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The trophoblast (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
: to feed; and : germinator) is the outer layer of cells of the
blastocyst The blastocyst is a structure formed in the early embryonic development of mammals. It possesses an inner cell mass (ICM) also known as the ''embryoblast'' which subsequently forms the embryo, and an outer layer of trophoblast cells called the ...
. Trophoblasts are present four days after
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proc ...
in humans. They provide nutrients to the
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
and develop into a large part of the
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 mat ...
. They form during the first stage of
pregnancy Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins. Pregnancy usually occurs by sexual intercourse, but ...
and are the first cells to differentiate from the
fertilized Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proc ...
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point t ...
to become extraembryonic structures that do not directly contribute to the embryo. After
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrul ...
, the trophoblast is contiguous with the
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from th ...
of the embryo and is referred to as the trophectoderm. After the first differentiation, the cells in the human embryo lose their
totipotency Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...
and are no longer
totipotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...
stem cells because they cannot form a trophoblast. They are now
pluripotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...
stem cells.


Structure

The trophoblast proliferates and differentiates into two cell layers at approximately six days after fertilization for humans.


Function

Trophoblasts are specialized cells of the
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 mat ...
that play an important role in embryo implantation and interaction with the
decidua The decidua is the modified mucosal lining of the uterus (that is, modified endometrium) that forms every month, in preparation for pregnancy. It is shed off each month when there is no fertilised egg to support. The decidua is under the infl ...
lized maternal
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. Th ...
. The core of placental villi contain mesenchymal cells and placental blood vessels that are directly connected to the fetus’ circulation via the
umbilical cord In placental mammals, the umbilical cord (also called the navel string, birth cord or ''funiculus umbilicalis'') is a conduit between the developing embryo or fetus and the placenta. During prenatal development, the umbilical cord is physiol ...
. This core is surrounded by two layers of trophoblasts, the
cytotrophoblast "Cytotrophoblast" is the name given to both the inner layer of the trophoblast (also called layer of Langhans) or the cells that live there. It is interior to the syncytiotrophoblast and external to the wall of the blastocyst in a developing embr ...
and the
syncytiotrophoblast Syncytiotrophoblast (from the Greek 'syn'- "together"; 'cytio'- "of cells"; 'tropho'- "nutrition"; 'blast'- "bud") is the epithelial covering of the highly vascular embryonic placental villi, which invades the wall of the uterus to establish ...
. The cytotrophoblast is a layer of mono-nucleated cells that resides underneath the syncytiotrophoblast. The syncytiotrophoblast is composed of fused cytotrophoblasts which then form a layer that covers the placental surface. The syncytiotrophoblast is in direct contact with the maternal blood that reaches the placental surface. It then facilitates the exchange of nutrients, wastes and gases between the maternal and fetal systems. In addition, cytotrophoblasts in the tips of villi can differentiate into another type of trophoblast called the
extravillous trophoblast Extravillous trophoblasts (EVTs), are one form of differentiated trophoblast cells of the placenta. They are invasive mesenchymal cells which function to establish critical tissue connection in the developing placental-uterine interface. EVTs der ...
. Extravillous trophoblasts grow out from the placenta and penetrate into the decidualized uterus. This process is essential not only for physically attaching the placenta to the mother, but also for altering the vasculature in the uterus. This alteration allows an adequate blood supply to the growing fetus as pregnancy progresses. Some of these trophoblasts even replace the
endothelial cells The endothelium is a single layer of squamous endothelial cells that line the interior surface of blood vessels and lymphatic vessels. The endothelium forms an interface between circulating blood or lymph in the lumen and the rest of the ve ...
in the uterine
spiral arteries Spiral arteries are small arteries which temporarily supply blood to the endometrium of the uterus during the luteal phase of the menstrual cycle. In histology, identifying the presence of these arteries is one of the most useful techniques in ...
as they remodel these vessels into wide bore conduits that are independent of maternal
vasoconstriction Vasoconstriction is the narrowing of the blood vessels resulting from contraction of the muscular wall of the vessels, in particular the large arteries and small arterioles. The process is the opposite of vasodilation, the widening of blood vesse ...
. This ensures that the fetus receives a steady supply of blood, and the placenta is not subjected to fluctuations in oxygen that could cause it damage.


Clinical significance

The invasion of a specific type of trophoblast (extravillous trophoblast) into the maternal
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. Th ...
is a vital stage in the establishment of
pregnancy Pregnancy is the time during which one or more offspring develops ( gestates) inside a woman's uterus (womb). A multiple pregnancy involves more than one offspring, such as with twins. Pregnancy usually occurs by sexual intercourse, but ...
. Failure of the trophoblast to invade sufficiently is important in the development of some cases of
pre-eclampsia Pre-eclampsia is a disorder of pregnancy characterized by the onset of high blood pressure and often a significant amount of protein in the urine. When it arises, the condition begins after 20 weeks of pregnancy. In severe cases of the disease ...
. Invasion of the trophoblast too deeply may cause conditions such as
placenta accreta Placenta accreta occurs when all or part of the placenta attaches abnormally to the '' myometrium'' (the muscular layer of the uterine wall). Three grades of abnormal placental attachment are defined according to the depth of attachment and invas ...
, placenta increta, or placenta percreta.
Gestational trophoblastic disease Gestational trophoblastic disease (GTD) is a term used for a group of pregnancy-related tumours. These tumours are rare, and they appear when cells in the womb start to proliferate uncontrollably. The cells that form gestational trophoblastic tu ...
is a pregnancy-associated concept, forming from the villous and extravillous trophoblast cells in the placenta.
Choriocarcinoma Choriocarcinoma is a malignant, trophoblastic cancer, usually of the placenta. It is characterized by early hematogenous spread to the lungs. It belongs to the malignant end of the spectrum in gestational trophoblastic disease (GTD). It is also ...
are trophoblastic tumors that form in the uterus from villous cells. Trophoblast stem cells (TSCs) are cells that can regenerate and they are similar to
embryonic stem cell Embryonic stem cells (ESCs) are pluripotent stem cells derived from the inner cell mass of a blastocyst, an early-stage pre- implantation embryo. Human embryos reach the blastocyst stage 4–5 days post fertilization, at which time they consis ...
s (ESCs) in the fact that they come from early on in the trophoblast lifetime. In the
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 mat ...
, these stem cells are able to differentiate into any trophoblast cell because they are pluripotent.


Additional images

File:Gray10.png, Blastodermic
vesicle Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features ...
of ''Vespertilio murinus''. File:Gray11.png, Section through embryonic disk of ''Vespertilio murinus''. File:Gray35.png, Transverse section of a chorionic villus. File:Gray39.png, Scheme of placental circulation. File:HumanEmbryogenesis.svg, The initial stages of
human embryogenesis Human embryonic development, or human embryogenesis, is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of ...
File:Histopathology of tubal pregnancy.jpg, Histopathology of a chorionic villus, in a tubal pregnancy, with labeled cytotrophoblasts and syncytiotrophoblasts.


See also

*
Syncytiotrophoblast Syncytiotrophoblast (from the Greek 'syn'- "together"; 'cytio'- "of cells"; 'tropho'- "nutrition"; 'blast'- "bud") is the epithelial covering of the highly vascular embryonic placental villi, which invades the wall of the uterus to establish ...
* Hydatidiform mole


References


External links

* {{Authority control Animal developmental biology