Embryonic signals and placental hormones: molecular bases and potential use for the diagnosis and followup of gestation in farm animals

Keywords: embryo, fetoplacental unit, pregnancyassociated glycoproteins


During early pregnancy, the endocrine function of the embryo and foeto-maternal unit (endometrium and trophectoderm) involves the synthesis of a series of growth factors (e.g. Insuline-like Growth Factors, Epidermal Growth Factor, Transforming Growth Factors, Platelet-Derived Growth Factor and Fibroblast Growth Factors) and several steroid (e.g. progesterone, estrogens and prostaglandins) and proteins (e.g. Early Pregnancy Factor, Interferon tau and Pregnancy-Associated Glycoproteins) hormones. Some of these molecules interfere with the implantation of the embryo and with the establishment of pregnancy and corpus luteum maintenance during gestation. Indeed, pregnancy diagnosis is an important part in reproduction management of ruminants. The pregnancy associated glycoproteins (PAG) are synthetized in the rumiant by the mono- and binucleates cells of the trophectoderm. Part of them are released into maternal blood circulation where they can be assayed by RIA and ELISA techniques, as pregnancy diagnosis biomarker. The assay of PAG can also bring very interesting information for researchers working in programs focused on the study of embrionic and fetal moralities, as well as on embryo technology (FIV, clonage) animal nutrition, or infections diseases resulting in pathologies affecting the pregnancy.


Download data is not yet available.


Abd-Elnaeim MM, Derar IR, Wilsher S, Allen WR, Leiser R, Schuler G. 2009. Immunohistochemical localization of oestrogen receptors alpha and beta, progesterone receptor and aromatase in the equine placenta. Reprod Domest Anim. 44:312-319. (doi: 10.1111/j.1439-0531.2008.01073.x)

Akizawa, H.; Nagatomo, H.; Odagiri, H.; Kohri, N.; Yamauchi, N.; Yanagawa, Y.; Nagano, M.; Takahashi, M.; Kawahara, M. 2016. Conserved Roles of Fibroblast Growth Factor Receptor 2 Signaling in the Regulation of Inner Cell Mass Development in Bovine Blastocysts. Mol. Reprod. Dev. 83:516–525. (doi: 10.1002/mrd.22646)

Akpan, I.; Pro, B.; Platanias, L.C. 2018. Transforming Growth Factor Superfamily Ligands and Links to Tumorigenesis. Leuk Lymphoma. 59:1282–1283. (doi: 10.1080/10428194.2018.1461865)

Ali A, Swanepoel CM, Winger QA, Rozance PJ, Anthony RV. 2020. Chorionic somatomammotropin RNA interference alters fetal liver glucose utilization. J Endocrinol. 247:251-262. (doi: 10.1530/JOE-20-0375).

Alvarez-Oxiley A.V., Sousa N.M., Beckers J.F. 2008. Native and Recombinant bovine Placental Lactogen. Reproductive Biology. 8:85-106. (doi:10.1016/s1642-431x(12)60006-0)

Alvarez-Oxiley A.V., Sousa N.M., Hornick J.L., Touati K., van Der Weijden G.C., Taverne M.A.M., Szenci O., Sulon J., Debliquy P., Beckers J.F. 2007. Radioimmunoassay of bovine placental lactogen (bPL) using recombinant and native preparations: determination of fetal concentrations across gestation. Reproduction Fertility and Development. 19:1-7. (doi: 10.1071/rd06173)

Alvarez-Oxiley A.V., Sousa N.M., Hornick J.L., Touati K., van Der Weijden G.C., Taverne M.A.M., Szenci O., Beckers J.F. 2010. Effect of exogenous circulating anti-bPL antibodies on bovine placental lactogen measurements in fetal samples. Acta Veterinaria Scandinavica. 52:9. (doi: 10.1186/1751-0147-52-9)

Alvarez-Oxiley A.V., Sousa N.M., Hornick J.L., Touati K., van Der Weijden G.C., Taverne M.A.M., Szenci O., Sulon J., Debliquy P., Beckers J.F. 2007. Radioimmunoassay of bovine placental lactogen (bPL) using recombinant and native preparations: determination of fetal concentrations across gestation. Reproduction Fertility and Development. 19:1-7. (doi: 10.1071/rd06173)

Anchordoquy, J.M., Anchordoquy, J.P., Testa, J.A., Sirini, M.Á., Furnus, C.C. 2015. Influence of Vascular Endothelial Growth Factor and Cysteamine on in Vitro Bovine Oocyte Maturation and Subsequent Embryo Development. Cell Biol. Int. 39:1090–1098. (doi: 10.1002/cbin.10481)

Anthony R.V., Pratt S.L., Liang R., Holland M.D. 1995. Placental–fetal hormonal interactions: impact on fetal growth. J Anim Sci. 73:1861–1871. (doi: 10.2527/1995.7361861x)

Antoniazzi AQ, Webb BT, Romero JJ, Ashley RL, Smirnova NP, Henkes LE, Bott RC, Oliveira JF, Niswender GD, Bazer FW, Hansen TR. 2013. Endocrine delivery of interferon t protects the corpus luteum from prostaglandin F2a-induced luteolysis in ewes. Biology of Reproduction, 88:1–12. (doi: 10.1095/biolreprod.112.105684)

Arck P, Hansen PJ, Mulac Jericevic B, Piccinni MP, Szekeres-Bartho J. 2007. Progesterone during pregnancy: endocrine-immune cross talk in mammalian species and the role of stress. Am J Reprod Immunol, 58:268-279. (doi: 10.1111/j.1600-0897.2007.00512.x)

Arima, Y., and Bremel, R. D. 1983. Purification and characterization of bovine placental lactogen. Endocrinology. 113:2186–2194. (doi: 10.1210/endo-113-6-2186)

Athanasas-Platsis S, Quinn KA, Wong TY, Rolfe BE, Cavanagh AC, Morton H. 1989. Passive immunization of pregnant mice against early pregnancy factor causes loss of embryonic viability. J Reprod Fertil. 87:495-502. (doi: 10.1530/jrf.0.0870495)

Athanasas-Platsis, S., Zhang, B., Hillyard, N.C., Cavanagh, A.C., Csurhes, P.A., Morton, H. & Mccombe, P.A. 2003. Early pregnancy factor suppresses the infiltration of lymphocytes and macrophages in the spinal cord of rats during experimental autoimmune encephalomyelitis but has no effect on apoptosis. J. Neurol. Sci. 214:27-36. (doi: 10.1016/s0022-510x(03)00170-9)

Ayad A., Sousa N.M., Sulon J., Iguer-Ouada M., Beckers J.F. 2007a. Comparison of five radioimmunoassay systems for PAG measurement: ability to detect early pregnancy in cows. Reprod. Dom. Anim. 42:433-40. (doi: 10.1111/ j.1439-0531.2006.00804.x)

Ayad A., Sousa N.M., Sulon J., Hornick J.L., Watts J., Lopez-Gatius F., Iguerouada M., Beckers J.F. 2007b. Influence of progesterone concentrations on trophoblast and pituitary secretory functions during the first trimester of pregnancy in dairy cattle. Theriogenology. 67:1503-1511. (doi: 10.1016/j.theriogenology.2007.03.010)

Baker CM, Goetzmann LN, Cantlon JD, Jeckel KM, Winger QA, Anthony RV. 2016. Development of ovine chorionic somatomammotropin hormone-deficient pregnancies. Am J Physiol Regul Integr Comp Physiol. 310:R837-46. (doi: 10.1152/ajpregu.00311.2015).

Barbato O., Sousa N.M., Klisch K., Clerget E., Debenedetti A., Barile V.L., Malfatti A., Beckers J.F. 2008. Isolation of new pregnancy-associated glycoproteins from water buffalo (Bubalus bubalis) placenta by Vicia villosa affinity chromatography. Research in Veterinary Science. 85:457-466. (doi:10.1016/j.rvsc.2008.01.004)

Bazer FW, Thatcher WW. 2017. Chronicling the discovery of interferon tau. Reproduction. 154:F11-F20. (doi: 10.1530/REP-17-0257)

Beckers J.F. – 1983. L'hormone placentaire somatomammotrope bovine, Université de Liège, (Thèse d'agrégation), 207 p.

Beckers J.F., Fromont-Lienard C., van der Zwalmen P., Wouters Ballman P., Ectors. F. 1980. Isolement d'une hormone placentaire bovine présentant une activité analogue à la prolactine et à l'hormone de croissance. Ann. Med. Vet. 124:585-604.

Ben-Jonathan N., Mershon J.L., Allen D.L., Steinmetz R.W. 1996. Extrapituitary prolactin: distribution, regulation, functions, and clinical aspects. Endocr. Rev. 17:639-669. (doi: 10.1210/edrv-17-6-639)

Bériot M., Tchimbou A.F., Barbato O., Beckers J.F., de Sousa N.M. 2014. Identification of pregnancy-associated glycoproteins and a-fetoprotein in fallow deer (Dama dama) placenta. Acta Veterinaria Scandinavica. 56:4. (doi:10.1186/1751-0147-56-4)

Bole-Feysot C., Goffin V., Edery M., Binart N., Kelly P.A. 1998. Prolactin (PRL) and its receptor: actions, signal transduction pathways and phenotypes observed in PRL receptor knockout mice. Endocr. Rev. 19:225-268. (doi: 10.1210/edrv.19.3.0334)

Bott RC, Ashley RL, Henkes LE, Antoniazzi AQ, Bruemmer JE, Niswender GD, Bazer FW, Spencer TE, Smirnova NP, Anthony RV, Hansen TR. 2010. Uterine vein infusion of interferon t (IFNT) extends luteal life span in ewes. Biology of Reproduction. 82:725–735. (doi: 10.1095/biolreprod.109.079467)

Böttcher, R.T.; Niehrs, C. 2005. Fibroblast Growth Factor Signaling during Early Vertebrate Development. Endocr. Rev. 26: 63–77. (doi: 10.1210/er.2003-0040)

Brandt G.A., Parks T.E., Killian G., Ealy A.D, Green J.A. 2007. A cloning and expression analysis of pregnancy-associated glycoproteins expressed in trophoblasts of the white-tail deer placenta. Molecular Reproduction and Development. 74:1355–1362. (doi:10.1002/mrd.20669)

Breukelman S.P., Perenyi Z., Taverne M.A., Jonker H., van der Weijden G.C., Vos P.L., de Ruigh L., Dieleman S.J., Beckers J.F., Szenci O. 2012. Characterisation of pregnancy losses after embryo transfer by measuring plasma progesterone and bovine pregnancy-associated glycoprotein-1 concentrations. Veterinary Journal. 194:71–76. (doi:10.1016/j.tvjl.2012. 02.020)

Bukovsky A, Cekanova M, Caudle MR, Wimalasena J, Foster JS, Henley DC, Elder RF. 2003b. Expression and localization of estrogen receptor-alpha protein in normal and abnormal term placentae and stimulation of trophoblast differentiation by estradiol. Reprod Biol Endocrinol. 1:13. (doi: 10.1186/1477-7827-1-13)

Burton Graham J. and Fowden Abigail L. 2015. The placenta: a multifaceted, transient organPhil. Trans. R. Soc. B370: 20140066 (doi.org/10.1098/rstb.2014.0066)

Butler J.E., Hamilton W.C., Sasser R.G., Ruder C.A., Hass G.M., Williams R.J. 1982. Detection and partial characterization of two bovine pregnancyspecific proteins. Biology of Reproduction. 26:925–933. (doi:10.1095/ biolreprod26.5.925)

Buttle H.L., Forsyth I.A. 1976. Placental lactogen in the cow. J. Endocrinol. 68:141-148. (doi: 10.1677/joe.0.0680141)

Camacho-Arroyo I, Montor JM. 2012. Beyond reproductive effects of sex steroids. Mini Rev Med Chem. 12:1037-1039. (doi: 10.2174/138955712802762257)

Carter Anthony M. 2019. Evolution of placentation in cattle and antelopes. Anim. Reprod., v.16, n.1, p.3-17, Jan./Mar. (doi: 10.21451/1984-3143-AR2018-0145)

Cavanagh AC. 1996. Identification of early pregnancy factor as chaperonin 10: implications for understanding its role. Rev Reprod. 1:28-32. (doi: 10.1530/ror.0.0010028)

Chai N., Patel Y., Jacobson K., McMahon J., McMahon A., Rappolee D.A. 1998. FGF is an essential regulator of the fifth cell division in preimplantation mouse embryos. Dev. Biol. 198:105-115. (doi: 10.1006/dbio.1997.8858)

Chan J.S., Robertson H.A., Friesen H.G. 1978a. Distribution of binding sites for ovine placental lactogen in the sheep. Endocrinology. 102:632-40. (doi: 10.1210/endo-102-2-632).

Chan J.S., Robertson H.A., Friesen H.G. 1978b. Maternal and fetal concentrations of ovine placental lactogen measured by radioimmunoassay. Endocrinology. 102:1606-13. (doi: 10.1210/endo-102-5-1606).

Charlier M., Hue D., Boisnard M., Martal J., Gaye P. 1991. Cloning and structural analysis of two distinct families of ovine interferon-alpha genes encoding functional class II and trophoblast (OTP) alpha-interferons. Mol. Cell Endocrinol. 76:161-171. (doi: 10.1016/0303-7207(91)90270-3)

Charpigny G, Reinaud P, Huet JC, Guillomot M, Charlier M, Pernollet JC, Martal J. 1988. High homology between a trophoblastic protein (trophoblastin) isolated from ovine embryo and alpha-interferons. FEBS Lett. 228:12-6. (doi:10.1016/0014-5793(88)80574-x)

Chen Quangang, Xiaorong Zhu, Renjin Chen, Jing Liu, Peng Liu, Ankang Hu, Lianlian WU, Hui Hua, Honghua Yuan. 2016. Early Pregnancy Factor Enhances the Generation and Function of CD4+CD25+ Regulatory T Cells. The Tohoku Journal of Experimental Medicine. Volume 240 Issue 3 Pages 215-220. (doi.org/10.1620/tjem.240.215)

Chene N., Martal J. – 1996. Contrôle du développement embryonnaire et reconnaissance maternelle de la gestation. Point Vét. 28:901-910.

Citri, A.; Yarden, Y. 2006. EGF-ERBB Signalling: Towards the Systems Level. Nat. Rev. Mol. Cell Biol. 7:505–516. (doi: 10.1038/nrm1962)

Clemmons, D.R.; Maile, L.A. 2005. Interaction between Insulin-like Growth Factor-I Receptor and AlphaVbeta3 Integrin Linked Signaling Pathways: Cellular Responses to Changes in Multiple Signaling Inputs. Mol. Endocrinol. 19:1–11. (doi: 10.1210/me.2004-0376)

Clerget E., Sousa N.M., Bella A., Maghuin-RogisterbJ G., Becker J.F. 2008. Placental glycoproteins in the mammals. Annales d'Endocrinologie. 69:18-29. (doi: 10.1016/j.ando.2007.10.003)

Cooke, F.N.T.; Pennington, K.A.; Yang, Q.; Ealy, A.D. 2009. Several Fibroblast Growth Factors Are Expressed during Pre-Attachment Bovine Conceptus Development and Regulate Interferon-Tau Expression from Trophectoderm. Reproduction. 137: 259–269. (doi: 10.1530/REP-08-0396)

Cordoba M.C., Sartori R., Fricke P.M. 2001. Assessment of a commercially available early conception factor (ECF) test for determining pregnancy status of dairy cattle. Journal of Dairy Science. 84:1884-1889. (doi:10.3168/jds.S0022-0302(01)74629-2)

Coskun S., Sanbuissho A., Lin Y.C., Rikihisa Y. 1991. Fertilizability and subsequent developmental ability of bovine oocytes matured in medium contai ning epidermal growth factor (EGF). Theriogenology. 36:485-494. (doi: 10.1016/0093-691x(91)90477-u)

Currie W.B., Card C.E., Michel F.J., Ignotz G. 1990. Purification, partial characterization, and development of a specific radioimmunoassay for goat placental lactogen. Journal of Reproduction and Fertility. 90:25-36. (doi: 10.1530/jrf.0.0900025)

Da Rosa, P.R.A., De Cesaro, M.P., Pereira Dau, A.M., Duggavathi, R., Bordignon, V., Gonçalves, P.B.D. 2017. Reversible Meiotic Arrest of Bovine Oocytes by EGFR Inhibition and Follicular Hemisections. Theriogenology. 99:53–62. (doi: 10.1016/j.theriogenology.2017.05.014)

Dai G., Lu L., Tang S., Peal M.J., Soares M.J. 2002. Prolactin family miniarray: a tool for evaluating uteroplacental-trophoblast endocrine cell phenotypes. Reproduction. 124:755-765. (doi: 10.1530/rep.0.1240755)

De la Llosa-Hermier MP, Leboulleux P, Chene N, Martal J. 1983. Inhibitory effect of ovine and human placental lactogens on progesterone catabolism in luteinized rat ovaries in vitro. Placenta. 4:479-87.

Denley A, Cosgrove LJ, Booker GW, Wallace JC & Forbes BE 2005. Molecular interactions of the IGF system. Cytokine and Growth Factor Reviews 16:421–439. (doi.org/10.1016/j.cytogfr.2005.04.004)

Dichmann, D.S.; Miller, C.P.; Jensen, J.; Scott Heller, R.; Serup, P. 2003. Expression and Misexpression of Members of the FGF and TGFbeta Families of Growth Factors in the Developing Mouse Pancreas. Dev. Dyn. 226:663–674. (doi: 10.1002/dvdy.10270)

Diczfalusy E. 1964. Endocrine functions of the human fetoplacental unit. Fed. Proc. 23:791-798.

Diógenes, M.N.; Guimarães, A.L.S.; Leme, L.O.; Dode, M.A.N. 2017. Bovine in Vitro Embryo Production: The Effects of Fibroblast Growth Factor 10 (FGF10). J. Assist. Reprod. Genet. 34:383–390. (doi: 10.1007/s10815-016-0852-8)

Do H.J., Kim J.H., Abeydeera L.R., Han Y.M., Matteri R.L., Green J.A., Roberts R.M., Day B.N., Prather R.S. 2001. Expression of pregnancy-associated glycoprotein 1 and 2 genes in in vivo, in vitro and parthenogene tically derived preimplantation pig embryos. Zygote. 9:245-250. (doi: 10.1017/s0967199401001265)

Ealy A.D. y Wooldridge L.K. 2017. The evolution of interferon-tau. Reproduction. 154:F1-F10. (doi: 10.1530/REP-17-0292)

El Amiri B, Sousa NM, Alvarez Oxiley A, Hadarbach D, Beckers JF. 2015. Pregnancy-associated glycoprotein (PAG) concentration in plasma and milk samples for early pregnancy diagnosis in Lacaune dairy sheep. Res Vet Sci. 99:30-6. (doi: 10.1016/j.rvsc.2014.12.016)

Eley RM, Thatcher WW, Bazer FW. 1979. Hormonal and physical changes associated with bovine conceptus development. J Reprod Fertil. 55:181-190. (doi: 10.1530/jrf.0.0550181)

Estrada J.L., Jones E.E., Johnson B.H., Petters R.M. 1991. Effect of insulin-like growth factor-I on protein synthesis in porcine embryonic discs cultured in vitro. J. Reprod. Fertil. 93:53-61. (doi: 10.1530/jrf.0.0930053)

Ferrara, N., Chen, H., Davis-Smyth, T., Gerber, H.P., Nguyen, T.N., Peers, D., Chisholm, V., Hillan, K.J., Schwall, R.H. 1998. Vascular Endothelial Growth Factor Is Essential for Corpus Luteum Angiogenesis. Nat. Med. 4:336–340. (doi: 10.1038/nm0398-336)

Fillion C., Chaouat G., Reinaud P., Charpigny J.C., Martal J. 1991. Immunoregulatory effects of ovine trophoblastin protein (oTP): all five isoforms suppress PHA-induced lymphocyte proliferation.J Reprod Immunol. 19:237-49. (doi: 10.1016/0165-0378(91)90038-r)

Forsyth I.A . 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. (doi: 10.3168/jds.S0022-0302(86)80479-9)

Garbayo J.M., Benoit R., Alabart J.L, Folch J., Wattiez R., Falmagne P., Beckers J.F. 1998. Isolation and Partial Characterization of a Pregnancy-Associated Glycoprotein Family from the Goat Placenta. Biology of Reproduction. 58:109–115. (doi.org/10.1095/biolreprod58.1.109)

Garbayo J.M., Green J.A., Mannekin M., Beckers J.F., Kiesling D.O., Ealy A.D., Roberts R.M. 2000. Caprine pregnancy-associated glycoproteins (PAG): their cloning, expression and evolutionary relationship to other PAG. Molecular Reproduction and Development. 57:311-322. (doi:10.1002/ 1098-2795(200012)57:4!311::AID-MRD2O3.0.CO;2-F)

Germain-Lee, E.L.; Janicot, M.; Lammers, R.; Ullrich, A.; Casella, S.J. 1992. Expression of a Type I Insulin-like Growth Factor Receptor with Low Affinity for Insulin-like Growth Factor II. Biochem. J. 281 Pt 2:413–417. (doi: 10.1042/bj2810413)

Ghaffari Laleh, V., Ghaffari Laleh, R., Pirany, N. & Moghadaszadeh Ahrabi, M. 2008. Measurement of EPF for detection of cow pregnancy using rosette inhibition test. Theriogenology. 70:105-107. (doi.org/10.1016/j.theriogenology.2008.02.003)

Gifford C.A., Racicot K., Clark D.S., Austin K.J., Hansen T.R., Lucy M.C., Davies C.J., Ott T.L. 2007. Regulation of Interferon-stimulated genes in peripheral blood leukocytes in pregnant and bred, nonpregnant dairy cows. Journal of Dairy Science. 90:274–280. (doi:10.3168/jds.S0022-0302 (07)72628-0)

Giroto, A.B.; Fontes, P.K.; Franchi, F.F.; Dos Santos, P.H.; Razza, E.M.; Nogueira, M.F.G.; Maioli, M.A.; Nogueira, G.P.; Nunes, G.B.; Mingoti, G.Z. 2019. Use of Pregnancy-Associated Plasma Protein-A during Oocyte in Vitro Maturation Increases IGF-1 and Affects the Transcriptional Profile of Cumulus Cells and Embryos from Nelore Cows. Mol. Reprod. Dev. 86:1694–1704. (doi: 10.1002/mrd.23259)

Godkin J.D., Smith S.E., Johnson R.D., Doré J.J. 1997. The role of trophoblast interferons in the maintenance of early pregnancy in ruminants. Am. J. Reprod. Immunol. 37:137-143. (doi: 10.1111/j.1600-0897.1997.tb00202.x)

Gogolin-Ewens K.J., Lee C.S., Mercer W.R., Moseby A.M., Brandon M.R. 1986. Characterization of a sheep trophoblast-derived antigen first appearing at implantation. Placenta. 7:243-255. (doi.org/10.1016/S0143-4004(86)80162-X)

Gootwine E., Sise J.A., Penty J.M., Montgomery G.W. 1993. The duplicated gene copy of the ovine growth hormone gene contains a PvuII polymorphism in the second intron. Anim. Genet. 24:319-321. (doi: 10.1111/j.1365-2052.1993.tb00321.x)

Green J.A., Parks T.E., Avalle M.P., Telugu B.P., McLain A.L., Peterson A.J., McMillan W., Mathialagan N., Xie S., Hook R.R. 2005. The establishment of an ELISA for the detection of pregnancy-associated glycoproteins (PAGs) in the serum of pregnant cows and heifers. Theriogenology. 63:1481–1503. (doi:10.1016/j.theriogenology.2004.07. 011)

Green J.A., Xie S., Quan X., Bao B., Gan X., Mathialagan N., Beckers J.F., Roberts R.M. 2000. Pregnancy-associated bovine and ovine glycoproteins exhibit spatially and temporally distinct expression patterns during pregnancy. Biology of Reproduction. 62:1624–1631. (doi:10.1095/ biolreprod62.6.1624)

Green J.C., Okamura C.S., Poock S.E., Lucy M.C. 2010. Measurement of interferon-tau (IFN-t) stimulated gene expression in blood leukocytes for pregnancy diagnosis within 18–20d after insemination in dairy cattle. Animal Reproduction Science. 121:24–33. (doi:10.1016/j.anireprosci. 2010.05.010).

Grosso Mc, Rv Bellingeri, Ce Motta, Fe Alustiza, Ny Picco & Ab Vivas. 2015. Immunohistochemical distribution of early pregnancy factor in ovary, oviduct and placenta of pregnant gilts, Biotechnic & Histochemistry, 90:14-24, (doi: 10.3109/10520295.2014.931599)

Guillomot M., Reinaud P., Bonnardiere C., Charpigny G. 1998. Characterization of conceptus-produced goat interferon tau and analysis of its temporal and cellular distribution during early pregnancy. J. Reprod. Fertil. 112:149-156. (doi: 10.1530/jrf.0.1120149)

Guillomot M. L’implantation du blastocyste. 2001. In: Thibault C., Levasseur M.C. (Eds), La reproduction chez les mammifères et l’homme. 2e Ed. Ellipses: Paris, pp. 457-478.

Hajian, M.; Hosseini, S.M.; Ostadhosseini, S.; Nasr-Esfahani, M.H. 2016. Targeting the Transforming Growth Factor-_ Signaling during Pre-Implantation Development in Embryos of Cattle, Sheep and Goats. Growth Factors, 34:141–148. (doi: 10.1080/08977194.2016.1206089)

Handwerger s. 1991. Clinical counterpoint: the physiology of placental lactogen in human pregnancy. endocr. rev. 12:329-336. (doi: 10.1210/edrv-12-4-329)

Hansen PJ. 1998. Regulation of uterine immune function by progesterone--lessons from the sheep. J Reprod Immunol, 40:63-79. (doi: 10.1016/s0165-0378(98)00035-7)

Hansen TR, Austin KJ, Perry DJ, Pru JK, Teixeira MG, Johnson GA. 1999. Mechanism of action of interferon-tau in the uterus during early pregnancy. J Reprod Fertil. 54:329-39.

Harris LK y Westwood M. 2012. Biology and significance of signalling pathways activated by IGF-II. Growth Factors 30:1–12. (doi.org/10.3109/08977194.2011.640325)

Hayden T.J., Thomas C.R., Forsyth I.A. 1979. Effect of number of young born (litter size) on milk yield of goats: Role for placental lactogen. J. Dairy Sci. 62:53-57. (doi: 10.3168/jds.S0022-0302(79)83201-4)

Helmer S.D., Hansen P.J., Anthony R.V., Thatcher W.W., Bazer F.W., Roberts R.M. 1987. Identification of bovine trophoblast protein-1, a secretory protein immunologically related to ovine trophoblast protein-1. J. Reprod. Fertil. 79:83-91. (doi: 10.1530/jrf.0.0790083)

Hillege, M.M.G.; Galli Caro, R.A.; Offringa, C.; de Wit, G.M.J.; Jaspers, R.T.; Hoogaars, W.M.H. 2020. TGF-_ Regulates Collagen Type I Expression in Myoblasts and Myotubes via Transient Ctgf and Fgf-2 Expression. Cells. 9:375. (doi: 10.3390/cells9020375)

Hughes A.L., Green J.A., Garbayo J.M., Roberts R.M. 2000. Adaptive diversification within a large family of recently duplicated, placentally-expressed genes. PNAS. 97:3319–3323. (doi:10.1073/pnas. 97.7.3319)

Hunter, M.G., Robinson, R.S., Mann, G.E., Webb, R. 2004. Endocrine and Paracrine Control of Follicular Development and Ovulation Rate in Farm Species. Anim. Reprod. Sci. 82–83:461–477. (doi: 10.1016/j.anireprosci.2004.05.013)

Ipsa, E.; Cruzat, V.F.; Kagize, J.N.; Yovich, J.L.; Keane, K.N. 2019. Growth Hormone and Insulin-Like Growth Factor Action in Reproductive Tissues. Front. Endocrinol. 10:777. (doi: 10.3389/fendo.2019.00777)

Ishiwata H., Katsuma S., Kizaki K., Patel O.V., Nakano H., Takahashi T., Imai K., Hirasawa A., Shiojima S., Ikawa H., Suzuki Y., Tsujimoto G., Izaike Y., Todoroki J., Hashizume K. 2003. Characterization of gene expression profiles in early bovine pregnancy using a custom cDNA microarray. Mol. Reprod. Dev. 65:9-18. (doi: 10.1002/mrd.10292)

Jaeger, L.A., Spiegel, A.K., Ing, N.H., Johnson, G.A.; Bazer, F.W., Burghardt, R.C. 2005. Functional Effects of Transforming Growth Factor Beta on Adhesive Properties of Porcine Trophectoderm. Endocrinology. 146:3933–3942. (doi: 10.1210/en.2005-0090)

Jeckel KM, Boyarko AC, Bouma GJ, Winger QA, Anthony RV. 2018. Chorionic somatomammotropin impacts early fetal growth and placental gene expression. J Endocrinol. 237:301-310. (doi: 10.1530/JOE-18-0093).

Johnson GA, Bazer FW, Seo H. 2021.The Early Stages of Implantation and Placentation in the Pig. Adv Anat Embryol Cell Biol. 234:61-89. (doi: 10.1007/978-3-030-77360-1_5)

Kaur H, Muhlhausler BS, Roberts CT, Gatford KL. 2021. The growth hormone-insulin like growth factor axis in pregnancy. J Endocrinol. Sep 1:JOE-21-0087.R1. (doi: 10.1530/JOE-21-0087)

Kessler M.A., Schuler L.A. 1997. Purification and properties of placental prolactin-related protein-I. Placenta. 18:29-36. (doi: 10.1016/s0143-4004(97)90068-0)

Khan O.A., Jiang H., Subramaniam P.S., Johnson H.M., Dhib-Jalbut S.S. 1998. Immunomodulating functions of recombinant ovine interferon tau:potential for therapy in multiple sclerosis and autoimmune disorders. Mult. Scler. 4:63-69. (doi: 10.1177/135245859800400204)

Kliem A., Tetens F., Klonisch T., Grealy M., Fischer B. 1998. Epidermal growth factor receptor and ligands in elongating bovine blastocysts. Mol. Reprod. Dev. 51:402-412. (doi: 10.1002/(SICI)1098-2795(199812)51:4<402::AID-MRD7>3.0.CO;2-9)

Kolle S., Sinowatz F., Boie G., Lincoln D., Waters M.J. 1997. Differential expression of the growth hormone receptor and its transcript in bovine uterus and placenta. Mol. Cell. Endocrinol. 131:127-136. (doi: 10.1016/s0303-7207(97)00097-x)

Lacroix M.C., Devinoy E., Cassy S., Servely J.L., Vidaud M., Kann G. 1999. Expression of growth hormone and its receptor in the placental and feto-maternal environment during early pregnancy in sheep. Endocrinology. 140:5587-5597. (doi: 10.1210/endo.140.12.7210)

Lacroix M.C., Devinoy E., Servely J.L., Puissant C., Kann G. 1996. Expression of the growth hormone gene in ovine placenta: detection and cellular localization of the protein. Endocrinology. 137:4886-4892. (doi: 10.1210/endo.137.11.8895361)

Laron Z. 2001. Insulin-like growth factor 1 (IGF-1): a growth hormone. Molecular Pathology 54 311–316. (doi.org/10.1136/mp.54.5.311)

Larson JH, Kumar CG, Everts RE, Green CA, Everts-van der Wind A, Band MR, Lewin HA. 2006. Discovery of eight novel divergent homologs expressed in cattle placenta. Physiol Genomics, 25:405-413. (doi:10.1152/physiolgenomics.00307.2005)

Larson R.C., Ignotz G.G., Currie W.B. 1992. Transforming growth factor β and basic fibroblast growth factor synergistically promote early bovine embryo development during the fourth cell cycle. Mol Reprod Dev. 33:432-435. doi: 10.1002/mrd.1080330409

Le Roith D, Bondy C, Yakar S, Liu JL, Butler A. 2001. The somatomedin hypothesis: 2001. Endocr Rev. 22:53-74. (doi: 10.1210/edrv.22.1.0419)

Lin J., Toft D.J., Bengtson N.W., Linzer D.I. 2000. Placental prolactins and the physiology of pregnancy. Recent Prog. Horm. Res. 55:37-51. (doi: 10.1016/s0009-2509(99)00304-8)

Llobat L. 2021. Pluripotency and Growth Factors in Early Embryonic Development of Mammals: A Comparative Approach. Vet. Sci. 8:78. (doi.org/10.3390/vetsci8050078)

Llobat, L., Marco-Jiménez, F., Peñaranda, D.S., Thieme, R.; Navarrete, A., Vicente, J.S. 2012. MRNA Expression in Rabbit Blastocyst and Endometrial Tissue of Candidate Gene Involved in Gestational Losses. Reprod. Domest. Anim., 47:281–287. (doi: 10.1111/j.1439-0531.2011.01855.x)

Lonergan P., Gutiérrez-Adán A., Pintado B., Fair T., Ward F., Fuente J.D., Boland M. 2000. Relationship between time of first cleavage and the expression of IGF-I growth factor, its receptor, and two housekeeping genes in bovine two-cell embryos and blastocysts produced in vitro. Mol Reprod Dev. 57:146-52. (doi: 10.1002/1098-2795(200010)57:2<146::AID-MRD5>3.0.CO;2-2)

Lynch K.A., Alexander R.M., Sasser R.G. 1992.The cloning and expression of the bovine pregnancy specific protein B (bPSPB) gene. Biol. Reprod. 46 (Suppl. 1):73.

Majewska M., Panasiewicz G., Majewski M., Szafranska B. 2006. Localization of chorionic pregnancy-associated glycoprotein family in the pig. Reproductive Biology. 6:205–230.

Martal J, Lacroix MC, Loudes C, Saunier M, Wintenberger-Torrès S. 1979. Trophoblastin, an antiluteolytic protein present in early pregnancy in sheep. J Reprod Fertil. 56:63-73. (doi: 10.1530/jrf.0.0560063)

Martal J. y Djiane J. 1977. The production of chorionic somatomammotrophin in sheep. J Reprod Fertil. 49:285-289. (doi: 10.1530/jrf.0.0490285)

Matsui M., Takahashi Y., Hishinuma M., Kanagawa H. 1995. Insulin and insulin-like growth factor-1 (IGF-1) stimulate the development of bovine embryos fertili zed in vitro. J. Vet. Med. Sci. 57:1109-1111. (doi: 10.1292/jvms.57.1109)

Mialon M.M., Camous S., Renand G., Martal J., Menissier F. 1993. Peripheral concentrations of a 60-kDa pregnancy serum protein during gestation and after calving and in relationship to embryonic mortality in cattle. Reproduction, Nutrition, Development. 33:269–282. (doi:10.1051/ rnd:19930309).

Michael, D.D.; Alvarez, I.M.; Ocón, O.M.; Powell, A.M.; Talbot, N.C.; Johnson, S.E.; Ealy, A.D. 2006. Fibroblast Growth Factor-2 Is Expressed by the Bovine Uterus and Stimulates Interferon-Tau Production in Bovine Trophectoderm. Endocrinology, 147, 3571–3579. (doi: 10.1210/en.2006-0234)

Milosavljevic M, Duello TM, Schuler LA. 1989. In situ localization of two prolactin-related messenger ribonucleic acids to binucleate cells of bovine placentomes. Endocrinology, 125:883-889. (doi:10.1210/endo-125-2-883).

Mitchell BF, Taggart MJ. 2009. Are animal models relevant to key aspects of human parturition? Am J Physiol Regal Integr Comp Physiol, 297:R525-545. (doi: 10.1152/ajpregu.00153.2009)

Moor R.M. y Rowson L.E. 1966. Local uterine mechanisms affecting luteal function in the sheep. J. Reprod. Fertil. 11:307-310. (doi: 10.1530/jrf.0.0110307)

Mor, A.; Mondal, S.; Reddy, I.J.; Nandi, S.; Gupta, P. 2018. Molecular Cloning and Expression of FGF2 Gene in Pre-Implantation Developmental Stages of in Vitro-Produced Sheep Embryos. Reprod. Domest. Anim., 53:895–903. (doi: 10.1111/rda.13182)

Morton H, Cavanagh AC, Athanasas-Platsis S, Quinn KA, Rolfe BE. 1992. Early pregnancy factor has immunosuppressive and growth factor properties. Reprod Fertil Dev. 4:411-22. (doi: 10.1071/rd9920411)

Morton H., Hegh V., Clumie G.J.A. – 1974. Immunosuppression detected in pregnant mice by rosette inhibition test. Nature. 249:459-460. (doi: 10.1038/249459a0)

Morton, H., McKay, D.A., Murphy, R.M., Somodevilla-Torres, M.J., Swanson, C.E., Cassady, A.I., Summers, K.M. & Cavanagh, A.C. 2000. Production of a recombinant form of early pregnancy factor that can prolong allogeneic skin graft survival time in rats. Immunol. Cell Biol. 78:603-607. (doi: 10.1046/j.1440-1711.2000.00951.x)

Nayak N.R., Giudice L.C. 2003. Comparative biology of the IGF system in endometrium, decidua, and placenta, and clinical implications for fetal growth and implantation disorders. Placenta. 24:281-296. (doi: 10.1053/plac.2002.0906)

Nebel R.L. 1988. On-farm milk progesterone tests. Journal of Dairy Science. 71:1682-1690. (doi:10.3168/jds.S0022-0302(88)79733-7)

Neill JD. 2005. Knobil and Neill's physiology of reproduction. 3rd ed. New York: Academic Press

Nishimura, T.; Nakatake, Y.; Konishi, M.; Itoh, N. 2000. Identification of a Novel FGF, FGF-21, Preferentially Expressed in the Liver. Biochim Biophys Acta, 1492:203–206. (doi: 10.1016/s0167-4781(00)00067-1)

Noel S, Herman A, Johnson GA, Gray CA, Stewart MD, Bazer FW, Spencer TE. 2003. Ovine placental lactogen specifically binds to endometrial glands of the ovine uterus. Biol Reprod, 68:772-780. (doi: 10.1095/biolreprod.102.009183)

Ocón-Grove, O.M.; Cooke, F.N.T.; Alvarez, I.M.; Johnson, S.E.; Ott, T.L.; Ealy, A.D. 2008. Ovine Endometrial Expression of Fibroblast Growth Factor (FGF) 2 and Conceptus Expression of FGF Receptors during Early Pregnancy. Domest. Anim. Endocrinol. 34:135–145. (doi: 10.1016/j.domaniend.2006.12.002)

Oh SY, Kim CJ, Park I, Romero R, Sohn YK, Moon KC, Yoon BH. 2005. Progesterone receptor isoform (A/B) ratio of human fetal membranes increases during term parturition. Am J Obstet Gynecol, 193:1156-1160. (doi: 10.1016/j.ajog.2005.05.071)

Oliveira JF, Henkes LE, Ashley RL, Purcell SH, Smirnova NP, Veeramachaneni DN, Anthony RV, Hansen TR. 2008. Expression of interferon (IFN)-stimulated genes in extrauterine tissues during early pregnancy in sheep is the consequence of endocrine IFN-τau release from the uterine vein. Endocrinology. 149:1252–1259. (doi: 10.1210/en.2007-0863)

Pepe GJ, Albrecht ED. 1995. Actions of placental and fetal adrenal steroid hormones in primate pregnancy. Endocr Rev, 16:608-648. (doi: 10.1210/edrv-16-5-608)

Perry, G. A., Perkins, S. D., Northrop, E. J., Rich, J., Epperson, K. M., Andrews, T. N., Kline, A. C., Quail, L. K., Walker, J. A., Wright, C. L., & Russell, J. R. 2021. Impact of trace mineral source on beef replacement heifer growth, reproductive development, and biomarkers of maternal recognition of pregnancy and embryo survival. Journal of animal science, 99(7), skab160. (doi.org/10.1093/jas/skab160)

Petraglia F, Florio P, Nappi C, Genazzani AR. 1996. Peptide signaling in human placenta and membranes: autocrine, paracrine, and endocrine mechanisms. Endocr Rev.17:156-86. (doi: 10.1210/edrv-17-2-156)

Pohler K.G., Geary T.W., Johnson C.L., Atkins J.A., Jinks E.M., Busch D.C., Green J.A., MacNeil M.D., Smith M.F. 2013. Circulating bovine pregnancy associated glycoproteins are associated with late embryonic/fetal survival but not ovulatory follicle size in suckled beef cows. Journal of Animal Science. 91:4158-4167. (doi:10.2527/jas. 2013-6348)

Powell-Braxton L., Hollingshead P., Warburton C., Dowd M., Pitts-Meek S., Dalton D., Gillett N., Stewart T.A. 1993. IGF-1 is required for normal embryonic growth in mice. Genes Dev. 7:2609-2617. (doi: 10.1101/gad.7.12b.2609)

Rabinowich, H., Suminami, Y., Reichert, T.E., Crowley-Nowick, P., Bell, M., Edwards, R. & Whiteside, T.L. 1996. Expression of cytokine genes or proteins and signaling molecules in lymphocytes associated with human ovarian carcinoma. Int. J. Cancer. 68: 276-284. (doi: 10.1002/(SICI)1097-0215(19961104)68:3<276::AID-IJC2>3.0.CO;2-Z)

Raeside JI. 2017. A brief account of the discovery of the fetal/placental unit for estrogen production in equine and human pregnancies: relation to human medicine. Yale J Biol Med. 90:449-461.

Rappolee D.A., Basilico C., Patel Y., Werb Z. 1994. Expression and function of FGF-4 in peri-implantation development in mouse embryos. Development. 120:2259-2269. (doi: 10.1242/dev.120.8.2259)

Reddy S. y Watkins W.B. 1978. Purification and some properties of ovine placental lactogen. J Endocrinol. 78:59-69. (doi: 10.1677/joe.0.0780059).

Richani, D.; Gilchrist, R.B. 2018. The Epidermal Growth Factor Network: Role in Oocyte Growth, Maturation and Developmental Competence. Hum. Reprod. Update. 24:1–14. (doi: 10.1093/humupd/dmx029)

Rinderknecht, E., Humbel, R.E. 1978. The Amino Acid Sequence of Human Insulin-like Growth Factor I and Its Structural Homology with Proinsulin. J. Biol. Chem. 253:2769–2776.

Roberts RM, Ezashi T, Rosenfeld CS, Ealy AD, Kubisch HM. 2003. Evolution of the interferon tau genes and their promoters, and maternal-trophoblast interactions in control of their expression. Reprod Suppl, 61:239-251.

Roberts, C.T., Owens, J.A., Sferruzzi-Perri, A.N. 2008. Distinct Actions of Insulin-like Growth Factors (IGFs) on Placental Development and Fetal Growth: Lessons from Mice and Guinea Pigs. Placenta. 29:S42–S47. (doi: 10.1016/j.placenta.2007.12.002)

Rowson L.E. y Moor R.M. 1967. The influence of embryonic tissue homogenate infused into the uterus, on the life-span of the corpus luteum in the sheep. J Reprod Fertil. 13:511-516. (doi: 10.1530/jrf.0.0130511)

Saraiva MV, Rossetto R, Brito IR, Celestino JJ, Silva CM, Faustino LR, Almeida AP, Bruno JB, Magalhães DM, Matos MH, Campello CC, Figueiredo JR. 2010. Dynamic medium produces caprine embryo from preantral follicles grown in vitro. Reprod Sci. 17:1135-43. (doi: 10.1177/1933719110379269)

Sasser R.G., Ruder C.A., Ivani K.A., Butler J.E., Hamilton W.C. 1986. Detection of pregnancy by radioimmunoassay of a novel pregnancy-specific protein in serum of cows and a profile of serum concentrations during gestation. Biology of Reproduction. 35:936-942. (doi:10.1095/biolreprod35.4.936)

Schuler G, Greven H, Kowalewski MP, Döring B, Ozalp GR, Hoffmann B. 2008. Placental steroids in cattle: hormones, placental growth factors or by-products of trophoblast giant cell differentiation? Exp Clin Endocrinol Diabetes. 116:429-436. (doi: 10.1055/s-2008-1042408)

Schuler G, Teichmann U, Taubert A, Failing K, Hoffmann B. 2005. Estrogen receptor beta (ERbeta) is expressed differently from ERalpha in bovine placentomes. Exp Clin Endocrinol Diabetes. 113:107-114. (doi: 10.1055/s-2004-830539)

Schuler G, Wirth C, Klisch K, Pfarrer C, Leiser R, Hoffmann B. 1999. Immunolocalization of progesterone receptors in bovine placentomes throughout mid and late gestation and at parturition. Biol Reprod. 61:797-801. (doi: 10.1095/biolreprod61.3.797)

Schuler G., Fürbass R., Klisch K. 2018. Placental contribution to the endocrinology of gestation and parturitio. Proceedings of the 10th International Ruminant Reproduction Symposium (IRRS 2018); Foz do Iguaçu, PR, Brazil, September 16th to 20th, 2018.

Schuler L., Kessler M. 1992. Bovine placental prolactin-related hormones, Trends in Endocrinology & Metabolism, Volume 3, Issue 9, 334-338. doi.org/10.1016/1043-2760(92)90112-E.

Schultz GA, Heyner S. 1993. Growth factors in preimplantation mammalian embryos. Oxf Rev Reprod Biol. 15:43-81.

Senger, P. L. 2003. Pathways to pregnancy and parturition. 2º rev. ed. Washington, Current Conceptions. 381 p.

Shirasuna K, Matsumoto H, Matsuyama S, Kimura K, Bollwein H, Miyamoto. 2015. Possible role of interferon tau on the bovine corpus luteum and neutrophils during the early pregnancy vein. Reproduction. 150:217–225. (doi: 10.1530/REP-15-0085)

Sibiak, R.; Jankowski, M.; Gutaj, P.; Mozdziak, P.; Kempisty, B.; Wender-Ożegowska, E. 2020. Placental Lactogen as a Marker of Maternal Obesity, Diabetes, and Fetal Growth Abnormalities: Current Knowledge and Clinical Perspectives. J. Clin. Med. 9:1142. (doi.org/10.3390/jcm9041142)

Silva E., Sterry R.A., Kolb D., Mathialagan N., McGrath M.F., Ballam J.M., Fricke P.M. 2007. Accuracy of a pregnancy-associated glycoprotein ELISA to determine pregnancy status of lactating dairy cows twenty-seven days after timed artificial insemination. Journal of Dairy Science. 90:4612-4622. (doi:10.3168/jds.2007-0276)

Silva LA, Ginther OJ. 2010. Local effect of the conceptus in uterine vascular perfusion during early pregnancy in heifers. Reproduction. 139:453–63. (doi: 10.1530/REP-09-0363)

Soares MJ. 2004. The prolactin and growth hormone families: pregnancy-specific hormones/cytokines at the maternal-fetal interface. Reprod Biol Endocrinol 2:51. (doi: 10.1186/1477-7827-2-51)

Sousa N.M., El Amiri B., Remy B., Drión P.V., Beckers J.F. 2002. Signaux embryonnaires de la gestation. SFB – Paris, 29 - 31 Oct. Proceedings, p. 95.

Strauss J.F., Martinez F., Kiriakidou M. 1996. Placental steroid hormone synthesis: unique features and unanswered questions. Biol. Reprod. 54:303-311. (doi: 10.1095/biolreprod54.2.303)

Sutter, N.B.; Bustamante, C.D.; Chase, K.; Gray, M.M.; Zhao, K.; Zhu, L.; Padhukasahasram, B.; Karlins, E.; Davis, S.; Jones, P.G. 2007. A Single IGF1 Allele Is a Major Determinant of Small Size in Dogs. Science. 316:112–115. (doi: 10.1126/science.1137045)

Szafranska B., Xie S., Green J., Roberts R.M. 1995. Porcine pregnancyassociated glycoproteins: new members of the aspartic proteinase gene family expressed in trophectoderm. Biology of Reproduction. 53:21–28. (doi:10.1095/biolreprod53.1.21)

Szenci O. 2021. Recent Possibilities for the Diagnosis of Early Pregnancy and Embryonic Mortality in Dairy Cows. Animals (Basel). 11:1666. (doi: 10.3390/ani11061666)

Tanner AR, Lynch CS, Ali A, Winger QA, Rozance PJ, Anthony RV. 2021. Impact of chorionic somatomammotropin RNA interference on uterine blood flow and placental glucose uptake in the absence of intrauterine growth restriction. Am J Physiol Regul Integr Comp Physiol. 320:R138-R148. (doi: 10.1152/ajpregu.00223.2020).

Taverne, Marcel and Noakes David E. 2019. Pregnancy and Its Diagnosis, Editor(s): David E. Noakes, Timothy J. Parkinson, Gary C.W. England, Veterinary Reproduction and Obstetrics (Tenth Edition), W.B. Saunders, Pages 78-114, (doi.org/10.1016/B978-0-7020-7233-8.00005-7)

Telugu B.P., Walker A., Green J. 2009. Characterization of the bovine pregnancy-associated glycoprotein gene family – analysis of gene sequences, regulatory regions within the promoter and expression of selected genes. BMC Genomics. 10:185. (doi:10.1186/1471-2164- 10-185)

Thibault, Charles y Levasseur, Marie-Claire. 2001. La reproduction chez les mammifères et l'homme. Paris: INRA: Ellipses; 928 pp.

Ushizawa K, Takahashi T, Hosoe M, Kaneyama K, Hashizume K. 2005. Cloning and expression of two new prolactin-related proteins, prolactin-related protein-VIII and -IX, in bovine placenta. Reprod Biol Endocrinol. 3:68. (doi:10.1186/1477-7827-3-68).

Ushizawa K, Takahashi T, Hosoe M, Kizaki K, Abe Y, Sasada H, Hashizume K. 2007a. Gene expression profiles of novel caprine placental prolactin-related proteins similar to bovine placental prolactin-related proteins. BMC Dev Biol. 7:16. (doi:10.1186/1471-213X-7-16).

Ushizawa K, Takahashi T, Hosoe M, Ohkoshi K., Hashizume K. 2007b. Expression and characterization of novel ovine orthologs of bovine placental prolactin-related proteins. BMC Mol Biol. 8:95. (doi:10.1186/1471-2199-8-95).

Valdés, G., Erices, R., Chacón, C., Corthorn, J. 2008. Angiogenic, Hyperpermeability and Vasodilator Network in Utero-Placental Units along Pregnancy in the Guinea-Pig (Cavia Porcellus). Reprod. Biol. Endocrinol. 6:13. (doi: 10.1186/1477-7827-6-13)

Valdez Magaña, G.; Rodríguez, A.; Zhang, H.; Webb, R.; Alberio, R. 2014. Paracrine Effects of Embryo-Derived FGF4 and BMP4 during Pig Trophoblast Elongation. Dev. Biol. 387:15–27. (doi: 10.1016/j.ydbio.2014.01.008)

Veldhuis J.D., Anderson S.M., Shah N., Bray M., Vick T., Gentili A., Mulligan T., Johnson M.L., Weltman A., Evans W.S., Iranmanesh A. 2001. Neurophysiological regulation and target-tissue impact of the pulsatile mode of growth hormone secretion in the human. Growth Horm. IGF Res. 11:25-37. (DOI: 10.1016/s1096-6374(01)80005-8)

Wallace RM, Pohler KG, Smith MF, Green JA. 2015. Placental PAGs: gene origins, expression patterns, and use as markers of pregnancy. Reproduction. Mar;149:R115-26. (doi: 10.1530/REP-14-0485)

Wasielak, M.; Wi˛esak, T.; Bogacka, I.; Jalali, B.M.; Bogacki, M. 2017. Maternal Effect Gene Expression in Porcine Metaphase II Oocytes and Embryos in Vitro: Effect of Epidermal Growth Factor, Interleukin-1_ and Leukemia Inhibitory Factor. Zygote. 25: 120–130. (doi: 10.1017/S0967199416000332)

Watson A.J., Watson P.H., Arcellana-Panlilio M., Warnes D., Walker S.K., Schultz G.A., Armstrong D.T., Seamark R.F. 1994. A Growth Factor Phenotype Map for Ovine Preimplantation Development. Biology of Reproduction. 50:725–733. (doi: 10.1095/biolreprod50.4.725)

Wiedemann I, Krebs T, Momberg N, Knorr C, Tetens J. 2018. mRNA expression profiling in cotyledons reveals significant up-regulation of the two bovine pregnancy-associated glycoprotein genes boPAG-8 and boPAG-11 in early gestation. Vet Med Sci, 4:341-350. (doi:10.1002/vms3.123).

Wooding FB, Roberts RM, Green JA. 2005. Light and electron microscope immunocytochemical studies of the distribution of pregnancy associated glycoproteins (PAGs) throughout pregnancy in the cow: possible functional implications. Placenta, 26:807-827. (doi:10.1016/j.placenta.2004.10.014).

Xie S., Green J., Bixby J.B., Szafranska B., DeMartini J.C., Hecht S., Roberts R.M. 1997. The diversity and evolutionary relationships of the pregnancyassociated glycoproteins, an aspartic proteinase subfamily consisting of many trophoblast-expressed genes. PNAS. 94:12809–12816. (doi:10. 1073/pnas.94.24.12809)

Xie S.C., Low B.G., Nagel R.J., Kramer K.K., Anthony R.V., Zoli A.P., Beckers J.F., Roberts R.M. 1991. Identification of the major pregnancy-specific antigens of cattle and sheep as inactive members of the aspartic proteinase family. PNAS. 88:10247–10251. (doi:10.1073/pnas.88.22.10247)

Yamada O., Todoroki J., Kizaki K., Takahashi T., Imai K., Patel O.V., Schuler L.A., Hashizume K. 2002. Expression of prolactin-related protein I at the fetomaternal interface during the implantation period in cows. Reproduction. 124: 427-437.

Yancopoulos GD, Davis S, Gale NW, Rudge JS, Wiegand SJ, Holash J. 2000. Vascular-specific growth factors and blood vessel formation. Nature. 14:242-8. (doi: 10.1038/35025215)

Yang L, Wang XL, Wan PC, Zhang LY, Wu Y, Tang DW, Zeng SM. 2010. Up-regulation of expression of interferon-stimulated gene 15 in the bovine corpus luteum during early pregnancy. Journal of Dairy Science. 93:1000–1011. (doi: 10.3168/jds.2009-2529)

Yoshida Y, Miyamura M, Hamano S, Yoshida M. 1998. Expression of growth factor ligand and their receptor mRNAs in bovine ova during in vitro maturation and after fertilization in vitro. J Vet Med Sci. 60:549-54. doi: 10.1292/jvms.60.549

Złotkowska, A., Adamczyk, S., Andronowska, A. 2019. Presence of Trophoblast in the Uterine Lumen Affects VEGF-C Expression in Porcine Endometrium. Theriogenology, 125:216–223. (doi: 10.1016/j.theriogenology.2018.11.007)

Zoli A.P., Beckers J.F., Wouters-Ballman P., Closset J., Falmagne P., Ectors F. 1991. Purification and characterization of a bovine pregnancy-associated glycoprotein. Biology of Reproduction. 45:1-10. (doi:10.1095/biolreprod45.1.1)

Zoli A.P., Guilbault L.A., Delahaut P., Ortiz W.B., Beckers J.F. 1992. Radioimmunoassay of a bovine pregnancy-associated glycoprotein in serum: its application for pregnancy diagnosis. Biology of Reproduction. 46:83–92. (doi:10.1095/biolreprod46.1.83)

How to Cite
Alvarez-Oxiley, Andrea. 2022. “Embryonic Signals and Placental Hormones: Molecular Bases and Potential Use for the Diagnosis and Followup of Gestation in Farm Animals”. Archivos Latinoamericanos De Producción Animal 30 (3), 155-78. https://doi.org/10.53588/alpa.300302.
Invited papers