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Placenta

The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation. It plays critical roles in facilitating nutrient, gas and waste exchange between the physically separate maternal and fetal circulations, and is an important endocrine organ, producing hormones that regulate both maternal and fetal physiology during pregnancy.[1] The placenta connects to the fetus via the umbilical cord, and on the opposite aspect to the maternal uterus in a species-dependent manner. In humans, a thin layer of maternal decidual (endometrial) tissue comes away with the placenta when it is expelled from the uterus following birth (sometimes incorrectly referred to as the 'maternal part' of the placenta). Placentas are a defining characteristic of placental mammals, but are also found in marsupials and some non-mammals with varying levels of development.[2]

Placenta
Human placenta shown in uterus connected to fetus with umbilical cord
Human placenta from just after birth with the umbilical cord in place
Details
Precursordecidua basalis, chorion frondosum
Identifiers
LatinPlacento
MeSHD010920
TEE5.11.3.1.1.0.5
Anatomical terminology
[edit on Wikidata]

Mammalian placentas probably first evolved about 150 million to 200 million years ago. The protein syncytin, found in the outer barrier of the placenta (the syncytiotrophoblast) between mother and fetus, has a certain RNA signature in its genome that has led to the hypothesis that it originated from an ancient retrovirus: essentially a virus that helped pave the transition from egg-laying to live-birth.[3][4][5]

The word placenta comes from the Latin word for a type of cake, from Greek πλακόεντα/πλακοῦντα plakóenta/plakoúnta, accusative of πλακόεις/πλακούς plakóeis/plakoús, "flat, slab-like",[6][7] with reference to its round, flat appearance in humans. The classical plural is placentae, but the form placentas is more common in modern English.

Phylogenetic diversity

Although all mammalian placentas have the same functions, there are important differences in structure and function in different groups of mammals. For example, human, bovine, equine and canine placentas are very different at both the gross and the microscopic levels. Placentas of these species also differ in their ability to provide maternal immunoglobulins to the fetus.[8]

Structure

Placental mammals, including humans, have a chorioallantoic placenta that forms from the chorion and allantois. In humans, the placenta averages 22 cm (9 inch) in length and 2–2.5 cm (0.8–1 inch) in thickness, with the center being the thickest, and the edges being the thinnest. It typically weighs approximately 500 grams (just over 1 lb). It has a dark reddish-blue or crimson color. It connects to the fetus by an umbilical cord of approximately 55–60 cm (22–24 inch) in length, which contains two umbilical arteries and one umbilical vein.[9] The umbilical cord inserts into the chorionic plate (has an eccentric attachment). Vessels branch out over the surface of the placenta and further divide to form a network covered by a thin layer of cells. This results in the formation of villous tree structures. On the maternal side, these villous tree structures are grouped into lobules called cotyledons. In humans, the placenta usually has a disc shape, but size varies vastly between different mammalian species.[10]

The placenta occasionally takes a form in which it comprises several distinct parts connected by blood vessels.[11] The parts, called lobes, may number two, three, four, or more. Such placentas are described as bilobed/bilobular/bipartite, trilobed/trilobular/tripartite, and so on. If there is a clearly discernible main lobe and auxiliary lobe, the latter is called a succenturiate placenta. Sometimes the blood vessels connecting the lobes get in the way of fetal presentation during labor, which is called vasa previa.

Gene and protein expression

About 20,000 protein coding genes are expressed in human cells and 70% of these genes are expressed in the normal mature placenta.[12][13] Some 350 of these genes are more specifically expressed in the placenta and fewer than 100 genes are highly placenta specific. The corresponding specific proteins are mainly expressed in trophoblasts and have functions related to pregnancy. Examples of proteins with elevated expression in placenta compared to other organs and tissues are PEG10 and the cancer testis antigen PAGE4 and expressed in cytotrophoblasts, CSH1 and KISS1 expressed in syncytiotrophoblasts, and PAPPA2 and PRG2 expressed in extravillous trophoblasts.

Physiology

Development

 
Placenta
 
The initial stages of human embryogenesis

The placenta begins to develop upon implantation of the blastocyst into the maternal endometrium, very early on in pregnancy at about week 4.[14]

The outer layer of the late blastocyst, is formed of trophoblasts, cells that form the outer layer of the placenta. This outer layer is divided into two further layers: the underlying cytotrophoblast layer and the overlying syncytiotrophoblast layer. The syncytiotrophoblast is a multinucleated continuous cell layer that covers the surface of the placenta. It forms as a result of differentiation and fusion of the underlying cytotrophoblasts, a process that continues throughout placental development. The syncytiotrophoblast contributes to the barrier function of the placenta.[15]

The placenta grows throughout pregnancy. Development of the maternal blood supply to the placenta is complete by the end of the first trimester of pregnancy week 14 (DM).[14]

Placental circulation

 
Maternal blood fills the intervillous space, nutrients, water, and gases are actively and passively exchanged, then deoxygenated blood is displaced by the next maternal pulse.

Maternal placental circulation

In preparation for implantation of the blastocyst, the endometrium undergoes decidualization. Spiral arteries in the decidua are remodeled so that they become less convoluted and their diameter is increased. The increased diameter and straighter flow path both act to increase maternal blood flow to the placenta. There is relatively high pressure as the maternal blood fills intervillous space through these spiral arteries which bathe the fetal villi in blood, allowing an exchange of gases to take place. In humans and other hemochorial placentals, the maternal blood comes into direct contact with the fetal chorion, though no fluid is exchanged. As the pressure decreases between pulses, the deoxygenated blood flows back through the endometrial veins.

Maternal blood flow begins between days 5–12,[16] and is approximately 600–700 ml/min at term.

Fetoplacental circulation

Deoxygenated fetal blood passes through umbilical arteries to the placenta. At the junction of umbilical cord and placenta, the umbilical arteries branch radially to form chorionic arteries. Chorionic arteries, in turn, branch into cotyledon arteries. In the villi, these vessels eventually branch to form an extensive arterio-capillary-venous system, bringing the fetal blood extremely close to the maternal blood; but no intermingling of fetal and maternal blood occurs ("placental barrier").[17]

Endothelin and prostanoids cause vasoconstriction in placental arteries, while nitric oxide causes vasodilation.[18] On the other hand, there is no neural vascular regulation, and catecholamines have only little effect.[18]

The fetoplacental circulation is vulnerable to persistent hypoxia or intermittent hypoxia and reoxygenation, which can lead to generation of excessive free radicals. This may contribute to pre-eclampsia and other pregnancy complications.[19] It is proposed that melatonin plays a role as an antioxidant in the placenta.[19]

This begins at day 17–22.[20]

Birth

Placental expulsion begins as a physiological separation from the wall of the uterus. The period from just after the child is born until just after the placenta is expelled is called the "third stage of labor". The placenta is usually expelled within 15–30 minutes of birth.

Placental expulsion can be managed actively, for example by giving oxytocin via intramuscular injection followed by cord traction to assist in delivering the placenta. Alternatively, it can be managed expectantly, allowing the placenta to be expelled without medical assistance. Blood loss and the risk of postpartum bleeding may be reduced in women offered active management of the third stage of labour, however there may be adverse effects and more research is necessary.[21]

The habit is to cut the cord immediately after birth, but it is theorised that there is no medical reason to do this; on the contrary, it is theorised that not cutting the cord helps the baby in its adaptation to extrauterine life, especially in preterm infants.[22]

Microbiome

The placenta is traditionally thought to be sterile, but recent research suggests that a resident, non-pathogenic, and diverse population of microorganisms may be present in healthy tissue. However, whether these microbes exist or are clinically important is highly controversial and is the subject of active research.[23][24][25][26]

Functions

Nutrition and gas exchange

The placenta intermediates the transfer of nutrients between mother and fetus. The perfusion of the intervillous spaces of the placenta with maternal blood allows the transfer of nutrients and oxygen from the mother to the fetus and the transfer of waste products and carbon dioxide back from the fetus to the maternal blood. Nutrient transfer to the fetus can occur via both active and passive transport.[27] Placental nutrient metabolism was found to play a key role in limiting the transfer of some nutrients.[28] Adverse pregnancy situations, such as those involving maternal diabetes or obesity, can increase or decrease levels of nutrient transporters in the placenta potentially resulting in overgrowth or restricted growth of the fetus.[29]

 
Animated schematic of the hearts and circulatory systems of a fetus and its mother – red and blue represent oxygenated and deoxygenated blood, respectively (animation)

Excretion

Waste products excreted from the fetus such as urea, uric acid, and creatinine are transferred to the maternal blood by diffusion across the placenta.

Immunity

The placenta functions as a selective barrier between maternal and fetal cells, preventing maternal blood, proteins and microbes (including bacteria and most viruses) from crossing the maternal-fetal barrier.[30] Deterioration in placental functioning, referred to as placental insufficiency, may be related to mother-to-child transmission of some infectious diseases.[31] A very small number of viruses including rubella virus, Zika virus and cytomegalovirus (CMV) can travel across the placental barrier, generally taking advantage of conditions at certain gestational periods as the placenta develops. CMV and Zika travel from the maternal bloodstream via placental cells to the fetal bloodstream.[30][32][33][34]

Beginning as early as 13 weeks of gestation, and increasing linearly, with the largest transfer occurring in the third trimester, IgG antibodies can pass through the human placenta, providing protection to the fetus in utero.[35][36] This passive immunity lingers for several months after birth, providing the newborn with a carbon copy of the mother's long-term humoral immunity to see the infant through the crucial first months of extrauterine life. IgM antibodies, because of their larger size, cannot cross the placenta,[37] one reason why infections acquired during pregnancy can be particularly hazardous for the fetus.[38]

Endocrine function

  • The first hormone released by the placenta is called the human chorionic gonadotropin (hCG) hormone. This is responsible for stopping the process at the end of menses when the corpus luteum ceases activity and atrophies. If hCG did not interrupt this process, it would lead to spontaneous abortion of the fetus. The corpus luteum also produces and releases progesterone and estrogen, and hCG stimulates it to increase the amount that it releases. hCG is the indicator of pregnancy that pregnancy tests look for. These tests will work when menses has not occurred or after implantation has happened on days seven to ten. hCG may also have an anti-antibody effect, protecting it from being rejected by the mother's body. hCG also assists the male fetus by stimulating the testes to produce testosterone, which is the hormone needed to allow the sex organs of the male to grow.
  • Progesterone helps the embryo implant by assisting passage through the fallopian tubes. It also affects the fallopian tubes and the uterus by stimulating an increase in secretions necessary for fetal nutrition. Progesterone, like hCG, is necessary to prevent spontaneous abortion because it prevents contractions of the uterus and is necessary for implantation.
  • Estrogen is a crucial hormone in the process of proliferation. This involves the enlargement of the breasts and uterus, allowing for growth of the fetus and production of milk. Estrogen is also responsible for increased blood supply towards the end of pregnancy through vasodilation. The levels of estrogen during pregnancy can increase so that they are thirty times what a non-pregnant woman mid-cycles estrogen level would be.
  • Human placental lactogen (hPL) is a hormone used in pregnancy to develop fetal metabolism and general growth and development. Human placental lactogen works with growth hormone to stimulate Insulin-like growth factor production and regulating intermediary metabolism. In the fetus, hPL acts on lactogenic receptors to modulate embryonic development, metabolism and stimulate production of IGF, insulin, surfactant and adrenocortical hormones. hPL values increase with multiple pregnancies, intact molar pregnancy, diabetes and Rh incompatibility. They are decreased with toxemia, choriocarcinoma, and Placental insufficiency.[39][40]

Immunological barrier

The placenta and fetus may be regarded as a foreign body inside the mother and must be protected from the normal immune response of the mother that would cause it to be rejected. The placenta and fetus are thus treated as sites of immune privilege, with immune tolerance.

For this purpose, the placenta uses several mechanisms :

However, the placental barrier is not the sole means of evading the immune system, as foreign fetal cells also persist in the maternal circulation, on the other side of the placental barrier.[43]

Other

The placenta also provides a reservoir of blood for the fetus, delivering blood to it in case of hypotension and vice versa, comparable to a capacitor.[44]

 
Ultrasound image of human placenta and umbilical cord (color Doppler rendering) with central cord insertion and three umbilical vessels, at 20 weeks of pregnancy

Clinical significance

 
Micrograph of a cytomegalovirus (CMV) infection of the placenta (CMV placentitis). The characteristic large nucleus of a CMV-infected cell is seen off-centre at the bottom-right of the image. H&E stain.

Numerous pathologies can affect the placenta.

Society and culture

The placenta often plays an important role in various cultures, with many societies conducting rituals regarding its disposal. In the Western world, the placenta is most often incinerated.[45]

Some cultures bury the placenta for various reasons. The Māori of New Zealand traditionally bury the placenta from a newborn child to emphasize the relationship between humans and the earth.[46] Likewise, the Navajo bury the placenta and umbilical cord at a specially chosen site,[47] particularly if the baby dies during birth.[48] In Cambodia and Costa Rica, burial of the placenta is believed to protect and ensure the health of the baby and the mother.[49] If a mother dies in childbirth, the Aymara of Bolivia bury the placenta in a secret place so that the mother's spirit will not return to claim her baby's life.[50]

The placenta is believed by some communities to have power over the lives of the baby or its parents. The Kwakiutl of British Columbia bury girls' placentas to give the girl skill in digging clams, and expose boys' placentas to ravens to encourage future prophetic visions. In Turkey, the proper disposal of the placenta and umbilical cord is believed to promote devoutness in the child later in life. In Transylvania, and Japan, interaction with a disposed placenta is thought to influence the parents' future fertility.[citation needed]

Several cultures believe the placenta to be or have been alive, often a relative of the baby. Nepalese think of the placenta as a friend of the baby; the orang Asli and Malay populations in Malay Peninsula regard it as the baby's older sibling.[49][51] Native Hawaiians believe that the placenta is a part of the baby, and traditionally plant it with a tree that can then grow alongside the child.[45] Various cultures in Indonesia, such as Javanese and Malay, believe that the placenta has a spirit and needs to be buried outside the family house. Some Malays would bury the baby's placenta with a pencil (if it is a boy) or a needle and thread (if it is a girl).[51]

In some cultures, the placenta is eaten, a practice known as placentophagy. In some eastern cultures, such as China, the dried placenta (ziheche 紫河车, literally "purple river car") is thought to be a healthful restorative and is sometimes used in preparations of traditional Chinese medicine and various health products.[52] The practice of human placentophagy has become a more recent trend in western cultures and is not without controversy; its practice being considered cannibalism is debated.

Some cultures have alternative uses for placenta that include the manufacturing of cosmetics, pharmaceuticals and food.[53]

Additional images

See also

References

  1. ^ Jin M, Xu Q, Li J, Xu S, Tang C (September 2022). "Micro-RNAs in Human Placenta: Tiny Molecules, Immense Power". Molecules. 27 (18): 5943. doi:10.3390/molecules27185943. PMC 9501247. PMID 36144676.
  2. ^ Pough, F. H.; Andrews, Robin M.; Cadle, John E.; Crump, Martha L.; Savitsky, Alan H.; Wells, Kentwood D. (2004). Herpetology (3rd ed.). Pearson. ISBN 978-0-13-100849-6.[page needed]
  3. ^ Mitra, Avir (31 January 2020). "How the placenta evolved from an ancient virus". WHYY. Retrieved 9 March 2020.
  4. ^ Chuong, Edward B. (9 October 2018). "The placenta goes viral: Retroviruses control gene expression in pregnancy". PLOS Biology. 16 (10): e3000028. doi:10.1371/journal.pbio.3000028. PMC 6177113. PMID 30300353.
  5. ^ Villarreal, Luis P. (January 2016). "Viruses and the placenta: the essential virus first view". APMIS. 124 (1–2): 20–30. doi:10.1111/apm.12485. PMID 26818259. S2CID 12042851.
  6. ^ Henry George Liddell, Robert Scott, "A Greek-English Lexicon", at Perseus 2012-04-05 at the Wayback Machine.
  7. ^ "placenta" 2016-01-30 at the Wayback Machine. Online Etymology Dictionary.
  8. ^ Bowen, R. Implantation and Development of the Placenta: Introduction and Index. From: Pathophysiology of the Reproductive System. Accessed: 7 July 2019.
  9. ^ Examination of the placenta 2011-10-16 at the Wayback Machine[dead link]
  10. ^ Placental Structure and Classification 2016-02-11 at the Wayback Machine
  11. ^ Fujikura, Toshio; Benson, Ralph C; Driscoll, Shirley G; et al. (1970), "The bipartite placenta and its clinical features", American Journal of Obstetrics and Gynecology, 107 (7): 1013–1017, doi:10.1016/0002-9378(70)90621-6, PMID 5429965, Bipartite placenta represented 4.2 per cent (366 of 8,505) of placentas of white women at the Boston Hospital for Women who were enrolled in the Collaborative Project.
  12. ^ "The human proteome in placenta - The Human Protein Atlas". www.proteinatlas.org. from the original on 2017-09-26. Retrieved 2017-09-26.
  13. ^ Uhlén, Mathias; Fagerberg, Linn; Hallström, Björn M.; Lindskog, Cecilia; Oksvold, Per; Mardinoglu, Adil; Sivertsson, Åsa; Kampf, Caroline; Sjöstedt, Evelina (2015-01-23). "Tissue-based map of the human proteome". Science. 347 (6220): 1260419. doi:10.1126/science.1260419. ISSN 0036-8075. PMID 25613900. S2CID 802377.
  14. ^ a b "Stages of Development of the Fetus - Women's Health Issues". Merck Manuals Consumer Version. Retrieved 2022-06-12.
  15. ^ "How Your Fetus Grows During Pregnancy". www.acog.org. Retrieved 2022-06-12.
  16. ^ Dashe, Jodi S.; Bloom, Steven L.; Spong, Catherine Y.; Hoffman, Barbara L. (2018). Williams Obstetrics. McGraw Hill Professional. ISBN 978-1-259-64433-7.[page needed]
  17. ^ Placental blood circulation 2011-09-28 at the Wayback Machine
  18. ^ a b Kiserud T, Acharya G (2004). "The fetal circulation". Prenatal Diagnosis. 24 (13): 1049–1059. doi:10.1002/pd.1062. PMID 15614842. S2CID 25040285.
  19. ^ a b Reiter, R. J.; Tan, D. X.; Korkmaz, A.; Rosales-Corral, S. A. (2013). "Melatonin and stable circadian rhythms optimize maternal, placental and fetal physiology". Human Reproduction Update. 20 (2): 293–307. doi:10.1093/humupd/dmt054. ISSN 1355-4786. PMID 24132226.
  20. ^ Williams book of obsteritcis.
  21. ^ Begley, CM; Gyte, GM; Devane, D; McGuire, W; Weeks, A; Biesty, LM (13 February 2019). "Active versus expectant management for women in the third stage of labour". Cochrane Database of Systematic Reviews. 2019 (2): CD007412. doi:10.1002/14651858.CD007412.pub5. PMC 6372362. PMID 30754073.
  22. ^ Mercer JS, Vohr BR, Erickson-Owens DA, Padbury JF, Oh W (2010). "Seven-month developmental outcomes of very low birth weight infants enrolled in a randomized controlled trial of delayed versus immediate cord clamping". Journal of Perinatology. 30 (1): 11–6. doi:10.1038/jp.2009.170. PMC 2799542. PMID 19847185.
  23. ^ Perez-Muñoz, Maria Elisa; Arrieta, Marie-Claire; Ramer-Tait, Amanda E.; Walter, Jens (2017). "A critical assessment of the "sterile womb" and "in utero colonization" hypotheses: implications for research on the pioneer infant icrobiome". Microbiome. 5 (1): 48. doi:10.1186/s40168-017-0268-4. ISSN 2049-2618. PMC 5410102. PMID 28454555.
  24. ^ Mor, Gil; Kwon, Ja-Young (2015). "Trophoblast-microbiome interaction: a new paradigm on immune regulation". American Journal of Obstetrics and Gynecology. 213 (4): S131–S137. doi:10.1016/j.ajog.2015.06.039. ISSN 0002-9378. PMC 6800181. PMID 26428492.
  25. ^ Prince, Amanda L.; Antony, Kathleen M.; Chu, Derrick M.; Aagaard, Kjersti M. (2014). "The microbiome, parturition, and timing of birth: more questions than answers". Journal of Reproductive Immunology. 104–105: 12–19. doi:10.1016/j.jri.2014.03.006. ISSN 0165-0378. PMC 4157949. PMID 24793619.
  26. ^ Hornef, M; Penders, J (2017). "Does a prenatal bacterial microbiota exist?". Mucosal Immunology. 10 (3): 598–601. doi:10.1038/mi.2016.141. PMID 28120852.
  27. ^ Wright C, Sibley CP (2011). "Placental Transfer in Health and Disease". In Kay H, Nelson M, Yuping W (eds.). The Placenta: From Development to Disease. John Wiley and Sons. pp. 66. ISBN 978-1-4443-3366-4.
  28. ^ Perazzolo S, Hirschmugl B, Wadsack C, Desoye G, Lewis RM, Sengers BG (February 2017). "The influence of placental metabolism on fatty acid transfer to the fetus". J. Lipid Res. 58 (2): 443–454. doi:10.1194/jlr.P072355. PMC 5282960. PMID 27913585.
  29. ^ Kappen C, Kruger C, MacGowan J, Salbaum JM (2012). "Maternal diet modulates placenta growth and gene expression in a mouse model of diabetic pregnancy". PLOS ONE. 7 (6): e38445. Bibcode:2012PLoSO...738445K. doi:10.1371/journal.pone.0038445. PMC 3372526. PMID 22701643.
  30. ^ a b Madhusoodanan, Jyoti (October 10, 2018). "Break on through: How some viruses infect the placenta". Knowable Magazine. doi:10.1146/knowable-101018-1.
  31. ^ Erlebacher, Adrian (2013-03-21). "Immunology of the Maternal-Fetal Interface". Annual Review of Immunology. 31 (1): 387–411. doi:10.1146/annurev-immunol-032712-100003. PMID 23298207. Retrieved 25 June 2021.
  32. ^ Pereira, Lenore (2018-09-29). "Congenital Viral Infection: Traversing the Uterine-Placental Interface". Annual Review of Virology. 5 (1): 273–299. doi:10.1146/annurev-virology-092917-043236. PMID 30048217. S2CID 51724379. Retrieved 24 June 2021.
  33. ^ Arora, Nitin; Sadovsky, Yoel; Dermody, Terence S.; Coyne, Carolyn B. (May 2017). "Microbial Vertical Transmission during Human Pregnancy". Cell Host & Microbe. 21 (5): 561–567. doi:10.1016/j.chom.2017.04.007. PMC 6148370. PMID 28494237.
  34. ^ Robbins, Jennifer R; Bakardjiev, Anna I (February 2012). "Pathogens and the placental fortress". Current Opinion in Microbiology. 15 (1): 36–43. doi:10.1016/j.mib.2011.11.006. PMC 3265690. PMID 22169833.
  35. ^ Palmeira, Patricia; Quinello, Camila; Silveira-Lessa, Ana Lúcia; Zago, Cláudia Augusta; Carneiro-Sampaio, Magda (2012). "IgG Placental Transfer in Healthy and Pathological Pregnancies". Clinical and Developmental Immunology. 2012: 985646. doi:10.1155/2012/985646. PMC 3251916. PMID 22235228.
  36. ^ Simister N. E., Story C. M. (1997). "Human placental Fc receptors and the transmission of antibodies from mother to fetus". Journal of Reproductive Immunology. 37 (1): 1–23. doi:10.1016/s0165-0378(97)00068-5. PMID 9501287.
  37. ^ Pillitteri, Adele (2009). Maternal and Child Health Nursing: Care of the Childbearing and Childrearing Family. Hagerstwon, MD: Lippincott Williams & Wilkins. p. 202. ISBN 978-1-58255-999-5.
  38. ^ "What infections can affect pregnancy?". National Institute of Child Health and Human Development. Retrieved 25 June 2021.
  39. ^ Handwerger S, Freemark M (2000). "The roles of placental growth hormone and placental lactogen in the regulation of human fetal growth and development". Journal of Pediatric Endocrinology & Metabolism. 13 (4): 343–56. doi:10.1515/jpem.2000.13.4.343. PMID 10776988. S2CID 28778529.
  40. ^ . www.ucsfhealth.org. May 17, 2009. Archived from the original on April 29, 2017. Retrieved July 21, 2017.
  41. ^ "Placenta 'fools body's defences'". BBC News. 10 November 2007. from the original on 29 April 2012.
  42. ^ Clark DA, Chaput A, Tutton D (March 1986). "Active suppression of host-vs-graft reaction in pregnant mice. VII. Spontaneous abortion of allogeneic CBA/J x DBA/2 fetuses in the uterus of CBA/J mice correlates with deficient non-T suppressor cell activity". J. Immunol. 136 (5): 1668–75. PMID 2936806.
  43. ^ Williams Z, Zepf D, Longtine J, Anchan R, Broadman B, Missmer SA, Hornstein MD (March 2008). "Foreign fetal cells persist in the maternal circulation". Fertil. Steril. 91 (6): 2593–5. doi:10.1016/j.fertnstert.2008.02.008. PMID 18384774.
  44. ^ Assad RS, Lee FY, Hanley FL (2001). "Placental compliance during fetal extracorporeal circulation". Journal of Applied Physiology. 90 (5): 1882–1886. doi:10.1152/jappl.2001.90.5.1882. PMID 11299282. S2CID 8785947.
  45. ^ a b "Why eat a placenta?". BBC News. 18 April 2006.
  46. ^ Metge, Joan (2005). "Working In/Playing With Three Languages". Sites: A Journal of Social Anthropology and Cultural Studies. 2 (2): 83–90. doi:10.11157/sites-vol2iss2id65.
  47. ^ Francisco, Edna (3 December 2004). "Bridging the Cultural Divide in Medicine". Science.
  48. ^ Shepardson, Mary (1978). "Changes in Navajo mortuary practices and beliefs". American Indian Quarterly. 4 (4): 383–96. doi:10.2307/1184564. JSTOR 1184564. PMID 11614175.
  49. ^ a b Buckley, Sarah J. . Mothering. Archived from the original on 6 January 2008. Retrieved 7 January 2008.
  50. ^ Davenport, Ann (June 2005). "The Love Offer". Johns Hopkins Magazine. 57 (3).
  51. ^ a b Anisah Barakbah (2017). Ensiklopedia Perbidanan Melayu. Universiti Islam Malaysia Press. pp. 236–237. ISBN 978-967-13305-9-3.
  52. ^ Falcao, Ronnie. "Medicinal Uses of the Placenta". from the original on 5 December 2008. Retrieved 25 November 2008.
  53. ^ Kroløkke, Charlotte; Dickinson, Elizabeth; Foss, Karen A (May 2018). "The placenta economy: From trashed to treasured bio-products". European Journal of Women's Studies. 25 (2): 138–153. doi:10.1177/1350506816679004. ISSN 1350-5068. S2CID 151874106.
  54. ^ "Placental Weights: Means, Standard Deviations, and Percentiles by Gestational Age". Placental and Gestational Pathology. 2017. p. 336. doi:10.1017/9781316848616.039. ISBN 978-1-316-84861-6.

External links

  • The placenta-specific proteome at the Human Protein Atlas
  • The Placenta, gynob.com, with quotes from Williams Obstetrics, 18th Edition, F. Gary Cunningham, M.D., Paul C. MacDonald, M.D., Norman F. Grant, M.D., Appleton & Lange, Publishers.

placenta, this, article, about, human, placenta, general, information, about, placenta, organ, biology, tion, ancient, roman, bread, food, expulsion, placenta, after, birth, placental, expulsion, placenta, other, animals, placenta, temporary, embryonic, later,. This article is about the human placenta For general information about the placenta as an organ in biology see Placentation For the ancient Roman Bread see Placenta food For the expulsion of the placenta after birth see placental expulsion For placenta in other animals see Placentalia The placenta is a temporary embryonic and later fetal organ that begins developing from the blastocyst shortly after implantation It plays critical roles in facilitating nutrient gas and waste exchange between the physically separate maternal and fetal circulations and is an important endocrine organ producing hormones that regulate both maternal and fetal physiology during pregnancy 1 The placenta connects to the fetus via the umbilical cord and on the opposite aspect to the maternal uterus in a species dependent manner In humans a thin layer of maternal decidual endometrial tissue comes away with the placenta when it is expelled from the uterus following birth sometimes incorrectly referred to as the maternal part of the placenta Placentas are a defining characteristic of placental mammals but are also found in marsupials and some non mammals with varying levels of development 2 PlacentaHuman placenta shown in uterus connected to fetus with umbilical cordHuman placenta from just after birth with the umbilical cord in placeDetailsPrecursordecidua basalis chorion frondosumIdentifiersLatinPlacentoMeSHD010920TEE5 11 3 1 1 0 5Anatomical terminology edit on Wikidata For a less technical description see the Simple English Wikipedia version at simple Placenta Mammalian placentas probably first evolved about 150 million to 200 million years ago The protein syncytin found in the outer barrier of the placenta the syncytiotrophoblast between mother and fetus has a certain RNA signature in its genome that has led to the hypothesis that it originated from an ancient retrovirus essentially a virus that helped pave the transition from egg laying to live birth 3 4 5 The word placenta comes from the Latin word for a type of cake from Greek plakoenta plakoῦnta plakoenta plakounta accusative of plakoeis plakoys plakoeis plakous flat slab like 6 7 with reference to its round flat appearance in humans The classical plural is placentae but the form placentas is more common in modern English Contents 1 Phylogenetic diversity 2 Structure 2 1 Gene and protein expression 3 Physiology 3 1 Development 3 2 Placental circulation 3 2 1 Maternal placental circulation 3 2 2 Fetoplacental circulation 3 3 Birth 3 4 Microbiome 4 Functions 4 1 Nutrition and gas exchange 4 2 Excretion 4 3 Immunity 4 4 Endocrine function 4 5 Immunological barrier 4 6 Other 5 Clinical significance 6 Society and culture 7 Additional images 8 See also 9 References 10 External linksPhylogenetic diversity EditAlthough all mammalian placentas have the same functions there are important differences in structure and function in different groups of mammals For example human bovine equine and canine placentas are very different at both the gross and the microscopic levels Placentas of these species also differ in their ability to provide maternal immunoglobulins to the fetus 8 Structure EditPlacental mammals including humans have a chorioallantoic placenta that forms from the chorion and allantois In humans the placenta averages 22 cm 9 inch in length and 2 2 5 cm 0 8 1 inch in thickness with the center being the thickest and the edges being the thinnest It typically weighs approximately 500 grams just over 1 lb It has a dark reddish blue or crimson color It connects to the fetus by an umbilical cord of approximately 55 60 cm 22 24 inch in length which contains two umbilical arteries and one umbilical vein 9 The umbilical cord inserts into the chorionic plate has an eccentric attachment Vessels branch out over the surface of the placenta and further divide to form a network covered by a thin layer of cells This results in the formation of villous tree structures On the maternal side these villous tree structures are grouped into lobules called cotyledons In humans the placenta usually has a disc shape but size varies vastly between different mammalian species 10 The placenta occasionally takes a form in which it comprises several distinct parts connected by blood vessels 11 The parts called lobes may number two three four or more Such placentas are described as bilobed bilobular bipartite trilobed trilobular tripartite and so on If there is a clearly discernible main lobe and auxiliary lobe the latter is called a succenturiate placenta Sometimes the blood vessels connecting the lobes get in the way of fetal presentation during labor which is called vasa previa Gene and protein expression Edit Further information Bioinformatics Gene and protein expression About 20 000 protein coding genes are expressed in human cells and 70 of these genes are expressed in the normal mature placenta 12 13 Some 350 of these genes are more specifically expressed in the placenta and fewer than 100 genes are highly placenta specific The corresponding specific proteins are mainly expressed in trophoblasts and have functions related to pregnancy Examples of proteins with elevated expression in placenta compared to other organs and tissues are PEG10 and the cancer testis antigen PAGE4 and expressed in cytotrophoblasts CSH1 and KISS1 expressed in syncytiotrophoblasts and PAPPA2 and PRG2 expressed in extravillous trophoblasts Physiology EditDevelopment Edit Placenta The initial stages of human embryogenesis Further information Placentation The placenta begins to develop upon implantation of the blastocyst into the maternal endometrium very early on in pregnancy at about week 4 14 The outer layer of the late blastocyst is formed of trophoblasts cells that form the outer layer of the placenta This outer layer is divided into two further layers the underlying cytotrophoblast layer and the overlying syncytiotrophoblast layer The syncytiotrophoblast is a multinucleated continuous cell layer that covers the surface of the placenta It forms as a result of differentiation and fusion of the underlying cytotrophoblasts a process that continues throughout placental development The syncytiotrophoblast contributes to the barrier function of the placenta 15 The placenta grows throughout pregnancy Development of the maternal blood supply to the placenta is complete by the end of the first trimester of pregnancy week 14 DM 14 Placental circulation Edit Maternal blood fills the intervillous space nutrients water and gases are actively and passively exchanged then deoxygenated blood is displaced by the next maternal pulse Maternal placental circulation Edit In preparation for implantation of the blastocyst the endometrium undergoes decidualization Spiral arteries in the decidua are remodeled so that they become less convoluted and their diameter is increased The increased diameter and straighter flow path both act to increase maternal blood flow to the placenta There is relatively high pressure as the maternal blood fills intervillous space through these spiral arteries which bathe the fetal villi in blood allowing an exchange of gases to take place In humans and other hemochorial placentals the maternal blood comes into direct contact with the fetal chorion though no fluid is exchanged As the pressure decreases between pulses the deoxygenated blood flows back through the endometrial veins Maternal blood flow begins between days 5 12 16 and is approximately 600 700 ml min at term Fetoplacental circulation Edit Further information Fetal circulation Deoxygenated fetal blood passes through umbilical arteries to the placenta At the junction of umbilical cord and placenta the umbilical arteries branch radially to form chorionic arteries Chorionic arteries in turn branch into cotyledon arteries In the villi these vessels eventually branch to form an extensive arterio capillary venous system bringing the fetal blood extremely close to the maternal blood but no intermingling of fetal and maternal blood occurs placental barrier 17 Endothelin and prostanoids cause vasoconstriction in placental arteries while nitric oxide causes vasodilation 18 On the other hand there is no neural vascular regulation and catecholamines have only little effect 18 The fetoplacental circulation is vulnerable to persistent hypoxia or intermittent hypoxia and reoxygenation which can lead to generation of excessive free radicals This may contribute to pre eclampsia and other pregnancy complications 19 It is proposed that melatonin plays a role as an antioxidant in the placenta 19 This begins at day 17 22 20 Birth Edit Main article Placental expulsion Placental expulsion begins as a physiological separation from the wall of the uterus The period from just after the child is born until just after the placenta is expelled is called the third stage of labor The placenta is usually expelled within 15 30 minutes of birth Placental expulsion can be managed actively for example by giving oxytocin via intramuscular injection followed by cord traction to assist in delivering the placenta Alternatively it can be managed expectantly allowing the placenta to be expelled without medical assistance Blood loss and the risk of postpartum bleeding may be reduced in women offered active management of the third stage of labour however there may be adverse effects and more research is necessary 21 The habit is to cut the cord immediately after birth but it is theorised that there is no medical reason to do this on the contrary it is theorised that not cutting the cord helps the baby in its adaptation to extrauterine life especially in preterm infants 22 Microbiome Edit Main article Placental microbiome The placenta is traditionally thought to be sterile but recent research suggests that a resident non pathogenic and diverse population of microorganisms may be present in healthy tissue However whether these microbes exist or are clinically important is highly controversial and is the subject of active research 23 24 25 26 Functions EditNutrition and gas exchange Edit The placenta intermediates the transfer of nutrients between mother and fetus The perfusion of the intervillous spaces of the placenta with maternal blood allows the transfer of nutrients and oxygen from the mother to the fetus and the transfer of waste products and carbon dioxide back from the fetus to the maternal blood Nutrient transfer to the fetus can occur via both active and passive transport 27 Placental nutrient metabolism was found to play a key role in limiting the transfer of some nutrients 28 Adverse pregnancy situations such as those involving maternal diabetes or obesity can increase or decrease levels of nutrient transporters in the placenta potentially resulting in overgrowth or restricted growth of the fetus 29 Animated schematic of the hearts and circulatory systems of a fetus and its mother red and blue represent oxygenated and deoxygenated blood respectively animation Excretion Edit Waste products excreted from the fetus such as urea uric acid and creatinine are transferred to the maternal blood by diffusion across the placenta Immunity Edit The placenta functions as a selective barrier between maternal and fetal cells preventing maternal blood proteins and microbes including bacteria and most viruses from crossing the maternal fetal barrier 30 Deterioration in placental functioning referred to as placental insufficiency may be related to mother to child transmission of some infectious diseases 31 A very small number of viruses including rubella virus Zika virus and cytomegalovirus CMV can travel across the placental barrier generally taking advantage of conditions at certain gestational periods as the placenta develops CMV and Zika travel from the maternal bloodstream via placental cells to the fetal bloodstream 30 32 33 34 Beginning as early as 13 weeks of gestation and increasing linearly with the largest transfer occurring in the third trimester IgG antibodies can pass through the human placenta providing protection to the fetus in utero 35 36 This passive immunity lingers for several months after birth providing the newborn with a carbon copy of the mother s long term humoral immunity to see the infant through the crucial first months of extrauterine life IgM antibodies because of their larger size cannot cross the placenta 37 one reason why infections acquired during pregnancy can be particularly hazardous for the fetus 38 Endocrine function Edit The first hormone released by the placenta is called the human chorionic gonadotropin hCG hormone This is responsible for stopping the process at the end of menses when the corpus luteum ceases activity and atrophies If hCG did not interrupt this process it would lead to spontaneous abortion of the fetus The corpus luteum also produces and releases progesterone and estrogen and hCG stimulates it to increase the amount that it releases hCG is the indicator of pregnancy that pregnancy tests look for These tests will work when menses has not occurred or after implantation has happened on days seven to ten hCG may also have an anti antibody effect protecting it from being rejected by the mother s body hCG also assists the male fetus by stimulating the testes to produce testosterone which is the hormone needed to allow the sex organs of the male to grow Progesterone helps the embryo implant by assisting passage through the fallopian tubes It also affects the fallopian tubes and the uterus by stimulating an increase in secretions necessary for fetal nutrition Progesterone like hCG is necessary to prevent spontaneous abortion because it prevents contractions of the uterus and is necessary for implantation Estrogen is a crucial hormone in the process of proliferation This involves the enlargement of the breasts and uterus allowing for growth of the fetus and production of milk Estrogen is also responsible for increased blood supply towards the end of pregnancy through vasodilation The levels of estrogen during pregnancy can increase so that they are thirty times what a non pregnant woman mid cycles estrogen level would be Human placental lactogen hPL is a hormone used in pregnancy to develop fetal metabolism and general growth and development Human placental lactogen works with growth hormone to stimulate Insulin like growth factor production and regulating intermediary metabolism In the fetus hPL acts on lactogenic receptors to modulate embryonic development metabolism and stimulate production of IGF insulin surfactant and adrenocortical hormones hPL values increase with multiple pregnancies intact molar pregnancy diabetes and Rh incompatibility They are decreased with toxemia choriocarcinoma and Placental insufficiency 39 40 Immunological barrier Edit Further information Immune tolerance in pregnancy The placenta and fetus may be regarded as a foreign body inside the mother and must be protected from the normal immune response of the mother that would cause it to be rejected The placenta and fetus are thus treated as sites of immune privilege with immune tolerance For this purpose the placenta uses several mechanisms It secretes neurokinin B containing phosphocholine molecules This is the same mechanism used by parasitic nematodes to avoid detection by the immune system of their host 41 There is presence of small lymphocytic suppressor cells in the fetus that inhibit maternal cytotoxic T cells by inhibiting the response to interleukin 2 42 However the placental barrier is not the sole means of evading the immune system as foreign fetal cells also persist in the maternal circulation on the other side of the placental barrier 43 Other Edit The placenta also provides a reservoir of blood for the fetus delivering blood to it in case of hypotension and vice versa comparable to a capacitor 44 Ultrasound image of human placenta and umbilical cord color Doppler rendering with central cord insertion and three umbilical vessels at 20 weeks of pregnancyClinical significance EditMain article Placental disease Micrograph of a cytomegalovirus CMV infection of the placenta CMV placentitis The characteristic large nucleus of a CMV infected cell is seen off centre at the bottom right of the image H amp E stain Numerous pathologies can affect the placenta Placenta accreta when the placenta implants too deeply all the way to the actual muscle of uterine wall without penetrating it Placenta praevia when the placement of the placenta is too close to or blocks the cervix Placental abruption premature detachment of the placenta Placentitis inflammation of the placenta such as by TORCH infections Society and culture EditThe placenta often plays an important role in various cultures with many societies conducting rituals regarding its disposal In the Western world the placenta is most often incinerated 45 Some cultures bury the placenta for various reasons The Maori of New Zealand traditionally bury the placenta from a newborn child to emphasize the relationship between humans and the earth 46 Likewise the Navajo bury the placenta and umbilical cord at a specially chosen site 47 particularly if the baby dies during birth 48 In Cambodia and Costa Rica burial of the placenta is believed to protect and ensure the health of the baby and the mother 49 If a mother dies in childbirth the Aymara of Bolivia bury the placenta in a secret place so that the mother s spirit will not return to claim her baby s life 50 The placenta is believed by some communities to have power over the lives of the baby or its parents The Kwakiutl of British Columbia bury girls placentas to give the girl skill in digging clams and expose boys placentas to ravens to encourage future prophetic visions In Turkey the proper disposal of the placenta and umbilical cord is believed to promote devoutness in the child later in life In Transylvania and Japan interaction with a disposed placenta is thought to influence the parents future fertility citation needed Several cultures believe the placenta to be or have been alive often a relative of the baby Nepalese think of the placenta as a friend of the baby the orang Asli and Malay populations in Malay Peninsula regard it as the baby s older sibling 49 51 Native Hawaiians believe that the placenta is a part of the baby and traditionally plant it with a tree that can then grow alongside the child 45 Various cultures in Indonesia such as Javanese and Malay believe that the placenta has a spirit and needs to be buried outside the family house Some Malays would bury the baby s placenta with a pencil if it is a boy or a needle and thread if it is a girl 51 In some cultures the placenta is eaten a practice known as placentophagy In some eastern cultures such as China the dried placenta ziheche 紫河车 literally purple river car is thought to be a healthful restorative and is sometimes used in preparations of traditional Chinese medicine and various health products 52 The practice of human placentophagy has become a more recent trend in western cultures and is not without controversy its practice being considered cannibalism is debated Some cultures have alternative uses for placenta that include the manufacturing of cosmetics pharmaceuticals and food 53 Additional images Edit Fetus of about 8 weeks enclosed in the amnion Magnified a little over two diameters Placenta with attached fetal membranes ruptured at the margin at the left in the image Micrograph of a placental infection CMV placentitis Micrograph of CMV placentitis A 3D Power Doppler image of vasculature in 20 week placenta Schematic view of the placenta Maternal side of a whole human placenta just after birth Fetal side of same placenta Close up of umbilical attachment to fetal side of freshly delivered placenta Placenta weight by gestational age 54 Ziheche 紫河车 dried human placenta used in traditional Chinese medicineSee also EditChoriovitelline placenta Caul Zygote Pregnancy in fishReferences Edit Jin M Xu Q Li J Xu S Tang C September 2022 Micro RNAs in Human Placenta Tiny Molecules Immense Power Molecules 27 18 5943 doi 10 3390 molecules27185943 PMC 9501247 PMID 36144676 Pough F H Andrews Robin M Cadle John E Crump Martha L Savitsky Alan H Wells Kentwood D 2004 Herpetology 3rd ed Pearson ISBN 978 0 13 100849 6 page needed Mitra Avir 31 January 2020 How the placenta evolved from an ancient virus WHYY Retrieved 9 March 2020 Chuong Edward B 9 October 2018 The placenta goes viral Retroviruses control gene expression in pregnancy PLOS Biology 16 10 e3000028 doi 10 1371 journal pbio 3000028 PMC 6177113 PMID 30300353 Villarreal Luis P January 2016 Viruses and the placenta the essential virus first view APMIS 124 1 2 20 30 doi 10 1111 apm 12485 PMID 26818259 S2CID 12042851 Henry George Liddell Robert Scott A Greek English Lexicon at Perseus Archived 2012 04 05 at the Wayback Machine placenta Archived 2016 01 30 at the Wayback Machine Online Etymology Dictionary Bowen R Implantation and Development of the Placenta Introduction and Index From Pathophysiology of the Reproductive System Accessed 7 July 2019 Examination of the placenta Archived 2011 10 16 at the Wayback Machine dead link Placental Structure and Classification Archived 2016 02 11 at the Wayback Machine Fujikura Toshio Benson Ralph C Driscoll Shirley G et al 1970 The bipartite placenta and its clinical features American Journal of Obstetrics and Gynecology 107 7 1013 1017 doi 10 1016 0002 9378 70 90621 6 PMID 5429965 Bipartite placenta represented 4 2 per cent 366 of 8 505 of placentas of white women at the Boston Hospital for Women who were enrolled in the Collaborative Project The human proteome in placenta The Human Protein Atlas www proteinatlas org Archived from the original on 2017 09 26 Retrieved 2017 09 26 Uhlen Mathias Fagerberg Linn Hallstrom Bjorn M Lindskog Cecilia Oksvold Per Mardinoglu Adil Sivertsson Asa Kampf Caroline Sjostedt Evelina 2015 01 23 Tissue based map of the human proteome Science 347 6220 1260419 doi 10 1126 science 1260419 ISSN 0036 8075 PMID 25613900 S2CID 802377 a b Stages of Development of the Fetus Women s Health Issues Merck Manuals Consumer Version Retrieved 2022 06 12 How Your Fetus Grows During Pregnancy www acog org Retrieved 2022 06 12 Dashe Jodi S Bloom Steven L Spong Catherine Y Hoffman Barbara L 2018 Williams Obstetrics McGraw Hill Professional ISBN 978 1 259 64433 7 page needed Placental blood circulation Archived 2011 09 28 at the Wayback Machine a b Kiserud T Acharya G 2004 The fetal circulation Prenatal Diagnosis 24 13 1049 1059 doi 10 1002 pd 1062 PMID 15614842 S2CID 25040285 a b Reiter R J Tan D X Korkmaz A Rosales Corral S A 2013 Melatonin and stable circadian rhythms optimize maternal placental and fetal physiology Human Reproduction Update 20 2 293 307 doi 10 1093 humupd dmt054 ISSN 1355 4786 PMID 24132226 Williams book of obsteritcis Begley CM Gyte GM Devane D McGuire W Weeks A Biesty LM 13 February 2019 Active versus expectant management for women in the third stage of labour Cochrane Database of Systematic Reviews 2019 2 CD007412 doi 10 1002 14651858 CD007412 pub5 PMC 6372362 PMID 30754073 Mercer JS Vohr BR Erickson Owens DA Padbury JF Oh W 2010 Seven month developmental outcomes of very low birth weight infants enrolled in a randomized controlled trial of delayed versus immediate cord clamping Journal of Perinatology 30 1 11 6 doi 10 1038 jp 2009 170 PMC 2799542 PMID 19847185 Perez Munoz Maria Elisa Arrieta Marie Claire Ramer Tait Amanda E Walter Jens 2017 A critical assessment of the sterile womb and in utero colonization hypotheses implications for research on the pioneer infant icrobiome Microbiome 5 1 48 doi 10 1186 s40168 017 0268 4 ISSN 2049 2618 PMC 5410102 PMID 28454555 Mor Gil Kwon Ja Young 2015 Trophoblast microbiome interaction a new paradigm on immune regulation American Journal of Obstetrics and Gynecology 213 4 S131 S137 doi 10 1016 j ajog 2015 06 039 ISSN 0002 9378 PMC 6800181 PMID 26428492 Prince Amanda L Antony Kathleen M Chu Derrick M Aagaard Kjersti M 2014 The microbiome parturition and timing of birth more questions than answers Journal of Reproductive Immunology 104 105 12 19 doi 10 1016 j jri 2014 03 006 ISSN 0165 0378 PMC 4157949 PMID 24793619 Hornef M Penders J 2017 Does a prenatal bacterial microbiota exist Mucosal Immunology 10 3 598 601 doi 10 1038 mi 2016 141 PMID 28120852 Wright C Sibley CP 2011 Placental Transfer in Health and Disease In Kay H Nelson M Yuping W eds The Placenta From Development to Disease John Wiley and Sons pp 66 ISBN 978 1 4443 3366 4 Perazzolo S Hirschmugl B Wadsack C Desoye G Lewis RM Sengers BG February 2017 The influence of placental metabolism on fatty acid transfer to the fetus J Lipid Res 58 2 443 454 doi 10 1194 jlr P072355 PMC 5282960 PMID 27913585 Kappen C Kruger C MacGowan J Salbaum JM 2012 Maternal diet modulates placenta growth and gene expression in a mouse model of diabetic pregnancy PLOS ONE 7 6 e38445 Bibcode 2012PLoSO 738445K doi 10 1371 journal pone 0038445 PMC 3372526 PMID 22701643 a b Madhusoodanan Jyoti October 10 2018 Break on through How some viruses infect the placenta Knowable Magazine doi 10 1146 knowable 101018 1 Erlebacher Adrian 2013 03 21 Immunology of the Maternal Fetal Interface Annual Review of Immunology 31 1 387 411 doi 10 1146 annurev immunol 032712 100003 PMID 23298207 Retrieved 25 June 2021 Pereira Lenore 2018 09 29 Congenital Viral Infection Traversing the Uterine Placental Interface Annual Review of Virology 5 1 273 299 doi 10 1146 annurev virology 092917 043236 PMID 30048217 S2CID 51724379 Retrieved 24 June 2021 Arora Nitin Sadovsky Yoel Dermody Terence S Coyne Carolyn B May 2017 Microbial Vertical Transmission during Human Pregnancy Cell Host amp Microbe 21 5 561 567 doi 10 1016 j chom 2017 04 007 PMC 6148370 PMID 28494237 Robbins Jennifer R Bakardjiev Anna I February 2012 Pathogens and the placental fortress Current Opinion in Microbiology 15 1 36 43 doi 10 1016 j mib 2011 11 006 PMC 3265690 PMID 22169833 Palmeira Patricia Quinello Camila Silveira Lessa Ana Lucia Zago Claudia Augusta Carneiro Sampaio Magda 2012 IgG Placental Transfer in Healthy and Pathological Pregnancies Clinical and Developmental Immunology 2012 985646 doi 10 1155 2012 985646 PMC 3251916 PMID 22235228 Simister N E Story C M 1997 Human placental Fc receptors and the transmission of antibodies from mother to fetus Journal of Reproductive Immunology 37 1 1 23 doi 10 1016 s0165 0378 97 00068 5 PMID 9501287 Pillitteri Adele 2009 Maternal and Child Health Nursing Care of the Childbearing and Childrearing Family Hagerstwon MD Lippincott Williams amp Wilkins p 202 ISBN 978 1 58255 999 5 What infections can affect pregnancy National Institute of Child Health and Human Development Retrieved 25 June 2021 Handwerger S Freemark M 2000 The roles of placental growth hormone and placental lactogen in the regulation of human fetal growth and development Journal of Pediatric Endocrinology amp Metabolism 13 4 343 56 doi 10 1515 jpem 2000 13 4 343 PMID 10776988 S2CID 28778529 Human Placental Lactogen www ucsfhealth org May 17 2009 Archived from the original on April 29 2017 Retrieved July 21 2017 Placenta fools body s defences BBC News 10 November 2007 Archived from the original on 29 April 2012 Clark DA Chaput A Tutton D March 1986 Active suppression of host vs graft reaction in pregnant mice VII Spontaneous abortion of allogeneic CBA J x DBA 2 fetuses in the uterus of CBA J mice correlates with deficient non T suppressor cell activity J Immunol 136 5 1668 75 PMID 2936806 Williams Z Zepf D Longtine J Anchan R Broadman B Missmer SA Hornstein MD March 2008 Foreign fetal cells persist in the maternal circulation Fertil Steril 91 6 2593 5 doi 10 1016 j fertnstert 2008 02 008 PMID 18384774 Assad RS Lee FY Hanley FL 2001 Placental compliance during fetal extracorporeal circulation Journal of Applied Physiology 90 5 1882 1886 doi 10 1152 jappl 2001 90 5 1882 PMID 11299282 S2CID 8785947 a b Why eat a placenta BBC News 18 April 2006 Metge Joan 2005 Working In Playing With Three Languages Sites A Journal of Social Anthropology and Cultural Studies 2 2 83 90 doi 10 11157 sites vol2iss2id65 Francisco Edna 3 December 2004 Bridging the Cultural Divide in Medicine Science Shepardson Mary 1978 Changes in Navajo mortuary practices and beliefs American Indian Quarterly 4 4 383 96 doi 10 2307 1184564 JSTOR 1184564 PMID 11614175 a b Buckley Sarah J Placenta Rituals and Folklore from around the World Mothering Archived from the original on 6 January 2008 Retrieved 7 January 2008 Davenport Ann June 2005 The Love Offer Johns Hopkins Magazine 57 3 a b Anisah Barakbah 2017 Ensiklopedia Perbidanan Melayu Universiti Islam Malaysia Press pp 236 237 ISBN 978 967 13305 9 3 Falcao Ronnie Medicinal Uses of the Placenta Archived from the original on 5 December 2008 Retrieved 25 November 2008 Krolokke Charlotte Dickinson Elizabeth Foss Karen A May 2018 The placenta economy From trashed to treasured bio products European Journal of Women s Studies 25 2 138 153 doi 10 1177 1350506816679004 ISSN 1350 5068 S2CID 151874106 Placental Weights Means Standard Deviations and Percentiles by Gestational Age Placental and Gestational Pathology 2017 p 336 doi 10 1017 9781316848616 039 ISBN 978 1 316 84861 6 External links Edit Wikimedia Commons has media related to Placenta Look up placenta in Wiktionary the free dictionary The placenta specific proteome at the Human Protein Atlas The Placenta gynob com with quotes from Williams Obstetrics 18th Edition F Gary Cunningham M D Paul C MacDonald M D Norman F Grant M D Appleton amp Lange Publishers Retrieved from https en wikipedia org w index php title Placenta amp oldid 1136321358, wikipedia, wiki, book, books, library,

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