Human embryonic development

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
development. In biological terms, the
development of the human body entails
growth from a one-celled zygote to an
adult human being. Fertilization occurs
when the sperm cell successfully enters
and fuses with an egg cell (ovum). The
genetic material of the sperm and egg
then combine to form the single cell
zygote and the germinal stage of
development commences. Embryonic
development in the human, covers the The initial stages of human embryonic development
first eight weeks of development; at the (embryogenesis)
beginning of the ninth week the
embryo is termed a fetus. The eight
weeks has 23 stages.

Human embryology is the study of this development during the first eight weeks after fertilization. The
normal period of gestation (pregnancy) is about nine months or 40 weeks.

The germinal stage refers to the time from fertilization through the development of the early embryo until
implantation is completed in the uterus. The germinal stage takes around 10 days.[1] During this stage, the
zygote begins to divide, in a process called cleavage. A blastocyst is then formed and implants in the uterus.
Embryogenesis continues with the next stage of gastrulation, when the three germ layers of the embryo
form in a process called histogenesis, and the processes of neurulation and organogenesis follow.

In comparison to the embryo, the fetus has more recognizable external features and a more complete set of
developing organs. The entire process of embryogenesis involves coordinated spatial and temporal changes
in gene expression, cell growth and cellular differentiation. A nearly identical process occurs in other
species, especially among chordates.

Germinal stage

Fertilization

Fertilization takes place when the spermatozoon has successfully entered the ovum and the two sets of
genetic material carried by the gametes fuse together, resulting in the zygote (a single diploid cell). This
usually takes place in the ampulla of one of the fallopian tubes. The zygote contains the combined genetic
material carried by both the male and female gametes which consists of the 23 chromosomes from the
nucleus of the ovum and the 23 chromosomes from the nucleus of the sperm. The 46 chromosomes
undergo changes prior to the mitotic division which leads to the formation of the embryo having two cells.

Successful fertilization is enabled by three processes, which also act as controls to ensure species-specificity.
The first is that of chemotaxis which directs the movement of the sperm towards the ovum.[2] Secondly, an
adhesive compatibility between the sperm and the egg occurs. With the sperm adhered to the ovum, the
third process of acrosomal reaction takes place; the front part of the spermatozoan head is capped by an
acrosome which contains digestive enzymes to break down the zona pellucida and allow its entry.[3] The
entry of the sperm causes calcium to be released which blocks entry to other sperm cells.[3] A parallel
reaction takes place in the ovum called the zona reaction. This sees the release of cortical granules that
release enzymes which digest sperm receptor proteins, thus preventing polyspermy.[4] The granules also
fuse with the plasma membrane and modify the zona pellucida in such a way as to prevent further sperm
entry.

Cleavage

The beginning of the cleavage process is marked when the zygote

divides through mitosis into two cells. This mitosis continues and the
first two cells divide into four cells, then into eight cells and so on.
Each division takes from 12 to 24 hours. The zygote is large compared
to any other cell and undergoes cleavage without any overall increase
in size. This means that with each successive subdivision, the ratio of
nuclear to cytoplasmic material increases.[5]

Initially, the dividing cells, called blastomeres (blastos Greek for Eight-cell embryo, at three days
sprout), are undifferentiated and aggregated into a sphere enclosed
within the zona pellucida of the ovum. When eight blastomeres have
formed, they start to compact.[6] They begin to develop gap junctions, enabling them to develop in an
integrated way and co-ordinate their response to physiological signals and environmental cues.[7]

When the cells number around sixteen, the solid sphere of cells within the zona pellucida is referred to as a
morula.[8]

Blastulation

Cleavage itself is the first stage in blastulation, the process of forming

the blastocyst. Cells differentiate into an outer layer of cells called the
trophoblast, and an inner cell mass. With further compaction the
individual outer blastomeres, the trophoblasts, become
indistinguishable. They are still enclosed within the zona pellucida.
This compaction serves to make the structure watertight, containing the
fluid that the cells will later secrete. The inner mass of cells
differentiate to become embryoblasts and polarise at one end. They
close together and form gap junctions, which facilitate cellular Blastocyst with an inner cell
communication. This polarisation leaves a cavity, the blastocoel, mass and trophoblast
creating a structure that is now termed the blastocyst. (In animals other
than mammals, this is called the blastula).
The trophoblasts secrete fluid into the blastocoel. The resulting increase in size of the blastocyst causes it to
hatch through the zona pellucida, which then disintegrates.[5] This process is called zona hatching and it
takes place on the sixth day of embryo development, immediately before the implantation process. The
hatching of the human embryo is supported by proteases secreted by the cells of the blastocyst, which digest
proteins of the zona pellucida, giving rise to a hole. Then, due to the rhythmic expansion and contractions
of the blastocyst, an increase of the pressure inside the blastocyst itself occurs, the hole expands and finally
the blastocyst can emerge from this rigid envelope.

The inner cell mass will give rise to the pre-embryo,[9] the amnion, yolk sac and allantois, while the fetal
part of the placenta will form from the outer trophoblast layer. The embryo plus its membranes is called the
conceptus, and by this stage the conceptus has reached the uterus. The zona pellucida ultimately disappears
completely, and the now exposed cells of the trophoblast allow the blastocyst to attach itself to the
endometrium, where it will implant. The formation of the hypoblast and epiblast, which are the two main
layers of the bilaminar germ disc, occurs at the beginning of the second week.[10] Both the embryoblast and
the trophoblast will turn into two sub-layers.[11] The inner cells will turn into the hypoblast layer, which will
surround the other layer, called the epiblast, and these layers will form the embryonic disc that will develop
into the embryo.[10][11]

The trophoblast will also develop two sub-layers: the cytotrophoblast, which is in front of the
syncytiotrophoblast, which in turn lies within the endometrium.[10] Next, another layer called the
exocoelomic membrane or Heuser's membrane will appear and surround the cytotrophoblast, as well as the
primitive yolk sac.[11] The syncytiotrophoblast will grow and will enter a phase called lacunar stage, in
which some vacuoles will appear and be filled by blood in the following days.[10][11] The development of
the yolk sac starts with the hypoblastic flat cells that form the exocoelomic membrane, which will coat the
inner part of the cytotrophoblast to form the primitive yolk sac. An erosion of the endothelial lining of the
maternal capillaries by the syncytiotrophoblastic cells results in the formation of the maternal sinusoids from
where the blood will begin to penetrate and flow into and through the trophoblastic lacunae to give rise to
the uteroplacental circulation.[12][13] Subsequently, new cells derived from yolk sac will be established
between trophoblast and exocoelomic membrane and will give rise to extra-embryonic mesoderm, which
will form the chorionic cavity.[11]

At the end of the second week of development, some cells of the trophoblast penetrate and form rounded
columns into the syncytiotrophoblast. These columns are known as primary villi. At the same time, other
migrating cells form into the exocoelomic cavity a new cavity named the secondary or definitive yolk sac,
smaller than the primitive yolk sac.[11][12]

Implantation

After ovulation, the endometrial lining becomes transformed into a

secretory lining in preparation of accepting the embryo. It becomes
thickened, with its secretory glands becoming elongated, and is
increasingly vascular. This lining of the uterine cavity (or womb) is
now known as the decidua, and it produces a great number of large
decidual cells in its increased interglandular tissue. The blastomeres in
the blastocyst are arranged into an outer layer called the trophoblast.
The trophoblast then differentiates into an inner layer, the Trophoblast differentiation
cytotrophoblast, and an outer layer, the syncytiotrophoblast. The
cytotrophoblast contains cuboidal epithelial cells and is the source of
dividing cells, and the syncytiotrophoblast is a syncytial layer without cell boundaries.
The syncytiotrophoblast implants the blastocyst in the decidual epithelium by projections of chorionic villi,
forming the embryonic part of the placenta. The placenta develops once the blastocyst is implanted,
connecting the embryo to the uterine wall. The decidua here is termed the decidua basalis; it lies between
the blastocyst and the myometrium and forms the maternal part of the placenta. The implantation is assisted
by hydrolytic enzymes that erode the epithelium. The syncytiotrophoblast also produces human chorionic
gonadotropin, a hormone that stimulates the release of progesterone from the corpus luteum. Progesterone
enriches the uterus with a thick lining of blood vessels and capillaries so that it can oxygenate and sustain
the developing embryo. The uterus liberates sugar from stored glycogen from its cells to nourish the
embryo.[14] The villi begin to branch and contain blood vessels of the embryo. Other villi, called terminal or
free villi, exchange nutrients. The embryo is joined to the trophoblastic shell by a narrow connecting stalk
that develops into the umbilical cord to attach the placenta to the embryo.[11][15] Arteries in the decidua are
remodelled to increase the maternal blood flow into the intervillous spaces of the placenta, allowing gas
exchange and the transfer of nutrients to the embryo. Waste products from the embryo will diffuse across
the placenta.

As the syncytiotrophoblast starts to penetrate the uterine wall, the inner cell mass (embryoblast) also
develops. The inner cell mass is the source of embryonic stem cells, which are pluripotent and can develop
into any one of the three germ layer cells, and which have the potency to give rise to all the tissues and
organs.

Embryonic disc

The embryoblast forms an embryonic disc of two layers, the upper layer is called the epiblast and the lower
layer, the hypoblast. The disc is stretched between what will become the amniotic cavity and the yolk sac.
The epiblast is adjacent to the trophoblast and made of columnar cells; the hypoblast is closest to the
blastocyst cavity and made of cuboidal cells. The epiblast migrates away from the trophoblast downwards,
forming the amniotic cavity, the lining of which is formed from amnioblasts developed from the epiblast.
The hypoblast is pushed down and forms the yolk sac (exocoelomic cavity) lining. Some hypoblast cells
migrate along the inner cytotrophoblast lining of the blastocoel, secreting an extracellular matrix along the
way. These hypoblast cells and extracellular matrix are called Heuser's membrane (or the exocoelomic
membrane), and they cover the blastocoel to form the yolk sac (or exocoelomic cavity). Cells of the
hypoblast migrate along the outer edges of this reticulum and form the extraembryonic mesoderm; this
disrupts the extraembryonic reticulum. Soon pockets form in the reticulum, which ultimately coalesce to
form the chorionic cavity (extraembryonic coelom).

Gastrulation

Histogenesis of the three germ layers

The primitive streak, a linear collection of cells formed by the migrating epiblast, appears, and this marks
the beginning of gastrulation, which takes place around the seventeenth day (week 3) after fertilization. The
process of gastrulation reorganises the two-layer embryo into a three-layer embryo, and also gives the
embryo its specific head-to-tail, and front-to-back orientation, by way
of the primitive streak which establishes bilateral symmetry. A primitive
node (or primitive knot) forms in front of the primitive streak which is
the organiser of neurulation. A primitive pit forms as a depression in
the centre of the primitive node which connects to the notochord which
lies directly underneath. The node has arisen from epiblasts of the
amniotic cavity floor, and it is this node that induces the formation of
the neural plate which serves as the basis for the nervous system.
Artificially colored – gestational
The neural plate will form opposite the primitive streak from sac, yolk sac and embryo
ectodermal tissue which thickens and flattens into the neural plate. The (measuring 3 mm at five weeks)
epiblast in that region moves down into the streak at the location of the
primitive pit where the process
called ingression, which leads to the
formation of the mesoderm takes
place. This ingression sees the cells
from the epiblast move into the
primitive streak in an epithelial-
mesenchymal transition; epithelial
cells become mesenchymal stem
cells, multipotent stromal cells that
can differentiate into various cell
types. The hypoblast is pushed out
of the way and goes on to form the
amnion. The epiblast keeps moving
Embryo attached to placenta in amniotic cavity
and forms a second layer, the
mesoderm. The epiblast has now
differentiated into the three germ
layers of the embryo, so that the bilaminar disc is now a trilaminar disc, the gastrula.

The three germ layers are the ectoderm, mesoderm and endoderm, and are formed as three overlapping flat
discs. It is from these three layers that all the structures and organs of the body will be derived through the
processes of somitogenesis, histogenesis and organogenesis.[16] The embryonic endoderm is formed by
invagination of epiblastic cells that migrate to the hypoblast, while the mesoderm is formed by the cells that
develop between the epiblast and endoderm. In general, all germ layers will derive from the epiblast.[11][15]
The upper layer of ectoderm will give rise to the outermost layer of skin, central and peripheral nervous
systems, eyes, inner ear, and many connective tissues.[17] The middle layer of mesoderm will give rise to
the heart and the beginning of the circulatory system as well as the bones, muscles and kidneys. The inner
layer of endoderm will serve as the starting point for the development of the lungs, intestine, thyroid,
pancreas and bladder.

Following ingression, a blastopore develops where the cells have ingressed, in one side of the embryo and it
deepens to become the archenteron, the first formative stage of the gut. As in all deuterostomes, the
blastopore becomes the anus whilst the gut tunnels through the embryo to the other side where the opening
becomes the mouth. With a functioning digestive tube, gastrulation is now completed and the next stage of
neurulation can begin.

Neurulation
 Following
gastrulation, the
ectoderm gives
rise to epithelial
and neural tissue,
and the gastrula is
now referred to as
the neurula. The
neural plate that
has formed as a
thickened plate
from the
ectoderm,
continues to
broaden and its Neural tube development
Neural plate
ends start to fold
upwards as neural
folds. Neurulation refers to this folding process whereby the neural plate is transformed into the neural tube,
and this takes place during the fourth week. They fold, along a shallow neural groove which has formed as
a dividing median line in the neural plate. This deepens as the folds continue to gain height, when they will
meet and close together at the neural crest. The cells that migrate through the most cranial part of the
primitive line form the paraxial mesoderm, which will give rise to the somitomeres that in the process of
somitogenesis will differentiate into somites that will form the sclerotomes, the syndetomes,[18] the
myotomes and the dermatomes to form cartilage and bone, tendons, dermis (skin), and muscle. The
intermediate mesoderm gives rise to the urogenital tract and consists of cells that migrate from the middle
region of the primitive line. Other cells migrate through the caudal part of the primitive line and form the
lateral mesoderm, and those cells migrating by the most caudal part contribute to the extraembryonic
mesoderm.[11][15]

The embryonic disc begins flat and round, but eventually elongates to have a wider cephalic part and
narrow-shaped caudal end.[10] At the beginning, the primitive line extends in cephalic direction and 18 days
after fertilization returns caudally until it disappears. In the cephalic portion, the germ layer shows specific
differentiation at the beginning of the fourth week, while in the caudal portion it occurs at the end of the
fourth week.[11] Cranial and caudal neuropores become progressively smaller until they close completely
(by day 26) forming the neural tube.[19]

Development of organs and organ systems

Organogenesis is the development of the organs that begins during the third to eighth week, and continues
until birth. Sometimes full development, as in the lungs, continues after birth. Different organs take part in
the development of the many organ systems of the body.

Blood

Haematopoietic stem cells that give rise to all the blood cells develop from the mesoderm. The development
of blood formation takes place in clusters of blood cells, known as blood islands, in the yolk sac. Blood
islands develop outside the embryo, on the umbilical vesicle, allantois, connecting stalk, and chorion, from
mesodermal hemangioblasts.
In the centre of a blood island, hemangioblasts form the
haematopoietic stem cells that are the precursor to all types of blood
cell. In the periphery of a blood island the hemangioblasts
differentiate into angioblasts, the precursors to the blood vessels.[20]

Heart and circulatory system

The heart is the first functional organ to develop and starts to beat
and pump blood at around 22 days.[21] Cardiac myoblasts and
blood islands in the splanchnopleuric mesenchyme on each side of
the neural plate, give rise to the cardiogenic region.[11]: 165 This is a
horseshoe-shaped area near to the head of the embryo. By day 19,
following cell signalling, two strands begin to form as tubes in this
region, as a lumen develops within them. These two endocardial
tubes grow and by day 21 have migrated towards each other and
fused to form a single primitive heart tube, the tubular heart. This is
enabled by the folding of the embryo which pushes the tubes into
the thoracic cavity.[22] Nine-week-old human embryo from
an ectopic pregnancy
Also at the same time that the endocardial tubes are forming,
vasculogenesis (the
development of the circulatory
system) has begun. This starts
on day 18 with cells in the
splanchnopleuric mesoderm
differentiating into angioblasts
that develop into flattened
endothelial cells. These join to
form small vesicles called
angiocysts which join up to
form long vessels called
angioblastic cords. These cords
develop into a pervasive
network of plexuses in the
formation of the vascular
network. This network grows
by the additional budding and
sprouting of new vessels in the
process of angiogenesis.[22]
Following vasculogenesis and
the development of an early
vasculature, a stage of vascular
remodelling takes place.

The tubular heart quickly forms five distinct regions. From head to tail, these are the infundibulum, bulbus
cordis, primitive ventricle, primitive atrium, and the sinus venosus. Initially, all venous blood flows into the
sinus venosus, and is propelled from tail to head to the truncus arteriosus. This will divide to form the aorta
and pulmonary artery; the bulbus cordis will develop into the right (primitive) ventricle; the primitive
ventricle will form the left ventricle; the primitive atrium will become the front parts of the left and right atria
and their appendages, and the sinus venosus will develop into the posterior part of the right atrium, the
sinoatrial node and the coronary sinus.[21]
Cardiac looping begins to shape the heart as one of the processes of morphogenesis, and this completes by
the end of the fourth week. Programmed cell death (apoptosis) at the joining surfaces enables fusion to take
place.[22] In the middle of the fourth week, the sinus venosus receives blood from the three major veins: the
vitelline, the umbilical and the common cardinal veins.

During the first two months of development, the interatrial septum begins to form. This septum divides the
primitive atrium into a right and a left atrium. Firstly it starts as a crescent-shaped piece of tissue which
grows downwards as the septum primum. The crescent shape prevents the complete closure of the atria
allowing blood to be shunted from the right to the left atrium through the opening known as the ostium
primum. This closes with further development of the system but before it does, a second opening (the
ostium secundum) begins to form in the upper atrium enabling the continued shunting of blood.[22]

A second septum (the septum secundum) begins to form to the right of the septum primum. This also leaves
a small opening, the foramen ovale which is continuous with the previous opening of the ostium secundum.
The septum primum is reduced to a small flap that acts as the valve of the foramen ovale and this remains
until its closure at birth. Between the ventricles the septum inferius also forms which develops into the
muscular interventricular septum.[22]

Digestive system

The digestive system starts to develop from the third week and by the twelfth week, the organs have
correctly positioned themselves.

Respiratory system

The respiratory system develops from the lung bud, which appears in the ventral wall of the foregut about
four weeks into development. The lung bud forms the trachea and two lateral growths known as the
bronchial buds, which enlarge at the beginning of the fifth week to form the left and right main bronchi.
These bronchi in turn form secondary (lobar) bronchi; three on the right and two on the left (reflecting the
number of lung lobes). Tertiary bronchi form from secondary bronchi.

While the internal lining of the larynx originates from the lung bud, its cartilages and muscles originate from
the fourth and sixth pharyngeal arches.[23]

Urinary system

Kidneys

Three different kidney systems form in the developing embryo: the pronephros, the mesonephros and the
metanephros. Only the metanephros develops into the permanent kidney. All three are derived from the
intermediate mesoderm.

Pronephros
The pronephros derives from the intermediate mesoderm in the cervical region. It is not functional and
degenerates before the end of the fourth week.

Mesonephros

The mesonephros derives from intermediate mesoderm in the upper thoracic to upper lumbar segments.
Excretory tubules are formed and enter the mesonephric duct, which ends in the cloaca. The mesonephric
duct atrophies in females, but participate in development of the reproductive system in males.

Metanephros

The metanephros appears in the fifth week of development. An outgrowth of the mesonephric duct, the
ureteric bud, penetrates metanephric tissue to form the primitive renal pelvis, renal calyces and renal
pyramids. The ureter is also formed.

Bladder and urethra

Between the fourth and seventh weeks of development, the urorectal septum divides the cloaca into the
urogenital sinus and the anal canal. The upper part of the urogenital sinus forms the bladder, while the lower
part forms the urethra.[23]

Reproductive system

Integumentary system

The superficial layer of the skin, the epidermis, is derived from the ectoderm. The deeper layer, the dermis,
is derived from mesenchyme.

The formation of the epidermis begins in the second month of development and it acquires its definitive
arrangement at the end of the fourth month. The ectoderm divides to form a flat layer of cells on the surface
known as the periderm. Further division forms the individual layers of the epidermis.

The mesenchyme that will form the dermis is derived from three sources:

The mesenchyme that forms the dermis in the limbs and body wall derives from the lateral
plate mesoderm
The mesenchyme that forms the dermis in the back derives from paraxial mesoderm
The mesenchyme that forms the dermis in the face and neck derives from neural crest
cells[23]

Nervous system

Late in the fourth week, the superior part of the neural tube bends ventrally as the cephalic flexure at the
level of the future midbrain—the mesencephalon.[24] Above the mesencephalon is the prosencephalon
(future forebrain) and beneath it is the rhombencephalon (future hindbrain).

Cranial neural crest cells migrate to the pharyngeal arches as neural stem cells, where they develop in the
process of neurogenesis into neurons.
The optical vesicle (which eventually becomes the optic nerve,
retina and iris) forms at the basal plate of the prosencephalon.
The alar plate of the prosencephalon expands to form the
cerebral hemispheres (the telencephalon) whilst its basal plate
becomes the diencephalon. Finally, the optic vesicle grows to
form an optic outgrowth.

Development of physical features

Development of brain in eight-week-old

Face and neck
embryo

From the third to the eighth week the face and neck develop.

Ears

The inner ear, middle ear and outer ear have distinct embryological origins.

Inner ear

At about 22 days into development, the ectoderm on each side of the rhombencephalon thickens to form
otic placodes. These placodes invaginate to form otic pits, and then otic vesicles. The otic vesicles then form
ventral and dorsal components.

The ventral component forms the saccule and the cochlear duct. In the sixth week of development the
cochlear duct emerges and penetrates the surrounding mesenchyme, travelling in a spiral shape until it forms
2.5 turns by the end of the eighth week. The saccule is the remaining part of the ventral component. It
remains connected to the cochlear duct via the narrow ductus reuniens.

The dorsal component forms the utricle and semicircular canals.

Middle ear

The first pharyngeal pouch lengthens and expands to form the tubotympanic recess. This recess
differentiates to form most of the tympanic cavity of the middle ear, and all of the Eustachian or auditory
tube. The narrow auditory tube connects the tympanic cavity to the pharynx.[25]

The bones of the middle ear, the ossicles, derive from the cartilages of the pharyngeal arches. The malleus
and incus derive from the cartilage of the first pharyngeal arch, whereas the stapes derives from the cartilage
of the second pharyngeal arch.

Outer ear
The external auditory meatus develops from the dorsal portion of the first pharyngeal cleft. Six auricular
hillocks, which are mesenchymal proliferations at the dorsal aspects of the first and second pharyngeal
arches, form the auricle of the ear.[23]

Eyes

The eyes begin to develop from the third week to the tenth week.

Limbs

At the end of the fourth week limb development begins. Limb buds
appear on the ventrolateral aspect of the body. They consist of an
outer layer of ectoderm and an inner part consisting of mesenchyme
which is derived from the parietal layer of lateral plate mesoderm.
Ectodermal cells at the distal end of the buds form the apical
ectodermal ridge, which creates an area of rapidly proliferating
mesenchymal cells known as the progress zone. Cartilage (some of
Movements of embryo at nine weeks
which ultimately becomes bone) and muscle develop from the
gestational age
mesenchyme.[23]

Clinical significance
Toxic exposures in the embryonic period can be the cause of major congenital malformations, since the
precursors of the major organ systems are now developing.

Each cell of the preimplantation embryo has the potential to form all of the different cell types in the
developing embryo. This cell potency means that some cells can be removed from the preimplantation
embryo and the remaining cells will compensate for their absence. This has allowed the development of a
technique known as preimplantation genetic diagnosis, whereby a small number of cells from the
preimplantation embryo created by IVF, can be removed by biopsy and subjected to genetic diagnosis. This
allows embryos that are not affected by defined genetic diseases to be selected and then transferred to the
mother's uterus.

Sacrococcygeal teratomas, tumours formed from different types of tissue, that can form, are thought to be
related to primitive streak remnants, which ordinarily disappear.[10][11][13]

First arch syndromes are congenital disorders of facial deformities, caused by the failure of neural crest cells
to migrate to the first pharyngeal arch.

Spina bifida a congenital disorder is the result of the incomplete closure of the neural tube.

Vertically transmitted infections can be passed from the mother to the unborn child at any stage of its
development.

Hypoxia a condition of inadequate oxygen supply can be a serious consequence of a preterm or premature
birth.

See also
Embryo loss
 Aorta-gonad-mesonephros
CDX2
Developmental biology
Drosophila embryogenesis
Embryomics
Human tooth development
List of human cell types derived from the germ layers
Potential person
Recapitulation theory
The Human Embryo

Additional images

Representing different Early stage of the Phase of the Top of the form of the
stages of gastrulation process gastrulation process embryo
embryogenesis

Establishment of Spinal cord at five Head and neck at 32

embryo medium weeks days

References
1. "germinal stage" (http://medical-dictionary.thefreedictionary.com/germinal+stage). Mosby's
Medical Dictionary, 8th edition. Elsevier. Retrieved 6 October 2013.
2. Marlow, Florence L (2020). Maternal Effect Genes in Development (https://books.google.co
m/books?id=JX3nDwAAQBAJ&pg=PA124). Academic Press. p. 124. ISBN 978-
0128152218.
3. Singh, Vishram (2013). Textbook of Clinical Embryology – E-book (https://books.google.com/
books?id=ABBtAwAAQBAJ&pg=PA35). Elsevier Health Sciences. p. 35. ISBN 978-
8131236208.
 4. Standring, Susan (2015). Gray's Anatomy E-Book: The Anatomical Basis of Clinical Practice
(https://books.google.com/books?id=b7FVCgAAQBAJ&pg=PA163). Elsevier Health
Sciences. p. 163. ISBN 978-0702068515.
5. Forgács, G.; Newman, Stuart A. (2005). "Cleavage and blastula formation" (https://books.goo
gle.com/books?id=rUyVWQhk7CkC&pg=PA27). Biological physics of the developing
embryo. Cambridge University Press. p. 27. ISBN 978-0-521-78337-8.
6. Standring, Susan (2015). Gray's Anatomy E-Book: The Anatomical Basis of Clinical Practice
(https://books.google.com/books?id=b7FVCgAAQBAJ&pg=PA165). Elsevier Health
Sciences. p. 166. ISBN 978-0702068515.
7. Brison, D. R.; Sturmey, R. G.; Leese, H. J. (2014). "Metabolic heterogeneity during
preimplantation development: the missing link?" (https://doi.org/10.1093%2Fhumupd%2Fdm
u018). Human Reproduction Update. 20 (5): 632–640. doi:10.1093/humupd/dmu018 (https://
doi.org/10.1093%2Fhumupd%2Fdmu018). ISSN 1355-4786 (https://www.worldcat.org/issn/1
355-4786). PMID 24795173 (https://pubmed.ncbi.nlm.nih.gov/24795173).
8. Boklage, Charles E. (2009). How New Humans Are Made: Cells and Embryos, Twins and
Chimeras, Left and Right, Mind/Self/Soul, Sex, and Schizophrenia (https://books.google.co
m/books?id=j7GfgRIgP9YC&pg=PT233). World Scientific. p. 217. ISBN 978-981-283-513-0.
9. "28.2 Embryonic Development – Anatomy and Physiology" (https://opentextbc.ca/anatomyan
dphysiology/chapter/28-2-embryonic-development/). opentextbc.ca.
10. Carlson, Bruce M. (1999) [1t. Pub. 1997]. "Chapter 4: Formation of germ layers and initial
derivatives". Human Embryology & Developmental Biology. Mosby, Inc. pp. 62–68. ISBN 0-
8151-1458-3.
11. Sadler, T.W.; Langman, Jan (2012) [1st. Pub. 2001]. "Chapter 3: Primera semana del
desarrollo: de la ovulación a la implantación". In Seigafuse, sonya (ed.). Langman,
Embriología médica. Lippincott Williams & Wilkins, Wolters Kluwer. pp. 29–42. ISBN 978-
84-15419-83-9.
12. Moore, Keith L.; Persaud, V.N. (2003) [1t. Pub. 1996]. "Chapter 3: Formation of the bilaminar
embryonic disc: second week". The Developing Human, Clinically Oriented Embryology. W
B Saunders Co. pp. 47–51. ISBN 0-7216-9412-8.
13. Larsen, William J.; Sherman, Lawrence S.; Potter, S. Steven; Scott, William J. (2001) [1t.
Pub. 1998]. "Chapter 2: Bilaminar embryonic disc development and establishment of the
uteroplacental circulation". Human Embryology. Churchill Livingstone. pp. 37–45. ISBN 0-
443-06583-7.
14. Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life (http://w
ww.phschool.com/el_marketing.html). Boston: Pearson Prentice Hall. ISBN 0-13-250882-6.
15. Smith Agreda, Víctor; Ferrés Torres, Elvira; Montesinos Castro-Girona, Manuel (1992).
"Chapter 5: Organización del desarrollo: Fase de germinación". Manual de embriología y
anatomía general. Universitat de València. pp. 72–85. ISBN 84-370-1006-3.
16. Schünke, Michael; Ross, Lawrence M.; Schulte, Erik; Schumacher, Udo; Lamperti, Edward
D. (2006). Thieme Atlas of Anatomy: General Anatomy and Musculoskeletal System (https://b
ooks.google.com/books?id=NK9TgTaGt6UC&pg=PA6). Thieme. ISBN 978-3-13-142081-7.
17. "Pregnancy week by week" (http://www.mayoclinic.com/health/prenatal-care/PR00112/NSE
CTIONGROUP=2). Retrieved 28 July 2010.
18. Brent AE, Schweitzer R, Tabin CJ (April 2003). "A somitic compartment of tendon
progenitors" (https://doi.org/10.1016%2FS0092-8674%2803%2900268-X). Cell. 113 (2):
235–48. doi:10.1016/S0092-8674(03)00268-X (https://doi.org/10.1016%2FS0092-8674%28
03%2900268-X). PMID 12705871 (https://pubmed.ncbi.nlm.nih.gov/12705871).
S2CID 16291509 (https://api.semanticscholar.org/CorpusID:16291509).
19. Larsen, W J (2001). Human Embryology (3rd ed.). Elsevier. p. 87. ISBN 0-443-06583-7.
20. Sadler, T.W. (2010). Langman's Medical Embryology (11th ed.). Baltimore: Lippincott
Williams &Wilkins. pp. 79–81. ISBN 9780781790697.
21. Betts, J. Gordon (2013). Anatomy & physiology (http://cnx.org/content/m46676/latest/?collecti
on=col11496/latest). pp. 787–846. ISBN 978-1938168130.
22. Larsen, W J (2001). Human Embryology (3rd ed.). Elsevier. pp. 170–190. ISBN 0-443-
06583-7.
23. Sadler, Thomas W. (2012). Langman's medical embryology. Langman, Jan. (12th ed.).
Philadelphia: Wolters Kluwer Health/Lippincott Williams & Wilkins. ISBN 9781451113426.
OCLC 732776409 (https://www.worldcat.org/oclc/732776409).
24. Zhou, Yi; Song, Hongjun; Ming, Guo-Li (2023-07-28). "Genetics of human brain
development" (https://pubmed.ncbi.nlm.nih.gov/37507490). Nature Reviews. Genetics.
doi:10.1038/s41576-023-00626-5 (https://doi.org/10.1038%2Fs41576-023-00626-5).
ISSN 1471-0064 (https://www.worldcat.org/issn/1471-0064). PMID 37507490 (https://pubme
d.ncbi.nlm.nih.gov/37507490).
25. Larsen's human embryology (5. ed.). Philadelphia, Pa: Elsevier, Churchill Livingstone. 2015.
p. 485. ISBN 9781455706846.

External links
Virtual human embryo (https://www.ehd.org/virtual-human-embryo)
Photo of blastocyst in utero (https://www.pbs.org/wgbh/nova/miracle/images/stem_blastocys
t.jpg)
Slideshow: In the Womb (https://web.archive.org/web/20161222152312/http://www.lifehugge
r.com/doc/99/womb)

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