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      A Review of Mechanisms of Implantation

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          ABSTRACT

          Implantation is a highly organized process that involves an interaction between a receptive uterus and a competent blastocyst. In humans, natural fecundity suggests that the chance of conception per cycle is relatively low (~30%) and two-third of lost pregnancies occur because of implantation failure. Defective implantation leads to adverse pregnancy outcomes including infertility, spontaneous miscarriage, intrauterine fetal growth restriction and preeclampsia. With use of advanced scientific technologies, gene expression analysis and genetically-engineered animal models have revealed critical cellular networks and molecular pathways. But, because of ethical restrictions and the lack of a mechanistic experiment, comprehensive steps in human implantation have still not been completely understood. This review primarily focuses on the recent advances in mechanisms of implantation. Because infertility is an emerging issue these days, gaining an understanding the molecular and hormonal signaling pathway will improve the outcome of natural pregnancy and assisted reproductive technology.

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          Mice lacking progesterone receptor exhibit pleiotropic reproductive abnormalities.

          J Lydon, F DeMayo, C. R. Funk … (1995)
          Although progesterone has been recognized as essential for the establishment and maintenance of pregnancy, this steroid hormone has been recently implicated to have a functional role in a number of other reproductive events. The physiological effects of progesterone are mediated by the progesterone receptor (PR), a member of the nuclear receptor superfamily of transcription factors. In most cases the PR is induced by estrogen, implying that many of the in vivo effects attributed to progesterone could also be the result of concomitantly administered estrogen. Therefore, to clearly define those physiological events that are specifically attributable to progesterone in vivo, we have generated a mouse model carrying a null mutation of the PR gene using embryonic stem cell/gene targeting techniques. Male and female embryos homozygous for the PR mutation developed normally to adulthood. However, the adult female PR mutant displayed significant defects in all reproductive tissues. These included an inability to ovulate, uterine hyperplasia and inflammation, severely limited mammary gland development, and an inability to exhibit sexual behavior. Collectively, these results provide direct support for progesterone's role as a pleiotropic coordinator of diverse reproductive events that together ensure species survival.
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            Alteration of reproductive function but not prenatal sexual development after insertional disruption of the mouse estrogen receptor gene.

            D B Lubahn, J. Moyer, T Golding … (1993)
            Estrogen receptor and its ligand, estradiol, have long been thought to be essential for survival, fertility, and female sexual differentiation and development. Consistent with this proposed crucial role, no human estrogen receptor gene mutations are known, unlike the androgen receptor, where many loss of function mutations have been found. We have generated mutant mice lacking responsiveness to estradiol by disrupting the estrogen receptor gene by gene targeting. Both male and female animals survive to adulthood with normal gross external phenotypes. Females are infertile; males have a decreased fertility. Females have hypoplastic uteri and hyperemic ovaries with no detectable corpora lutea. In adult wild-type and heterozygous females, 3-day estradiol treatment at 40 micrograms/kg stimulates a 3- to 4-fold increase in uterine wet weight and alters vaginal cornification, but the uteri and vagina do not respond in the animals with the estrogen receptor gene disruption. Prenatal male and female reproductive tract development can therefore occur in the absence of estradiol receptor-mediated responsiveness.
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              Time of implantation of the conceptus and loss of pregnancy.

              Marsha Wilcox, Aaron C Weinberg, Richard Baird (1999)
              Implantation of the conceptus is a key step in pregnancy, but little is known about the time of implantation or the relation between the time of implantation and the outcome of pregnancy. We collected daily urine samples for up to six months from 221 women attempting to conceive after ceasing to use contraception. Ovulation was identified on the basis of the ratio of urinary estrogen metabolites to progesterone metabolites, which changes rapidly with luteinization of the ovarian follicle. The time of implantation was defined by the appearance of chorionic gonadotropin in maternal urine. There were 199 conceptions, for 95 percent of which (189) we had sufficient data for analysis. Of these 189 pregnancies, 141 (75 percent) lasted at least six weeks past the last menstrual period, and the remaining 48 pregnancies (25 percent) ended in early loss. Among the pregnancies that lasted six weeks or more, the first appearance of chorionic gonadotropin occurred 6 to 12 days after ovulation; 118 women (84 percent) had implantation on day 8, 9, or 10. The risk of early pregnancy loss increased with later implantation (P<0.001). Among the 102 conceptuses that implanted by the ninth day, 13 percent ended in early loss. This proportion rose to 26 percent with implantation on day 10, to 52 percent on day 11, and to 82 percent after day 11. In most successful human pregnancies, the conceptus implants 8 to 10 days after ovulation. The risk of early pregnancy loss increases with later implantation.
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                Author and article information

                Journal
                Journal ID (nlm-ta): Dev Reprod
                Journal ID (iso-abbrev): Dev Reprod
                Journal ID (publisher-id): Dev Reprod
                Journal ID (publisher-id): DR
                Title: Development & Reproduction
                Publisher: The Korean Society of Developmental Biology
                ISSN (Print): 2465-9525
                ISSN (Electronic): 2465-9541
                Publication date (Print): December 2017
                Publication date (Electronic): 31 December 2017
                Volume: 21
                Issue: 4
                Pages: 351-359
                Affiliations
                [1 ] Dept. of Obstetrics and Gynecology, College of Medicine, Dankook University, Cheonan 31116, Korea
                Author notes
                [† ]Corresponding Author : Jong-Soo Kim, Ph.D., Department of Obstetrics & Gynecology, College of Medicine, Dankook University, 359 Manhyangro, Dongnam-gu, Cheonan 31116, Korea. Tel: +82-41-550-6159, Fax: +82-41-556-3878, E-mail: pluripotent77@ 123456gmail.com
                Article
                Publisher ID: dr-21-4-351
                DOI: 10.12717/DR.2017.21.4.351
                PMC ID: 5769129
                PubMed ID: 29359200
                SO-VID: ebee4c30-fbd7-433a-b476-1b15e92e1fef
                Copyright © ⓒ Copyright 2017 The Korean Society of Developmental Biology
                License:

                This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                Date received : 20 November 2017
                Date revision received : 06 December 2017
                Date accepted : 08 December 2017
                Categories
                Subject: Review
                Custom metadata
                crossmark 2017-12-31

                Keywords: implantation,blastocyst,endometrium,uterine receptivity
                Data availability:
                Keywords: implantation, blastocyst, endometrium, uterine receptivity

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