic inhibition of luteinising hormone and prolactin release changes during pregnancy in pony mares. J. Endocrinol. 169:511–518.
- Bagna B, Schwabe C, Anderson LL. Effect of relaxin on facilitation of parturition in dairy heifers. J. Reprod. Fertil. 91:605–615.
- Banks DR, Paape SR, Stabenfeldt GH. Prolactin in the Cat: I. Pseudopregnancy, pregnancy and lactation. Biol. Reprod. 28:923–932.
- Barrena JC, Griffith DR. Role of maternal and foetal adrenal activity in the development of the rat mammary gland near term. J. Endocrinol. 68:439–443.
- Barrington GM, McFadden TB, Huyler MT, Besser TE. Regulation of colostrogenesis in cattle. Livest. Prod. Sci. 70:95–104.
- Bathgate R, Moniac N, Bartlick B, Schumacher M, Fields M, Ivell R. Expression and regulation of relaxin-like factor gene transcripts in the bovine ovary: differentiation-dependent expression in theca cell cultures. Biol. Reprod. 61:1090–1098.
- Baumrucker CR, Bruckmaier RM. Colostrogenesis: IgG1 transcytosis mechanisms. J. Mammary Gland Biol. Neoplasia. 19:103–117.
- Baumrucker CR, Dechow CD, Macrina AL, Gross JJ, Bruckmaier RM. Mammary immunoglobulin transfer rates following prepartum milking. J. Dairy Sci. 99:9254–9262.
- Baumrucker CR, Macrina AL, Bruckmaier RM. Colostrogenesis: role and mechanism of the bovine Fc receptor of the neonate (FcRn). J. Mammary Gland Biol. Neoplasia. 26:419–453.
- Baumrucker CR, Burkett AM, Magliaro-Macrina AL, Dechow CD. Colostrogenesis: mass transfer of immunoglobulin G1 into colostrum. J. Dairy Sci. 93:3031–3038.
- Baumrucker CR, Zbinden RS, Van Dorland HA, Remmelink GJ, Kemp B, Van Knegsel ATM, Bruckmaier RM. Continuous milking of dairy cows disrupts timing of peak IgG concentration appearance in mammary secretions. J. Dairy Res. 81:403–409.
- Bazer FW, Vallet JL, Roberts RM, Sharp DC, Thatcher WW. Role of conceptus secretory products in establishment of pregnancy. J. Reprod. Fertil. 76:841–850.
- Bigler NA, Bruckmaier RM, Gross JJ. Implications of placentation type on species-specific colostrum properties in mammals. J. Anim. Sci. 100(12):skac287.
- Blanks AM, Brosens JJ. Progesterone action in the myometrium and decidua in preterm birth. Facts Views Vis. Obgyn. 4:33–43.
- Blum JW. Nutritional physiology of neonatal calves. J. Anim. Physiol. Anim .Nutr. 90:1–11.
- Bongers-Binder S, Burgardt A, Seeger H, Voelter W, Lippert TH. Distribution of immunoreactive relaxin in the genital tract and in the mammary gland of non-pregnant women. Clin. Exp. Obstet. Gynecol. 18:161–164.
- Borelli E, Jonsson NN, Denholm KS. Associations between putative risk factors and poor colostrum yield in Holstein Friesian cattle. Res. Vet. Sci. 149:74–81.
- Brandon MR, Watson DL, Lascelles AK. The mechanism of transfer of immunoglobulin into mammary secretion of cows. Aust. J. Exp. Biol. Med. Sci. 49(6):613–623.
- Braun BC, Zschockelt L, Dehnhard M, Jewgenow K. Progesterone and estradiol in cat placenta--biosynthesis and tissue concentration. J.. Steroid Biochem Mol. Biol. 132:295–302.
- Capuco AV, Tucker HA. Progesterone inhibition of glucocorticoid binding to mammary tissue from lactating and nonlactating cows. Proc. Soc. Exp. Biol. Med. 164:386–393.
- Casey TM, Plaut K. The role of glucocorticoids in secretory activation and milk secretion, a historical perspective. J. Mammary Gland Biol. Neoplasia. 12:293–304.
- Castro N, Capote J, Bruckmaier RM, Argüello A. Management effects on colostrogenesis in small ruminants: a review. J. Appl. Anim. Res. 39:85–93.
- Challis JRG, Bloomfield FH, Booking AD, Casciani V, Chisaka H, Connor K, Dong X, Gluckman P, Harding JE, Johnstone J. Fetal signals and parturition. J. Obstet. Gynaecol. Res. 31:492–499.
- Challis JR, Harrison FA, Heap RB. The extraction of oestrogens, and the rate of secretion of oestrone and oestradiol-17beta by the uterus in the pregnant sheep. J. Endocrinol. 61:277–284.
- Chavatte P, Holtan D, Ousey JC, Rossdale PD. Biosynthesis and possible biological roles of progestagens during equine pregnancy and in the newborn foal. Equine Vet. J. 24:89–95.
- Chilliard Y, Delouis C, Smith MC, Sauvant D, Morand-Fehr P. Mammary metabolism in the goat during normal or hormonally-induced lactation. Reprod. Nutr. Dev. 26:607–615.
- Collier RJ, Tucker HA. Regulation of cortisol uptake in mammary tissue of cows. J. Dairy Sci. 61:1709–1714.
- Concannon PW, Butler WR, Hansel W, Knight PJ, Hamilton JM. Parturition and lactation in the bitch: serum progesterone, cortisol and prolactin. Biol. Reprod. 19:1113–1118.
- Concannon PW, Powers ME, Holder W, Hansel W. Pregnancy and parturition in the bitch. Biol. Reprod. 16:517–526.
- Conley AJ. Review of the reproductive endocrinology of the pregnant and parturient mare. Theriogenology 86:355–365.
- Conley AJ, Ball BA. Steroids in the establishment and maintenance of pregnancy and at parturition in the mare. J. Reprod. Fertil. 158:197–208.
- Conley AJ, Reynolds LP. Steroidogenesis and the initiation of parturition. p. 399–413.
- Conneely M, Berry DP, Sayers R, Murphy JP, Lorenz I, Doherty ML, Kennedy E. Factors associated with the concentration of immunoglobulin G in the colostrum of dairy cows. Animal 7:1824–1832.
- Costa MA. The endocrine function of human placenta: an overview. Reprod. Biomed. Online. 32:14–43.
- Currie WB, Gorewit RC, Michel FJ. Endocrine changes, with special emphasis on oestradiol-17β, prolactin and oxytocin, before and during labour and delivery in goats. J. Reprod. Fertil. 82:299–308.
- Currie WB, Thorburn GD. Parturition in goats: studies on the interactions between the foetus, placenta, prostaglandin F and progesterone before parturition, at term or at parturition induced prematurely by corticotrophin infusion of the fetus. J. Endocrinol. 73:263–278.
- Daels PF. Induction of lactation and adoption of the orphan foal. Page in AAEP Resort Symposium, Rome IT.
- Dai Y, Ivell R, Liu X, Janowski D, Anand-Ivell R. Relaxin-family peptide receptors 1 and 2 are fully functional in the bovine. Front. Physiol. 8:359.
- Davis AJ, Fleet IR, Goode JA, Hamon MH, Maule Walker FM, Peaker M. Changes in mammary function at the onset of lactation in the goat: correlation with hormonal changes. J. Physiol. 288:33–44.
- Davis SL, Reichert LE, Niswender GD. Serum levels of prolactin in sheep as measured by radioimmunoassay. Biol. Reprod. 4:145–153.
- Delouis C. Physiology of colostrum production. Ann. Rech. Vet. 9:193–203.
- Devillers N, Farmer C, Mounier AM, Le Dividich J, Prunier A. Hormones, IgG and lactose changes around parturition in plasma, and colostrum or saliva of multiparous sows. Reprod. Nutr. Dev. 44:381–396.
- Eissa HM, El-Belely MS. Sequential changes in plasma progesterone, total oestrogens and corticosteroids in cow throughout pregnancy and around parturition. Br. Vet. J. 146:24–29.
- Ellendorff F, Taverne M, Elsaesser F, Forsling M, Parvizi N, Naaktgeboren C, Smidt D. Endocrinology of parturition in the pig. Anim. Reprod. Sci. 2:323–334.
- Estergreen VL, Frost OL, Gomes WR, Erb RE, Bullard JF. Effect of ovariectomy on pregnancy maintenance and parturition in dairy cows. J. Dairy Sci. 50:1293–1295.
- Farmer C, Quesnel H. Current knowledge on the control of onset and cessation of colostrogenesis in swine. J. Anim. Sci. 98:S133–S139.
- Farmer C, Sorensen MT, Petitclerc D. Inhibition of prolactin in the last trimester of gestation decreases mammary gland development in gilts. J. Anim. Sci. 78:1303–1309.
- Fleet IR, Goode JA, Hamon MH, Laurie MS, Linzell JL, Peaker M. Secretory activity of goat mammary glands during pregnancy and the onset of lactation. J. Physiol. 251:763–773.
- Forsyth IA. 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.
- Frankshun AL, Ho TY, Reimer DC, Chen J, Lasano S, Steinetz BG, Bartol FF, Bagnell CA. Characterization and biological activity of relaxin in porcine milk. Reproduction 141(3):373–380.
- Gavin K, Neibergs H, Hoffman A, Kiser JN, Cornmesser MA, Haredasht SA, Martínez-López B, Wenz JR, Moore DA. Low colostrum yield in Jersey cattle and potential risk factors. J Dairy Sci. 101(7):6388–6398.
- Gräf KJ. Serum estrogen, progesterone and prolactin concentrations in cyclic, pregnant and lactating beagle dogs. J. Reprod. Fertil. 52(1):9–14.
- Graham JD, Clarke CL. Physiological action of progesterone in target tissues. Endocr Rev. 18(4):502–519.
- Gross JJ, Kessler EC, Bjerre-Harpoth V, Dechow C, Baumrucker CR, Bruckmaier RM. Peripartal progesterone and prolactin have little effect on the rapid transport of immunoglobulin G into colostrum of dairy cows. J. Dairy Sci. 97:2923–2931.
- Grunert E, Ahlers D, Heuwieser W. The role of endogenous estrogens in the maturation process of the bovine placenta. Theriogenology 31:1081–1091.
- Hennessy DP, Coghlan JP, Hardy KJ, Scoggins BA, Wintour EM. The origin of cortisol in the blood of fetal sheep. J. Endocrinol. 95:71–79.
- Hilliard J. Corpus luteum function in guinea pigs, hamsters, rats, mice and rabbits. Biol. Reprod. 8:203–221.
- Hoffmann B, Höveler R, Nohr B, Hasan SH. Investigations on hormonal changes around parturition in the dog and the occurrence of pregnancy-specific non conjugated oestrogens. Exp. Clin. Endocrinol. 102:185–189.
- Holt JA. Regulation of progesterone production in the rabbit corpus luteum. Biol. Reprod. 40:201–208.
- Hombach-Klonisch S, Tetens F, Kauffold J, Steger K, Fischer B, Klonisch T. Molecular cloning and localization of caprine relaxin-like factor (RLF) mRNA within the goat testis. Mol. Reprod. Dev. 53:135–141.
- Huang SC, Hu ZL, Hasler-Rapacz J, Rapacz J. Preferential mammary storage and secretion of immunoglobulin gamma (IgG) subclasses in swine. J. Reprod. Immunol. 21:15–28.
- Hurley WL. Mammary gland growth in the lactating sow. Livest. Prod. Sci. 70:149–157.
- Inoue R, Tsukahara T. Composition and physiological functions of the porcine colostrum. Anim. Sci. J. 92:e13618.
- Igwebuike UM. A review of uterine structural modifications that influence conceptus implantation and development in sheep and goats. Anim. Reprod. Sci. 112:1–7.
- Ireland FA, Loch WE, Worthy K, Anthony RV. Effects of bromocriptine and perphenazine on prolactin and progesterone concentrations in pregnant pony mares during late gestation. J. Reprod. Fertil. 92:179–186.
- Ivell R, Balvers M, Pohnke Y, Telgmann R, Bartsch O, Milde-Langosch K, Bamberger AM, Einspanier A. Immunoexpression of the relaxin receptor LGR7 in breast and uterine tissues of humans and primates. Reprod. Biol. Endocrinol. 1:114.
- Jahn GA, Rastrilla AM, Deis RP. Correlation of growth hormone secretion during pregnancy with circulating prolactin in rats. J. Reprod. Fertil. 98:327–333.
- Jewgenow K, Meyer HHD. Comparative binding affinity study of progestins to the cytosol progestin receptor of endometrium in different mammals. Gen. Comp. Endocrinol. 110:118–124.
- Kehoe SI, Jayarao BM, Heinrichs AJ. A survey of bovine colostrum composition and colostrum management practices on Pennsylvania dairy farms. J. Dairy Sci. 90(9):4108–4116.
- Klein C. The role of relaxin in mare reproductive physiology: a comparative review with other species. Theriogenology 86:451–456.
- Kota SK, Gayatri K, Jammula S, Kota SK, Krishna SVS, Meher LK, Modi KD. Endocrinology of parturition. Indian J. Endocrinol. Metab. 17:50–59.
- Kowalewski MP. Luteal regression vs. prepartum luteolysis: regulatory mechanisms governing canine corpus luteum function. Reprod. Biol. 14:89–102.
- Kuhn NJ. Lactogenesis: the search for trigger mechanisms in different species. p. 165–192.
- Legacki EL, Corbin CJ, Ball BA, Scoggin KE, Stanley SD, Conley AJ. Steroidogenic enzyme activities in the pre- and post-parturient equine placenta. Reproduction 155:51–59.
- Legacki EL, Corbin CJ, Ball BA, Wynn M, Loux S, Stanley SD, Conley AJ. Progestin withdrawal at parturition in the mare. Reproduction 152:323–331.
- Legacki EL, Scholtz EL, Ball BA, Stanley SD, Berger T, Conley AJ. The dynamic steroid landscape of equine pregnancy mapped by mass spectrometry. Reproduction 151:421–430.
- Li XQ, Zhu P, Myatt L, Sun K. Roles of glucocorticoids in human parturition: a controversial fact?. Placenta 35:291–296.
- Liggins GC, Fairclough RJ, Grieves SA, Forster CS, Knox BS. Parturition in the sheep. Ciba Found Symp. 47:5–30.
- Lyons WR. Experiments with hypophyseal lactogenic hormone on normal ovariectomized and hypophysectomized dogs. Proc. Soc. Exp. Biol. Med. 31:303–305.
- Magliaro AL, Kensinger RS, Ford SA, O’Connor ML, Muller LD, Graboski R. Induced lactation in nonpregnant cows: profitability and response to bovine somatotropin. J. Dairy Sci. 87:3290–3297.
- Malone L, Opazo JC, Ryan PL, Hoffmann FG. Progressive erosion of the Relaxin1 gene in bovids. Gen. Comp. Endocrinol. 252:12–17.
- Mayer B, Doleschall M, Bender B, Bartyik J, Bosze Z, Frenyó LV, Kacskovics I. Expression of the neonatal Fc receptor (FcRn) in the bovine mammary gland. J Dairy Res. 72:107–112.
- McCue PM, Sitters S. Lactation. p. 2277–2290.
- McFadden TB, Akers RM, Kazmer GW. Alpha-lactalbumin in bovine serum: relationships with udder development and function. J. Dairy Sci. 70:259–264.
- McFadden TB, Besser TE, Barrington GM. Regulation of immunoglobulin transfer into mammary secretions of ruminants. p. 133–152.
- Menchetti L, Andoni E, Barbato O, Canali C, Quattrone A, Vigo D, Codini M, Curone G, Brecchia G. Energy homeostasis in rabbit does during pregnancy and pseudopregnancy. Anim. Reprod. Sci. 218:106505.
- Min G, Sherwood OD. Identification of specific relaxin-binding cells in the cervix, mammary glands, nipples, small intestine, and skin of pregnant pigs. Biol. Reprod. 55:1243–1252.
- Miura H, Yamazaki T, Kikuchi M, Sakaguchi M. Plasma steroid hormone concentrations and their relationships in Suffolk ewes during gestation and parturition. Anim. Sci. J. 90:1426–1431.
- Mixner JP, Turner CW. Role of estrogen in the stimulation of mammary lobule-alveolar growth by progesterone and by the mammogenic lobule-alveolar growth factor of the anterior pituitary. Endocrinology 30:591–597.
- Munford RE. 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. 28:17–34.
- Murphy BM, Drennan MJ, O’Mara FP, Earley B. Cow serum and colostrum immunoglobulin (IgG1) concentration of five suckler cow breed types and subsequent immune status of their calves. Irish J. Agric. Food Res. 44:205–213.
- Musah AI, Schwabe C, Anderson LL. Acute decrease in progesterone and increase in estrogen secretion caused by relaxin during late pregnancy in beef heifers. Endocrinology 120(1):317–324.
- Nagel C, Erber R, Bergmaier C, Wulf M, Aurich J, Möstl E, Aurich C. Cortisol and progestin release, heart rate and heart rate variability in the pregnant and postpartum mare, fetus and newborn foal. Theriogenology 78:759–767.
- Negatu Z, McNitt JI. Hormone profiles and nest-building behavior during the periparturient period in rabbit does. Anim. Reprod. Sci. 72:125–135.
- Neville MC, McFadden TB, Forsyth I. Hormonal regulation of mammary differentiation and milk secretion. J. Mammary Gland Biol. Neoplasia. 7:49–66.
- Neville MC, Morton J. Physiology and endocrine changes underlying human lactogenesis II. J. Nutr. 131:3005S-3008S.
- Neville MC, Morton J, Umemura S. Lactogenesis: the transition from pregnancy to lactation. Pediatr. Clin. North Am. 48:35–52.
- Nguyen DA, Neville MC. Tight junction regulation in the mammary gland. J Mammary Gland Biol Neoplasia 3:233–246.
- Nishikawa S, Moore RC, Nonomura N, Oka T. Progesterone and EGF inhibit mouse mammary gland prolactin receptor and beta-casein gene expression. Am. J. Physiol. 267(5 Pt 1):C1467–1472.
- Nishiyama T, Tsumagari S, Ito M, Kimura J, Watanabe G, Taya K, Takeishi M. Immunohistochemical study of steroidogenic enzymes in the ovary and placenta during pregnancy in the dog. Anat. Histol. Embryol. 28:125–129.
- Nocek JE, Braund DG, Warner RG. Influence of neonatal colostrum administration, immunoglobulin, and continued feeding of colostrum on calf gain, health, and serum protein. J. Dairy Sci. 67:319–333.
- Pang WW, Hartmann PE. Initiation of human lactation: secretory differentiation and secretory activation. J. Mammary Gland Biol. Neoplasia. 12:211–221.
- Parry LJ, Vodstrcil LA, Madden A, Amir SH, Baldwin K, Wlodek ME, Nicholas KR. Normal mammary gland growth and lactation capacity in pregnant relaxin-deficient mice. Reprod. Fertil. Dev. 21:549–560.
- Patil NA, Rosengren KJ, Separovic F, Wade JD, Bathgate RAD, Hossain MA. Relaxin family peptides: structure-activity relationship studies. Br. J. Pharmacol. 174:950–961.
- Peaker M, Taylor E, Tashima L, Redman TL, Greenwood FC, Bryant-Greenwood GD. Relaxin detected by immunocytochemistry and northern analysis in the mammary gland of the guinea pig. Endocrinology 125:693–698.
- Pitia AM, Minagawa I, Abe Y, Kizaki K, Hamano KI, Sasada H, Hashizume K, Kohsaka T. Evidence for existence of insulin-like factor 3 (INSL3) hormone-receptor system in the ovarian corpus luteum and extra-ovarian reproductive organs during pregnancy in goats. Cell Tissue Res. 385:173–189.
- Pritchett LC, Gay CC, Besser TE, Hancock DD. Management and production factors influencing immunoglobulin G1 concentration in colostrum from Holstein cows. J. Dairy Sci. 74:2336–2341.
- Quesnel H, Farmer C. Review: nutritional and endocrine control of colostrogenesis in swine. Animal 13(S1):s26–s34.
- Randall GCB, Kendall JZ, Tsang BK, Taverne MAM. Endocrine changes following infusion of fetal pigs with corticotropin in litters of reduced numbers. Anim. Reprod. Sci. 23:109–122.
- Root Kustritz MV. Clinical management of pregnancy in cats. Theriogenology 66:145–150.
- Ryan KJ. Endocrine control of gestational length. A time to be born. Am. J. Obstet. Gynecol. 109:299–306.
- Schmidt PM, Chakraborty PK, Wildt DE. Ovarian activity, circulating hormones and sexual behavior in the cat. II. Relationships during pregnancy, parturition, lactation and the postpartum estrus. Biol. Reprod. 28:657–671.
- Scholtz EL, Krishnan S, Ball BA, Moeller BC, Stanley SD, McDowell KJ, Hughes AL, McDonnell DP, Conley AJ. Pregnancy without progesterone in horses defines a second endogenous biopotent progesterone receptor agonist, 5α-dihydroprogesterone. Proc. Natl. Acad. Sci. U. S. A. 111:3365–3370.
- Schuler G, Fürbass R, Klisch K. Placental contribution to the endocrinology of gestation and parturition. Anim. Reprod. 15:822–842.
- Sheldrick EL, Ricketts AP, Flint APF. Placental production of progesterone in ovariectomized goats treated with a synthetic progestagen to maintain pregnancy. J. Reprod. Fertil. 60:339–348.
- Shynlova O, Nadeem L, Zhang J, Dunk C, Lye S. Myometrial activation: novel concepts underlying labor. Placenta 92:28–36.
- Smith JJ, Capuco AV, Beal WE, Akers RM. Association of prolactin and insulin receptors with mammogenesis and lobulo-alveolar formation in pregnant ewes. Int. J. Biochem. 21:73–81.
- Smith VG, Edgerton LA, Hafs HD, Convey EM. Bovine serum estrogens, progestins and glucocorticoids during late pregnancy, parturition and early lactation. J. Anim. Sci. 36:391–396.
- Smith DE, Hixon DL, Moore DW, Van Kirk EA, Alexander BM, Anthony RV, Moss GE. Effects of porcine relaxin on induced parturition in beef heifers. Domest. Anim. Endocrinol. 13:469–476.
- Smith KH, Musah AI, Cho SJ, Schwabe C, Anderson LL. Continuous infusion of relaxin on periparturient progesterone, oxytocin and relaxin plasma concentrations and time of parturition in beef heifers. Anim. Reprod. Sci. 46:15–25.
- Stark A, Vachkova E, Wellnitz O, Bruckmaier R, Baumrucker C. Colostrogenesis: candidate genes for IgG1 transcytosis mechanisms in primary bovine mammary epithelial cells. J. Anim. Physiol. Anim. Nutr. 97:1114–1124.
- Stark A, Wellnitz O, Dechow C, Bruckmaier R, Baumrucker C. Colostrogenesis during an induced lactation in dairy cattle. J. Anim. Physiol. Anim. Nutr. 99:356–366.
- Steinetz BG, Horton L, Lasano S. The source and secretion of immunoactive relaxin in rat milk. Exp. Biol. Med. 234:562–565.
- Stelwagen K, van Espen DC, Verkerk GA, McFadden HA, Farr VC. Elevated plasma cortisol reduces permeability of mammary tight junctions in the lactating bovine mammary epithelium. J. Endocrinol. 159:173–178.
- Stelwagen K, McFadden HA, Demmer J. Prolactin, alone or in combination with glucocorticoids, enhances tight junction formation and expression of the tight junction protein occludin in mammary cells. Mol. Cell. Endocrinol. 156:55–61.
- Stirrat LI, Sengers BG, Norman JE, Homer NZM, Andrew R, Lewis RM, Reynolds RM. Transfer and metabolism of cortisol by the isolated perfused human placenta. J. Clin. Endocrinol. Metab. 103:640–648.
- Sud SC, Tucker HA, Meites J. Estrogen-progesterone requirements for udder development in ovariectomized heifers. J. Dairy Sci. 51:210–214.
- Sun K, Adamson SL, Yang K, Challis JRG. Interconversion of cortisol and cortisone by 11β-hydroxysteroid dehydrogenases type 1 and 2 in the perfused human placenta. Placenta 20:13–19.
- Taraborrelli S. Physiology, production and action of progesterone. Acta Obstet. Gynecol. Scand. 94:8–16.
- Tashima LS, Mazoujian G, Bryant-Greenwood GD. Human relaxins in normal, benign and neoplastic breast tissue. J. Mol. Endocrinol. 12:351–364.
- Thompson GE. Cortisol and regulation of tight junctions in the mammary gland of the late-pregnant goat. J. Dairy Res. 63:305–308.
- Tsutsui T, Suzuki Y, Toyonaga M, Oba H, Mizutani T, Hori T. The role of the ovary for the maintenance of pregnancy in cats. Reprod. Domest. Anim. 44:120–124.
- Tucker HA. Physiological control of mammary growth, lactogenesis, and lactation. J. Dairy Sci. 64:1403–1421.
- Tucker HA. Symposium: hormonal regulation of milk synthesis. Hormones, mammary growth, and lactation: a 41-year perspective. J. Dairy Sci. 83:874–884.
- Vannuccini S, Bocchi C, Severi FM, Challis JR, Petraglia F. Endocrinology of human parturition. Ann. Endocrinol. 77:105–113.
- Verstegen-Onclin K, Verstegen J. Endocrinology of pregnancy in the dog: a review. Theriogenology 70:291–299.
- Wellnitz O, Bruckmaier RM. Invited review: the role of the blood–milk barrier and its manipulation for the efficacy of the mammary immune response and milk production. J. Dairy Sci. 104:6376–6388.
- Wildman DE, Chen C, Erez O, Grossman LI, Goodman M, Stark A, Romero R. Evolution of the mammalian placenta revealed by phylogenetic analysis. Proc. Natl. Acad. Sci. U S A. 103(9):3203–3208.
- Wilkinson TN, Speed TP, Tregear GW, Stark A, Bathgate RA. Evolution of the relaxin-like peptide family. BMC Evol Biol. 5:14.
- Willcox DL, Arthur PG, Hartmann PE, Whitely JL. Perinatal changes in plasma oestradiol-17b, cortisol and progesterone and the initiation of lactation in sows. Aust. J. Biol. Sci. 36:173–181.
- Willett LB, Smith KL, Schanbacher FL. Hormone induced lactation in the bovine III. Dynamics of injected and endogenous hormones. J. Dairy Sci. 59:504–514.
- Winn RJ, Baker MD, Merle CA, Sherwood OD. Individual and combined effects of relaxin, estrogen, and progesterone in ovariectomized gilts. II. Effects on mammary development. Endocrinology 135:1250–1255.
- Woodward TL, Beal WE, Akers RM. Cell interactions in initiation of mammary epithelial proliferation by oestradiol and progesterone in prepubertal heifers. J. Endocrinol. 136:149–157.
- Yoshida T, Suzuki H, Hattori Y, Noda K. Hormonal changes around the parturition in rats. Tohoku J. Exp. Med. 135:87–91.
- Zakar T, Hertelendy F. Progesterone withdrawal: key to parturition. Am. J. Obstet. Gynecol. 196:289–296.
- Zhao L, Roche PJ, Gunnersen JM, Hammond VE, Tregear GW, Wintour EM, Beck F. Mice without a functional relaxin gene are unable to deliver milk to their pups. Endocrinology 140:445–453.
- Ziecik AJ, Przygrodzka E, Jalali BM, Kaczmarek MM. Regulation of the porcine corpus luteum during pregnancy. Reproduction 156:R57–R67.