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ISSN 2358-2731/BJBS-2015-0139/2/4/3/199
Braz. J. Biol. Sci.
http://revista.rebibio.net
200 Bertasoli et al.
anaccessory structure of the placenta, which in mammals, the blood vessels within it
may be found in ungulates. It has a dome- carry blood between the embryo and the
shaped, coated with trophoblast cells placenta. The chorion is the outermost
absorbing situated opposite the openings of extraembryonic membrane in birds, reptiles
the uterine glands (Mossman, 1987). and mammals and is involved in respiratory
All the placenta of mammals gas exchange. Exceptionally in mammals, it
increases the contact area between the is part of the placenta and is additionally
maternal and fetal surfaces by intense involved with nutrition and waste disposal,
interdigitation of tissue. When this once itis a thin film that surrounds the
distribution extends in the major surface embryo and other extraembryonic
chorionic (Leiser and Kaufmann, 1994), or membranes (Wolpert et al., 2000).
when the villi are distributed regularly by The yolk sac develops as an
chorion, there are then the diffuse placenta attached structure of the embryonic midgut
(pigs and horses). and consists of a layer of endodermal
The placenta is a structurally epithelium followed by a vascularized fetal
complex organ. This complexity is mesenchyme (Leiser and Kaufmann, 1994).
promoted in part by the interaction of origin The chorionic epithelium is the
maternal and fetal tissues, and the presence decisive barrier to maternal-fetal exchanges
of a variety of intermediate layers that are (Leiser and Kaufmann, 1994). During
interposed between the maternal and fetal deployment, the internal layer of
vascular space (Schwarze, 1970). mesenchymal tissue derived from embryois
Therefore, we aimed to conduct a able to fulfill the chorion, soon after the
literature review in order to describe the mesenchyme develops its own
porcine placenta and its extraembryonic vascularization system. The cavity
annexes, providing detailed data for future enwrapped by the chorion is denominated
research in this area. exoceloma and contains the embryo, the
amnion, the allantoisand the yolk sac
Extraembryonic structures: chorion, (Assis-Neto et al., 2012; Mançanares et al.,
amnion, allantois and yolk sac 2013). In order to be responsible for
maternal-fetal exchange, the chorion must
The extraembryonic structures are present a functional circulation. However, it
the chorion, the amnion, the allantois and is an avascular tissue and vascularization is
the yolk sac (Figure 1). These embryonic provided by allantois vessels exclusively
adnexa are different membranous structures (Kaufmann and Burton, 1994).
and are deeply involved in placental The allantois corresponds to the
development (Bertassoli et al., 2012; extraembryonic urinary bladder developed
Bertassoli et al., 2015). These membranes, from the embryonic intestine andformed
however, undergo morphological changes, with simple layer of squamous cells. These
interacting, forming connections and cells are sustained by a thin basement
functional complexes or even involution in membrane and extraembryonic
the course of pregnancy. mesenchyme (Bjorkman, 1986; Assis-Neto
The amnion is a thin membrane that et al., 2012).
delimits a bag fulfilledwithfluid - the The amniotic membrane is
amniotic fluid, which protects the embryo responsible for fetus hydration and
against mechanical shocks. This cavity nutrition, lubrication of the birth canal and
derives from the division (rodents and mechanical protection (Leiser and
primates) or the folding (mammals in Kaufmann, 1994). The amnion is
general) of the embryonic ectoderm (Leiser composedby epithelial cells that show
and Kaufmann, 1994). uniform morphology and did not differ
The allantois arises from the among mammals (Steven, 1982). It is
invagination of the posterior part of the squamous, continuous and organized,
embryo’s gut. It acts with a respiratory forming a membrane similar to the allantoic
surface in bird and reptile embryos, while, epithelium, being also supported by a layer
Figure 2. Swine placenta. (A) Embryonic attachment in swine; (B) Blood flow: cross current;
(C) Membrane inter-hematic: epitheliochorial type. As described by Roa et al. (2012) and Montiel et al.
(2013).
Figure 3. Schema of swine placenta and uterine wall section showing areola as described by Olio et al.
(2014).
often has more than one aperture at this involution of the yolk sac during early gestation
place. bovine (Bos indicus). Acta Scientiae
Veterinariae, v. 40, n. 4, p. 1-9, 2012.
Final considerations Bazer, F. W. Uterine protein secretions:
relationship to development of the conceptus.
The study of the placenta and the Journal of Animal Science, v. 41, n. 5, p. 1376-
extraembryonic tissues in swine is of 1382, 1975.
upmost importance once pigs are relevant Bazer, F. W.; Vallet, J. L.; Roberts, R. M.;
animals for livestock. Based on the Sharp, D. C.; Thatcher, W. W. Role of conceptus
secretory products in establishment of
literature, as referred by many authors here
pregnancy. Journal of Reproduction and
cited, the placenta is vitally important Fertility, v. 76, n. 2, p. 841-850, 1986.
according as one of its functions is nutrient
Beaudoin, S.; Simon, L.; Simeoni, J.; Sacquin,
exchange. The swine placenta is therefore
P.; Bargy, F. Surgical approach of an early
classified as chorioallantoic, diffuse, mammalian embryo: the rabbit model. Fetal
pleated, epitheliochorial and cross to Diagnosis and Therapy, v. 13, n. 2, p. 82-85,
counter-current. Amnion, chorion, allantois 1998.
and yolk sacare intimately related to fetal Bertassoli, B. M.; Oliveira, F. D.; Santos, A. C.;
development and embryonic losses, since Miglino, M. A.; Assis-Neto, A. C. Expressões
the placental development in pigs starts gênicas nas membranas extraembrionárias e
around the 18th day of gestation. suas conexões na espécie de murina e bovina.
Modifications in the swine placenta, like Sabios: Revista de Saúde e Biologia, v. 7, n. 2,
the presence of areolas, might have arisen p. 90-101, 2012.
as domestic species adaptions in order to Bertassoli, B. M.; Santos, A. C.; Fratini, P.; Will,
supply nourishing needs during the S. E. A. L.; Assis-Neto, A. C. Morphological
development of concept. analysis of the extra-embryonic membranes of
domestic pig (Sus scrofa) at 20 days of
Acknowledgements gestation. Journal of Basic and Applied
Research International, v. 10, n. 1, p. 14-20,
2015.
Thanks to FAPESP (Fundação de
Amparo a Pesquisa do Estado de São Bjorkman, N. Placentation. In: Dellman, H. D.;
Brown, E. M. (Eds.). Textbook of veterinary
Paulo) for Financial Support.
histology. Philadelphia: Lea & Febger, 1986. p.
351-359.
Conflict of interest statement
Chen, T. T.; Bazer, F. W.; Gebhardt, B. M.;
Roberts, R. M. Uterine secretion in mammals:
Authors declare that they have no synthesis and placental transport of a purple
conflict of interests. acid phosphatase in pig. Biology of
Reproduction, v. 13, n. 2, p. 304-313, 1975.
References Cummins, C.; Carrington, S.; Fitzpatrick, E.;
Duggan, V. Ascending placentitis in the mare: a
Abd-Elnaeim, M. M. M.; Leiser, R.; Allen, review. Irish Veterinary Journal, v. 61, n. 5,
W. R. Structural and haematological aspects of p. 307-313, 2008.
the equine placenta in mid-pregnancy.
Dantzer, V.; Leiser, R. Microvasculature of
Havemeyer Foundation Monograph Series,
regular and irregular areolae of the areola-gland
v. 10, n. 1, p. 39-42, 2003.
subunit of the porcine placenta: structural and
Abromavich, C. The morphology and functional aspects. Anatomy and Embryology,
distribution of the rosettes on the foetal placenta v. 188, n. 2, p. 257-267, 1993.
of the pig. Anatomical Records, v. 33, n. 2,
Drieux, H.; Thiery, G. Placenttion chez les
p. 67-72, 1926.
mammiferes domestiques. II. Placenta dês
Amoroso, E. C. Placentation. In: Parkes, A. S. suidés. Recueil de Médecine Vétérinaire,
(Ed.). Marshall’s physiology of reproduction. v. 125, n. 10, p. 437-455, 1949.
London: Green and Co., 1952. v. 2. p. 127-311.
Friess, A. E.; Sinowatz, F.; Skolek-Winnisch,
Assis-Neto, A. C.; Oliveira, F. D.; Constantino, R.; Traütner, W. The placenta of the pig. II. The
M. V. P.; Miglino, M. A. Morphology and
ultrastructure of the areola. Anatomy and Oliveira, G. B.; Vale, A. M.; Santos, A. C.;
Embryology, v. 163, n. 1, p. 43-53, 1981. Moura, C. E. B.; Rocha, H. A. O.; Oliveira, M.
Haar, J. L.; Ackerman, G. A. Phase and electron F. Composition and significance of
microscopic study of vasculogenesis and glycosaminoglycans in the uterus and placenta
erythropoieses in the yolk sac of the mouse. The of mammals. Brazilian Archives of Biology
Anatomicical Records, v. 170, n. 2, p. 199-224, and Technology, v. 58, n. 4, p. 512-520, 2015.
1971. Available from: <http://www.scielo.br/pdf/babt/
v58n4/1516-8913-babt-201500281.pdf>.
Hafez, E. S. S. E.; Hafez, B. Reprodução
Accessed in: Aug. 22, 2015.
animal. 7. ed. São Paulo: Manole, 2004.
Perry, J. P. The mammalian fetal membranes.
Hitzig, W. H. Über die entwicklung der
Journal of Reproduction and Fertility, v. 62,
scweineplacenta. Acta Anatomica, v. 7, n. 1-2,
n. 2, p. 321-335, 1981.
p. 33-81, 1949.
Raub, T. J.; Bazer, F. W.; Roberts, R. M.
Kauffmann, P.; Burton, G. Anatomy and genesis
Localization of the iron transport glycoprotein,
of the placenta. In: Knobil, E.; Neill, J. D. The
uteroferrin, in the porcine endometrium and
physiology of reproduction. New York: Raven
placenta by using immunocolloidal gold.
Press, 1994. p. 441-484.
Anatomy and Embryology, v. 171, n. 2,
Leiser, R.; Kauffmann. P. Placental structure: in p. 253-258, 1985.
a comparative aspect. Experimental and
Roa, I.; Smok, C. S.; Pietro, G. R. Placenta:
Clinical Endocrinology, v. 102, n. 3, p. 122-
anatomia e histologia comparada. International
134, 1994.
Journal of Morphology, v. 30, n. 4, p. 1490-
Mançanares, C. A. F.; Leiser, R.; Favaron, P. O.; 1496, 2012.
Carvalho, A. F.; Oliveira, V. C.; Santos, J. M.;
Santos, A. C.; Bertassoli, B. M.; Oliveira, F. D.;
Miglino, M. A.; Ambrósio, C. E. A
Assis-Neto, A. C.; Miglino, M. A. Circulação
Morphological analysis of the transition
vitelina: análise comparativa. Revista
between the embryonic primitive intestine and
Científica Eletrônica de Medicina
yolk sac in bovine embryos and fetuses.
Veterinária, v. 18, n. 2, p. 1-21, 2012.
Microscopy Research and Techniques, v. 76,
<http://faef.revista.inf.br/imagens_arquivos/arqu
n. 7, p. 756-766, 2013.
ivos_destaque/3NMxF5f36DMzZIw_2013-6-
Miglino, M. A.; Pereira, F. T. V.; Santos, T. C.; 24-17-5-40.pdf>. Accessed in: Aug. 22, 2015.
Carvalho, A. F. A morfologia placentária dos
Schwarze, E. Compêndio de anatomia
suínos domésticos. Arquivos de Ciências
veterinária: embriologia. Zaragoza: Acribia,
Veterinárias e Zoologia da UNIPAR, v. 4, n. 1,
1970.
p. 71-76, 2001.
Schlafer, D. H.; Fisher, P. J.; Davies, C. J. The
Montiel, J. F.; Kaune, H.; Maliqueo, M.
bovine placenta before and after birth: placental
Maternal-fetal unit interactions and eutherian
development and function in health and disease.
neocortical development and evolution.
Animal Reproduction Science, v. 61, n. 60,
Frontiers in Neuroanatomy, v. 7, n. 22, p. 1-
p. 145-160, 2000.
14, 2013. Available from:
<http://journal.frontiersin.org/article/10.3389/fn Schlosnagle, D. C.; Bazer, F. W.; Tsibris,
ana.2013.00022/>. Accessed in: Aug. 22, 2015. J. C. M.; Roberts, R. M. An iron-containing
phosphatase induced by progesterone in the
Mossman, H. W. Comparative morphogenesis
uterine fluids of pigs. The Journal of
of the fetal membranes and accessory uterine
Biological Chemistry, v. 249, n. 23, p. 7574-
structures. Contributions in Embriology, v. 26,
7579, 1974.
n. 158, p. 133-247, 1937.
Steven, D. H. Placentation in mare. Journal of
Mossman, H. W. Vertebrate fetal membranes.
Reproduction and Fertility, v. 31, sup., p. 41-
New Brunswick: Rutgers University Press,
55, 1982.
1987.
Tommasi-Jr, H. L. P.; Santos, A. C.; Miglino,
Olio, R. L.; Lobo, L. M.; Pereira, M. A.; Santos,
A. C.; Assis-Neto, A. C. The origin of
A. C.; Viana, D. C.; Favaron, P. O.; Miglino,
hematopoiesis and vasculogenesis in the yolk
M. A. Accessory placental structures - a review.
sac and the onset of intraembryonic blood flow
Open Journal of Animal Sciences, v. 4, n. 5, p.
and cardiac function in mammalians. Revista
305-312, 2014. Available from:
Científica Eletrônica de Medicina
<http://www.scirp.org/journal/PaperInformation
Veterinária, v. 19, n. 1, p. 1-15, 2012. Available
.aspx?PaperID=50831>. Accessed in: Aug. 22,
from: <http://faef.revista.inf.br/
2015.
imagens_arquivos/arquivos_destaque/EOcEFc
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