|Embryology - 23 Nov 2017 Expand to Translate|
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- 1 Introduction
- 2 Some Recent Findings
- 3 Trophoblast and Placental Villi
- 4 Cytotrophoblasts
- 5 Extravillous Cytotrophoblasts
- 6 Syncytiotrophoblasts
- 7 HLA-G
- 8 Trophoblast Infiltration
- 9 Trophoblast Cell Lines
- 10 History
- 11 Abnormalities
- 12 References
- 13 Glossary Links
(Greek, trophe = "nutrition" and -blast, a primordial cell) During early development the trophoblast cells have an important contribution to extraembryonic tissues (fetal placenta and membranes) and processes of early development (adplantation, implantation and endocrine support of pregnancy).
|Human Blastocyst (day 5), trophoblast cells form the peripheral flattened epithelial layer of cells directly under the zona pellucida.||Mouse blastocyst labelled with trophoblast marker.|
In humans, week 1 blastocyst formation the outer layer of cells (adjacent to the zona pellucida) form a flat squamous epithelial layer of cells, the trophoblast layer. Week 2 following blastocyst hatching the trophoblast layer is involved with initial adhesion to the uterine wall and subsequent implantation within the wall. During this period the trophoblast layer proliferates and differentiates into two distinct layers (syncitiotrophoblast and cytotrophoblast).
Following implantation, trophoblast cells continue to contribute to the placenta. Prenatal diagnosis by invasive chorionic villus sampling and [[ non-invasive cervical cell sampling uses mainly DNA from these cells.
|Historical - Who named the trophoblast cell?|
|From an 1905 paper on tubal implantation - "Hubrecht in 1889 published a monograph on the placentation of the hedgehog, showing that the developing ovum in this animal, after sequestration in a crypt, becomes imbedded in the sub-epithelial portion of the mucoa of the uterus by the action of the non-foetal ectodermal cells of the blastocyst. These cells Hubrecht named trophoblast because of their nutritive function, for by their agency the blastocyst “burrows” into the maternal tissues, both destroying and absorbing them."|
- Links: Blastocyst | Implantation | Week 2 | Week 3 | Placenta Development | Prenatal Diagnosis | Trophoblast - Protein Expression | Ectopic Implantation Research
Some Recent Findings
|More recent papers|
This table shows an automated computer PubMed search using the listed sub-heading term.
References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.
Tianying Pan, Guolin He, Meng Chen, Chenyi Bao, Yan Chen, Guangyu Liu, Mi Zhou, Shuying Li, Wenming Xu, Xinghui Liu Abnormal CYP11A1 gene expression induces excessive autophagy, contributing to the pathogenesis of preeclampsia. Oncotarget: 2017, 8(52);89824-89836 PubMed 29163791
Shang Li, Junyu Zhai, Jiansheng Liu, Yan Hong, Weixiu Zhao, Aimin Zhao, Kang Sun, Yanzhi Du, Zi-Jiang Chen BMAL1 facilitates trophoblast migration and invasion via SP1-DNMT1/DAB2IP pathway in recurrent spontaneous abortion. Oncotarget: 2017, 8(52);89451-89464 PubMed 29163762
V Mahajan, H S Banga, G Filia, M P Gupta, K Gupta Comparison of diagnostic tests for the detection of bovine brucellosis in the natural cases of abortion. Iran J Vet Res: 2017, 18(3);183-189 PubMed 29163647
Thy Ph Nguyen, Hannah Ej Yong, Tejasvy Chollangi, Shaun P Brennecke, Susan J Fisher, Euan M Wallace, Peter R Ebeling, Padma Murthi Altered downstream target gene expression of the placental Vitamin D receptor in human idiopathic fetal growth restriction. Cell Cycle: 2017;1-29 PubMed 29161966
Shuang Hou, Jie-Fu Chen, Min Song, Yazhen Zhu, Yu Jen Jan, Szu Hao Chen, Tzu-Hua Weng, Dean-An Ling, Shang-Fu Chen, Tracy Ro, An-Jou Liang, Tom Lee, Helen Jin, Man Li, Lian Liu, Yu-Sheng Hsiao, Peilin Chen, Hsiao-Hua Yu, Ming-Song Tsai, Margareta D Pisarska, Angela Chen, Li-Ching Chen, Hsian-Rong Tseng Correction to Imprinted NanoVelcro Microchips for Isolation and Characterization of Circulating Fetal Trophoblasts: Toward Noninvasive Prenatal Diagnostics. ACS Nano: 2017; PubMed 29160690
Trophoblast and Placental Villi
Early placental development cartoon showing trophoblast contribution to placental villi.
- SYN - syncytiotrophoblasts
- sCTB - subsyncytial cytotrophoblasts (this layer grows increasingly discontinuous in later trimesters)
- EVT - extravillous cytotrophoblasts (anchor the villous tree in the decidua)
Hyperglycosylated human Chorionic Gonadotropin (hCG) promotes the growth of cytotrophoblast cells and the endometrial invasion by these cells during implantation.
Maintain open maternal spiral arteries emptying into the maternal lacunae
At term, depending on maternal decidual localization, there are several identifiable subtypes:
- interstitial mononuclear (and multinuclear) - dispersed in decidual mesenchyme
- endovascular - in spiral arteries lumen (or replacing endothelial cells)
- intramural - in spiral arteries tunica media
- "epithelial" lines - maternal decidua basalis basal plate with maternal endothelial cells in a mosaic fashion
- form by fusion of rapidly dividing cytotrophoblast cells
- secrete proteolytic enzymes, enzymes break down extracellular matrix around cells
- Allow passage of blastocyst into endometrial wall, totally surround the blastocyst
- generate spaces, lacunae, that fill with maternal blood
- secrete Human Chorionic Gonadotropin (hCG), hormone, maintains decidua and Corpus Luteum, basis of pregnancy diagnostic test, present in urine is diagnostic of pregnancy
- Later in development placenta will secrete hCG
Cell–cell Fusion Activity
Two pairs of envelope genes of retroviral origin, syncytins, have fusogenic properties. Human endogenous retroviruses (HERVs) make up about 8% of the human genome. Recent study has shown that placentae from intrauterine growth restriction have impaired cell fusion and differentiation that correlates with reduced levels of HERV envelope genes.
Human - syncytin 1 and syncytin 2
- Syncytin 2 is an envelope gene from the human endogenous retrovirus FRD (HERV-FRD)
- Syncytin 2 receptor is Major Facilitator Superfamily Domain Containing 2 (MFSD2) (at chromosomal position 1p34.2)
- belongs to family of presumptive carbohydrate transporters with 10-12 membrane-spanning domains
- Syncytin 1 (-FRD) receptor is ASCT-1/-2
Mouse - syncytin A and syncytin B
- syncytin A is essential for trophoblast cell differentiation and syncytiotrophoblast morphogenesis
- Links: OMIM Syncytin-2
Human Chorionic Gonadotropin
hCG sources - produced by villous syncytiotrophoblast cells, hyperglycosylated hCG produced by cytotrophoblast cells, free beta-subunit made by multiple primary non-trophoblastic malignancies, and pituitary hCG made by the gonadotrope cells of the anterior pituitary.
- levels peak at 8 to 10 weeks of pregnancy, then decline and are lower for rest of pregnancy
- 0-1 week: 0-50 mIU/ml
- 1-2 weeks: 40-300 mIU/ml
- 3-4 weeks: 500-6,000 mIU/ml
- 1-2 months: 5,000-200,000 mIU/ml
- 2-3 months: 10,000-100,000 mIU/ml
- 2nd trimester: 3,000-50,000 mIU/ml
- 3rd trimester: 1,000-50,000 mIU/ml
Non-pregnant females: <5.0 mIU/ml Postmenopausal females: <9.5 mIU/ml
An acronym for histocompatibility antigen, class I, G (also called: Human Leukocyte Antigen G, (HLA-6.0; HLA60, T-CELL A LOCUS, TCA) and is expressed on placental cytotrophoblast cells and other adult tissues. This distinct tissue distribution differs from the other HLA antigens (HLA-A, HLA-B, HLA-C) leading to the description as a non-classical class I antigen. May have a role in protecting the fetus from the maternal immune response. For example, expression of the HLA-G class, instead of HLA-A and HLA-B, may be a mechanism for avoiding clearance by maternal natural killer cells.
Human gene is located at 6p22.1 and there exist several protein isoforms from alternative splicing of messenger RNAs, membrane-bound isoforms (HLA-G 1-4) and soluble soluble (HLA-G 5-6). The molecule is a heterodimer consisting of both a heavy chain and a light chain (beta-2 microglobulin). The membrane-bound isoform heavy chain is anchored in the membrane and increased expression of the soluble form is related to higher implantation rates. Changes in HLA-G expression have been associated with increased miscarriage rates. Killer cell immunoglobulin-like receptor (KIR) 2DL4 (KIR2DL4) has been shown to act as a receptor for the soluble HLA-G, leading to a stimulation of resting natural killer (NK) cells.
| Human trophoblast invasion
(A) The placenta connects the fetus to the uterine wall and establishes a vascular connection between mother and child. The placenta is structured as villous tree and is in direct contact with maternal blood and, thus referred to as hemochorial. The site where the placenta comes in direct contact with the maternal decidua is called the fetal-maternal interface.
(B) During early pregnancy, vCTBs fuse to form multinucleated STs, which surround the placental villus. STs transport nutrients and gases from the maternal to the fetal circulation and represent the major endocrine unit of the placenta by secreting hormones such as chorionic gonadotropin, placental growth hormone or placental lactogen. AV form cell columns that attach to the maternal decidua and give rise to the EVT lineage. Invasive EVTs can be divided into iCTBs, which invade the decidual stroma and become terminally differentiated multinucleated GCs, or eCTBs.
| Trophoblast plugs within Spiral Arteries Week 9, GA week 11.
Trophoblast Cell Lines
A useful component of current placentation and trophoblast research are a number of trophoblast cell lines that have been derived from a number of different sources including term placenta, choriocarcinomas and transformation of primary isolated cells. BeWo cell line is the earliest human choriocarcinoma, established in 1968. Most appear to be examples of extravillous trophoblast cells, some examples of various human "trophoblast" cell lines are listed below.
- BeWo - human choriocarcinoma, hypotetraploid (modal number = 86; range = 71 to 178) (ATCC CCL-98)
- JEG-3 - choriocarcinoma HTR8/SVneo a transformed extravillous trophoblast line
- SGHPL-4 -  SV40 (pSV3neo) transformed primary cell line express cytokeratin-7, HLA class I antigen, HLA-G, BC-1, CD9, human chorionic gonadotrophin, and human placental lactogen.
- TEV-1 - (HPV16) E6/E7 infection first-trimester extravillous trophoblast cell line
- ACH-3P - fusion of primary human first trimester trophoblasts (GA week 12) with a human choriocarcinoma cell line (AC1-1).
- HChEpC1b - retroviral infection by E6/E7/hTERT
The name "Trophoblast" was used for the first time by Ambrosius Arnold Willem Hubrecht (1853 – 1915) at the meeting of the Anatomical Congress at Wiirzburg in 1888, and its earliest definition is found in the report of that meeting in Nos. 17 and 18 of the Anatomischer Anzeiger, Bd. III. "We there read, concerning a very early stage of the hedgehog (p. 510) : Die aussere Wand der Keimblase ist verdickt (drei bis vierschichtig) und besitzt wabige Lacunen. Fur diese aussere (epiblastische) Schicht sei der. Name Trophoblast gewahlt."
- Links: The Trophoblast - A Rejoinder (1904) | Ambrosius Arnold Willem Hubrecht | Hubrecht Collection | Embryologists | Embryology History
A type of placental abnormality when only the conceptus trophoblast layers proliferates and not the embryoblast, no embryo develops, this is called a "hydatidiform mole", which is due to the continuing presence of the trophoblastic layer, this abnormal conceptus can also implant in the uterus. The trophoblast cells will secrete human chorionic gonadotropin (hCG), as in a normal pregnancy, and may appear maternally and by pregnancy test to be "normal". Prenatal diagnosis by ultrasound analysis demonstrates the absence of a embryo.
There are several forms of hydatidiform mole: partial mole, complete mole and persistent gestational trophoblastic tumor. Many of these tumours arise from a haploid sperm fertilizing an egg without a female pronucleus (the alternative form, an embryo without sperm contribution, is called parthenogenesis). The tumour has a "grape-like" placental appearance without enclosed embryo formation. Following a first molar pregnancy, there is approximately a 1% risk of a second molar pregnancy.
This topic is also covered in Placenta - Abnormalities
- Malaria (More? Abnormal Development - Malaria)
- Chlamydia trachomatis
- Herpesvirus 8 (HHV-8)
- Group B streptococcus (GBS)
- Pu Zhang, Marco Zucchelli, Sara Bruce, Fredwell Hambiliki, Anneli Stavreus-Evers, Lev Levkov, Heli Skottman, Erja Kerkelä, Juha Kere, Outi Hovatta Transcriptome profiling of human pre-implantation development. PLoS ONE: 2009, 4(11);e7844 PubMed 19924284 | PMC2773928 | PLoS One
- Noriko Tanaka, Takumi Takeuchi, Queenie V Neri, Eric Scott Sills, Gianpiero D Palermo Laser-assisted blastocyst dissection and subsequent cultivation of embryonic stem cells in a serum/cell free culture system: applications and preliminary results in a murine model. J Transl Med: 2006, 4;20 PubMed 16681851 | PMC1479373 | J Transl Med.
- Chandni V Jain, Leena Kadam, Marie van Dijk, Hamid-Reza Kohan-Ghadr, Brian A Kilburn, Craig Hartman, Vicki Mazzorana, Allerdien Visser, Michael Hertz, Alan D Bolnick, Rani Fritz, D Randall Armant, Sascha Drewlo Fetal genome profiling at 5 weeks of gestation after noninvasive isolation of trophoblast cells from the endocervical canal. Sci Transl Med: 2016, 8(363);363re4 PubMed 27807286
- Nannan He, Liesbeth van Iperen, Danielle de Jong, Karoly Szuhai, Frans M Helmerhorst, Lucette A J van der Westerlaken, Susana M Chuva de Sousa Lopes Human Extravillous Trophoblasts Penetrate Decidual Veins and Lymphatics before Remodeling Spiral Arteries during Early Pregnancy. PLoS ONE: 2017, 12(1);e0169849 PubMed 28081266
- Mario Rothbauer, Nilaykumar Patel, Hajnalka Gondola, Monika Siwetz, Berthold Huppertz, Peter Ertl A comparative study of five physiological key parameters between four different human trophoblast-derived cell lines. Sci Rep: 2017, 7(1);5892 PubMed 28724925
- Gregor Weiss, Monika Sundl, Andreas Glasner, Berthold Huppertz, Gerit Moser The trophoblast plug during early pregnancy: a deeper insight. Histochem. Cell Biol.: 2016; PubMed 27510415
- Hao Liang, Qian Zhang, Junjie Lu, Genling Yang, Na Tian, Xiaojie Wang, Yi Tan, Dongmei Tan MSX2 Induces Trophoblast Invasion in Human Placenta. PLoS ONE: 2016, 11(4);e0153656 PubMed 27088357
- Renaud, SJ. entail., OVO-like 1 regulates progenitor cell fate in human trophoblast development PNAS
- Yasuhide Ohinata, Tomoyuki Tsukiyama Establishment of trophoblast stem cells under defined culture conditions in mice. PLoS ONE: 2014, 9(9);e107308 PubMed 25203285
- Anna Chuprin, Hilah Gal, Tal Biron-Shental, Anat Biran, Aliza Amiel, Shmuel Rozenblatt, Valery Krizhanovsky Cell fusion induced by ERVWE1 or measles virus causes cellular senescence. Genes Dev.: 2013, 27(21);2356-66 PubMed 24186980 | Genes Dev.
- Sandra M Blois, Irene Tirado-González, Julie Wu, Gabriela Barrientos, Briana Johnson, James Warren, Nancy Freitag, Burghard F Klapp, Ster Irmak, Suleyman Ergun, Gabriela S Dveskler Early expression of pregnancy-specific glycoprotein 22 (PSG22) by trophoblast cells modulates angiogenesis in mice. Biol. Reprod.: 2012, 86(6);191 PubMed 22423048
- Michael A Hubert, Susan L Sherritt, Cindy J Bachurski, Stuart Handwerger Involvement of transcription factor NR2F2 in human trophoblast differentiation. PLoS ONE: 2010, 5(2);e9417 PubMed 20195529 | PLoS One
- G E Lash, K Naruse, B A Innes, S C Robson, R F Searle, J N Bulmer Secretion of angiogenic growth factors by villous cytotrophoblast and extravillous trophoblast in early human pregnancy. Placenta: 2010, 31(6);545-8 PubMed 20338637
- Jennifer R Robbins, Kasia M Skrzypczynska, Varvara B Zeldovich, Mirhan Kapidzic, Anna I Bakardjiev Placental syncytiotrophoblast constitutes a major barrier to vertical transmission of Listeria monocytogenes. PLoS Pathog.: 2010, 6(1);e1000732 PubMed 20107601 | PLoS
- Laurence A Cole Biological functions of hCG and hCG-related molecules. Reprod. Biol. Endocrinol.: 2010, 8;102 PubMed 20735820
- G St J Whitley, J E Cartwright Cellular and molecular regulation of spiral artery remodelling: lessons from the cardiovascular field. Placenta: 2010, 31(6);465-74 PubMed 20359743
- Matthias Ruebner, Pamela L Strissel, Manuela Langbein, Fabian Fahlbusch, David L Wachter, Florian Faschingbauer, Matthias W Beckmann, Reiner Strick Impaired cell fusion and differentiation in placentae from patients with intrauterine growth restriction correlate with reduced levels of HERV envelope genes. J. Mol. Med.: 2010, 88(11);1143-56 PubMed 20664994
- Cécile Esnault, Stéphane Priet, David Ribet, Cécile Vernochet, Thomas Bruls, Christian Lavialle, Jean Weissenbach, Thierry Heidmann A placenta-specific receptor for the fusogenic, endogenous retrovirus-derived, human syncytin-2. Proc. Natl. Acad. Sci. U.S.A.: 2008, 105(45);17532-7 PubMed 18988732 | PNAS
- Anne Dupressoir, Cécile Vernochet, Olivia Bawa, Francis Harper, Gérard Pierron, Paule Opolon, Thierry Heidmann Syncytin-A knockout mice demonstrate the critical role in placentation of a fusogenic, endogenous retrovirus-derived, envelope gene. Proc. Natl. Acad. Sci. U.S.A.: 2009, 106(29);12127-32 PubMed 19564597
- Yuan Q Yao, David H Barlow, Ian L Sargent Differential expression of alternatively spliced transcripts of HLA-G in human preimplantation embryos and inner cell masses. J. Immunol.: 2005, 175(12);8379-85 PubMed 16339579
- Theodora Papamitsou, Alexandros Toskas, Kyriaki Papadopoulou, Antonia Sioga, Sotirios Lakis, Maria Chatzistamatiou, Zinon Economou, Luisa Adriopoulou Immunohistochemical study of immunological markers: HLAG, CD16, CD25, CD56 and CD68 in placenta tissues in recurrent pregnancy loss. Histol. Histopathol.: 2014, 29(8);1047-55 PubMed 24557735
- Sumati Rajagopalan, Yenan T Bryceson, Shanmuga P Kuppusamy, Daniel E Geraghty, Arnold van der Meer, Irma Joosten, Eric O Long Activation of NK cells by an endocytosed receptor for soluble HLA-G. PLoS Biol.: 2006, 4(1);e9 PubMed 16366734
- Philipp Velicky, Martin Knöfler, Jürgen Pollheimer Function and control of human invasive trophoblast subtypes: intrinsic vs. maternal control. Cell Adh Migr: 2015; PubMed 26418186
- R A Pattillo, G O Gey, E Delfs, R F Mattingly Human hormone production in vitro. Science: 1968, 159(3822);1467-9 PubMed 5753554
- M Y Choy, I T Manyonda The phagocytic activity of human first trimester extravillous trophoblast. Hum. Reprod.: 1998, 13(1O);2941-9 PubMed 9804259
- H L LaMarca, C M Ott, K Höner Zu Bentrup, C L Leblanc, D L Pierson, A B Nelson, A B Scandurro, G St J Whitley, C A Nickerson, C A Morris Three-dimensional growth of extravillous cytotrophoblasts promotes differentiation and invasion. Placenta: 2005, 26(10);709-20 PubMed 16226120
- Hui Chen Feng, Mei Yee Choy, Wen Deng, Hing Lok Wong, Wui Man Lau, Annie N Y Cheung, Hextan Y S Ngan, Sai Wah Tsao Establishment and characterization of a human first-trimester extravillous trophoblast cell line (TEV-1). J. Soc. Gynecol. Investig.: 2005, 12(4);e21-32 PubMed 15866109
- Ursula Hiden, Christian Wadsack, Nicole Prutsch, Martin Gauster, Ursula Weiss, Hans-Georg Frank, Ulrike Schmitz, Christa Fast-Hirsch, Markus Hengstschläger, Andy Pötgens, Angela Rüben, Martin Knöfler, Peter Haslinger, Berthold Huppertz, Martin Bilban, Peter Kaufmann, Gernot Desoye The first trimester human trophoblast cell line ACH-3P: a novel tool to study autocrine/paracrine regulatory loops of human trophoblast subpopulations--TNF-alpha stimulates MMP15 expression. BMC Dev. Biol.: 2007, 7;137 PubMed 18093301
- Hiroko Omi, Aikou Okamoto, Takashi Nikaido, Mitsuyoshi Urashima, Rie Kawaguchi, Nagayoshi Umehara, Kentaro Sugiura, Misato Saito, Tohru Kiyono, Tadao Tanaka Establishment of an immortalized human extravillous trophoblast cell line by retroviral infection of E6/E7/hTERT and its transcriptional profile during hypoxia and reoxygenation. Int. J. Mol. Med.: 2009, 23(2);229-36 PubMed 19148547
- Eugenia de la Torre, Melissa J Mulla, Andrew G Yu, Seung-Joon Lee, Paula B Kavathas, Vikki M Abrahams Chlamydia trachomatis infection modulates trophoblast cytokine/chemokine production. J. Immunol.: 2009, 182(6);3735-45 PubMed 19265152
- Mariantonietta Di Stefano, Maria Luisa Calabrò, Iole Maria Di Gangi, Santina Cantatore, Massimo Barbierato, Elisa Bergamo, Anfumbom Jude Kfutwah, Margherita Neri, Luigi Chieco-Bianchi, Pantaleo Greco, Loreto Gesualdo, Ahidjo Ayouba, Elisabeth Menu, Josè Ramòn Fiore In vitro and in vivo human herpesvirus 8 infection of placenta. PLoS ONE: 2008, 3(12);e4073 PubMed 19115001
- Amber Kaplan, Kathy Chung, Hande Kocak, Cristina Bertolotto, Andy Uh, Calvin J Hobel, Charles F Simmons, Kelly Doran, George Y Liu, Ozlem Equils Group B streptococcus induces trophoblast death. Microb. Pathog.: 2008, 45(3);231-5 PubMed 18599257
John D Aplin Developmental cell biology of human villous trophoblast: current research problems. Int. J. Dev. Biol.: 2010, 54(2-3);323-9 PubMed 19876840
Amy Ralston, Brian J Cox, Noriyuki Nishioka, Hiroshi Sasaki, Evelyn Chea, Peter Rugg-Gunn, Guoji Guo, Paul Robson, Jonathan S Draper, Janet Rossant Gata3 regulates trophoblast development downstream of Tead4 and in parallel to Cdx2. Development: 2010, 137(3);395-403 PubMed 20081188
Yohei Hayashi, Miho Kusuda Furue, Satoshi Tanaka, Michiko Hirose, Noriko Wakisaka, Hiroki Danno, Kiyoshi Ohnuma, Shiho Oeda, Yuko Aihara, Kunio Shiota, Atsuo Ogura, Shoichi Ishiura, Makoto Asashima BMP4 induction of trophoblast from mouse embryonic stem cells in defined culture conditions on laminin. In Vitro Cell. Dev. Biol. Anim.: 2010, 46(5);416-30 PubMed 20033790
Jonathan P Stoye Proviral protein provides placental function. Proc. Natl. Acad. Sci. U.S.A.: 2009, 106(29);11827-8 PubMed 19617545
Anne Dupressoir, Cécile Vernochet, Olivia Bawa, Francis Harper, Gérard Pierron, Paule Opolon, Thierry Heidmann Syncytin-A knockout mice demonstrate the critical role in placentation of a fusogenic, endogenous retrovirus-derived, envelope gene. Proc. Natl. Acad. Sci. U.S.A.: 2009, 106(29);12127-32 PubMed 19564597
Srinivas R Viswanathan, Craig H Mermel, Jun Lu, Chi-Wei Lu, Todd R Golub, George Q Daley microRNA expression during trophectoderm specification. PLoS ONE: 2009, 4(7);e6143 PubMed 19582159
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Cite this page: Hill, M.A. 2017 Embryology Trophoblast. Retrieved November 23, 2017, from https://embryology.med.unsw.edu.au/embryology/index.php/Trophoblast
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