Animal embryonic development

In developmental biology, animal embryonic development,

also known as animal embryogenesis, is the developmental
stage of an animal embryo. Embryonic development starts
with the fertilization of an egg cell (ovum) by a sperm cell
(spermatozoon).[1] Once fertilized, the ovum becomes a
single diploid cell known as a zygote. The zygote undergoes
mitotic divisions with no significant growth (a process known
as cleavage) and cellular differentiation, leading to
development of a multicellular embryo[2] after passing
through an organizational checkpoint during mid-
embryogenesis.[3] In mammals, the term refers chiefly to the
early stages of prenatal development, whereas the terms fetus
and fetal development describe later stages.[2][4]

The main stages of animal embryonic development are as

follows:

The zygote undergoes a series of cell divisions

(called cleavage) to form a structure called a morula.
The morula develops into a structure called a
blastula through a process called blastulation.
The blastula develops into a structure called a
gastrula through a process called gastrulation.
The gastrula then undergoes further development,
including the formation of organs (organogenesis). Diagram of stages of embryo
development to a larval and adult stage.
The embryo then transforms into the next stage of
development, the nature of which varies among different
animal species (examples of possible next stages include a fetus and a larva).

Fertilization and the zygote

The egg cell is generally asymmetric, having an animal pole (future ectoderm). It is covered with
protective envelopes, with different layers. The first envelope – the one in contact with the membrane of
the egg – is made of glycoproteins and is known as the vitelline membrane (zona pellucida in mammals).
Different taxa show different cellular and acellular envelopes englobing the vitelline membrane.[2][5]

Fertilization is the fusion of gametes to produce a new organism. In animals, the process involves a sperm
fusing with an ovum, which eventually leads to the development of an embryo. Depending on the animal
species, the process can occur within the body of the female in internal fertilization, or outside in the case
of external fertilization. The fertilized egg cell is known as the zygote.[2][5]
To prevent more than one sperm fertilizing the egg (polyspermy), fast block and slow block to
polyspermy are used. Fast block, the membrane potential rapidly depolarizing and then returning to
normal, happens immediately after an egg is fertilized by a single sperm. Slow block begins in the first
few seconds after fertilization and is when the release of calcium causes the cortical reaction, in which
various enzymes are released from cortical granules in the eggs plasma membrane, causing the expansion
and hardening of the outside membrane, preventing more sperm from entering.[6][5]

Cleavage and morula

Cell division with no significant growth, producing a
cluster of cells that is the same size as the original
zygote, is called cleavage. At least four initial cell
divisions occur, resulting in a dense ball of at least
sixteen cells called the morula. In the early mouse
embryo, the sister cells of each division remain
connected during interphase by microtubule
bridges.[7] The different cells derived from cleavage,
up to the blastula stage, are called blastomeres.
Depending mostly on the amount of yolk in the egg,
the cleavage can be holoblastic (total) or meroblastic
Cell divisions (cleavage)
(partial).[8][9]

Holoblastic cleavage occurs in animals with little yolk

in their eggs,[10] such as humans and other mammals who receive nourishment as embryos from the
mother, via the placenta or milk, such as might be secreted from a marsupium. Meroblastic cleavage
occurs in animals whose eggs have more yolk (i.e. birds and reptiles). Because cleavage is impeded in the
vegetal pole, there is an uneven distribution and size of cells, being more numerous and smaller at the
animal pole of the zygote.[8][9]

In holoblastic eggs, the first cleavage always occurs along the vegetal-animal axis of the egg, and the
second cleavage is perpendicular to the first. From here the spatial arrangement of blastomeres can follow
various patterns, due to different planes of cleavage, in various organisms:

Cleavage patterns followed by holoblastic and meroblastic eggs in animals

Holoblastic Meroblastic

Radial (sea urchin,[9][11] amphioxus) Discoidal (fish, monotremes,

Bilateral (tunicates,[9][12] amphibians[13]) birds,[9][15] reptiles[9])
Spiral (annelids, mollusks[9][14]) Superficial (insects[9][17])
Rotational (placental mammals, marsupials,[9][15]
nematodes[16])
The end of cleavage is known as midblastula transition and coincides with the onset of zygotic
transcription.
In amniotes, the cells of the morula are at first closely aggregated, but soon they become arranged into an
outer or peripheral layer, the trophoblast, which does not contribute to the formation of the embryo
proper, and an inner cell mass, from which the embryo is developed. Fluid collects between the
trophoblast and the greater part of the inner cell-mass, and thus the morula is converted into a vesicle,
called the blastodermic vesicle. The inner cell mass remains in contact, however, with the trophoblast at
one pole of the ovum; this is named the embryonic pole, since it indicates the location where the future
embryo will develop.[18][9]

Formation of the blastula

After the seventh cleavage has produced 128 cells, the morula becomes a blastula.[8] The blastula is
usually a spherical layer of cells (the blastoderm) surrounding a fluid-filled or yolk-filled cavity the
blastocoel.

Mammals at this stage form a structure called the blastocyst,[1] characterized by an inner cell mass that is
distinct from the surrounding blastula.[19][20][21] The blastocyst is similar in structure to the blastula but
their cells have different fates. In the mouse, primordial germ cells arise from the inner cell mass (the
epiblast) as a result of extensive genome-wide reprogramming.[22] Reprogramming involves global DNA
demethylation facilitated by the DNA base excision repair pathway as well as chromatin reorganization,
and results in cellular totipotency.[23][20]

Before gastrulation, the cells of the trophoblast become differentiated into two layers: The outer layer
forms a syncytium (i.e., a layer of protoplasm studded with nuclei, but showing no evidence of
subdivision into cells), termed the syncytiotrophoblast, while the inner layer, the cytotrophoblast, consists
of well-defined cells. As already stated, the cells of the trophoblast do not contribute to the formation of
the embryo proper; they form the ectoderm of the chorion and play an important part in the development
of the placenta. On the deep surface of the inner cell mass, a layer of flattened cells, called the endoderm,
is differentiated and quickly assumes the form of a small sac, called the yolk sac. Spaces appear between
the remaining cells of the mass and, by the enlargement and coalescence of these spaces, a cavity called
the amniotic cavity is gradually developed. The floor of this cavity is formed by the embryonic disk,
which is composed of a layer of prismatic cells – the embryonic ectoderm, derived from the inner cell
mass and lying in apposition with the endoderm.[18][20]

Formation of the germ layers

The embryonic disc becomes oval and then pear-shaped, the wider end being directed forward. Towards
the narrow, posterior end, an opaque primitive streak, is formed and extends along the middle of the disc
for about half of its length; at the anterior end of the streak there is a knob-like thickening termed the
primitive node or knot, (known as Hensen's knot in birds). A shallow groove, the primitive groove,
appears on the surface of the streak, and the anterior end of this groove communicates by means of an
aperture, the blastopore, with the yolk sac. The primitive streak is produced by a thickening of the axial
part of the ectoderm, the cells of which multiply, grow downward, and blend with those of the subjacent
endoderm. From the sides of the primitive streak a third layer of cells, the mesoderm, extends laterally
between the ectoderm and endoderm; the caudal end of the primitive streak forms the cloacal membrane.
The blastoderm now consists of three layers, an outer ectoderm, a middle mesoderm, and an inner
endoderm; each has distinctive characteristics and gives rise
to certain tissues of the body. For many mammals, it is
sometime during formation of the germ layers that
implantation of the embryo in the uterus of the mother
occurs.[18][20]

Formation of the gastrula

During gastrulation cells migrate to the interior of the
blastula, subsequently forming two (in diploblastic animals)
or three (triploblastic) germ layers. The embryo during this
process is called a gastrula. The germ layers are referred to as
the ectoderm, mesoderm and endoderm. In diploblastic
animals only the ectoderm and the endoderm are present.[8]*
Among different animals, different combinations of the
following processes occur to place the cells in the interior of
the embryo: Comparative vertebrate embryology.

Epiboly – expansion of one cell sheet over other

cells[2][9]
Ingression – migration of individual cells into the embryo (cells move with pseudopods)
[3][9]

Invagination – infolding of cell sheet into embryo, forming the mouth, anus, and
archenteron.[8][9]
Delamination – splitting or migration of one sheet into two sheets[9]
Involution – inturning of cell sheet over the basal surface of an outer layer[9]
Polar proliferation – Cells at the polar ends of the blastula/gastrula proliferate, mostly at
the animal pole.[9]
Other major changes during gastrulation:
Heavy RNA transcription using embryonic genes; up to this point the RNAs used were
maternal (stored in the unfertilized egg).
Cells start major differentiation processes, losing their totipotentiality.
In most animals, a blastopore is formed at the point where cells are migrating inward. Two major groups
of animals can be distinguished according to the blastopore's fate. In deuterostomes the anus forms from
the blastopore, while in protostomes it develops into the mouth.[9]

Formation of the early nervous system – neural groove, tube and notochord
In front of the primitive streak, two longitudinal ridges, caused by a folding up of the ectoderm, make
their appearance, one on either side of the middle line formed by the streak. These are named the neural
folds; they commence some little distance behind the anterior end of the embryonic disk, where they are
continuous with each other, and from there gradually extend backward, one on either side of the anterior
end of the primitive streak. Between these folds is a shallow median groove, the neural groove. The
groove gradually deepens as the neural folds become elevated, and ultimately the folds meet and coalesce
in the middle line and convert the groove into a closed tube, the neural tube or canal, the ectodermal wall
of which forms the rudiment of the nervous system. After the coalescence of the neural folds over the
anterior end of the primitive streak, the blastopore no longer opens on the surface but into the closed
canal of the neural tube, and thus a transitory communication, the neurenteric canal, is established
between the neural tube and the primitive digestive tube. The coalescence of the neural folds occurs first
in the region of the hind brain, and from there extends forward and backward; toward the end of the third
week, the front opening (anterior neuropore) of the tube finally closes at the anterior end of the future
brain, and forms a recess that is in contact, for a time, with the overlying ectoderm; the hinder part of the
neural groove presents for a time a rhomboidal shape, and to this expanded portion the term sinus
rhomboidalis has been applied. Before the neural groove is closed, a ridge of ectodermal cells appears
along the prominent margin of each neural fold; this is termed the neural crest or ganglion ridge, and
from it the spinal and cranial nerve ganglia and the ganglia of the sympathetic nervous system are
developed. By the upward growth of the mesoderm, the neural tube is ultimately separated from the
overlying ectoderm.[24][9]

The cephalic end of the neural groove exhibits several

dilatations that, when the tube is closed, assume the form of
the three primary brain vesicles, and correspond, respectively,
to the future forebrain (prosencephalon), midbrain
(mesencephalon), and hindbrain (rhombencephalon) (Fig.
18). The walls of the vesicles are developed into the nervous
tissue and neuroglia of the brain, and their cavities are
modified to form its ventricles. The remainder of the tube
forms the spinal cord (medulla spinalis); from its ectodermal
wall the nervous and neuroglial elements of the spinal cord
Dissection of human embryo
are developed, while the cavity persists as the central
canal.[24][9]

Formation of the early septum

The extension of the mesoderm takes place throughout the whole of the embryonic and extra-embryonic
areas of the ovum, except in certain regions. One of these is seen immediately in front of the neural tube.
Here the mesoderm extends forward in the form of two crescentic masses, which meet in the middle line
so as to enclose behind them an area that is devoid of mesoderm. Over this area, the ectoderm and
endoderm come into direct contact with each other and constitute a thin membrane, the buccopharyngeal
membrane, which forms a septum between the primitive mouth and pharynx.[18][9]

Early formation of the heart and other primitive structures

In front of the buccopharyngeal area, where the lateral crescents of mesoderm fuse in the middle line, the
pericardium is afterward developed, and this region is therefore designated the pericardial area. A second
region where the mesoderm is absent, at least for a time, is that immediately in front of the pericardial
area. This is termed the proamniotic area, and is the region where the proamnion is developed; in humans,
however, it appears that a proamnion is never formed. A third region is at the hind end of the embryo,
where the ectoderm and endoderm come into apposition and form the cloacal membrane.[18][9]
Somitogenesis
Somitogenesis is the process by which somites (primitive
segments) are produced. These segmented tissue blocks
differentiate into skeletal muscle, vertebrae, and dermis of
all vertebrates.[25]

Somitogenesis begins with the formation of somitomeres

(whorls of concentric mesoderm) marking the future
somites in the presomitic mesoderm (unsegmented
0:00
paraxial). The presomitic mesoderm gives rise to
successive pairs of somites, identical in appearance that
differentiate into the same cell types but the structures Sample 3D cell of somitogenesis
formed by the cells vary depending upon the
anteroposterior (e.g., the thoracic vertebrae have ribs, the
lumbar vertebrae do not). Somites have unique positional values along this axis and it is thought that
these are specified by the Hox homeotic genes.[25]

Toward the end of the second week after fertilization, transverse segmentation of the paraxial mesoderm
begins, and it is converted into a series of well-defined, more or less cubical masses, also known as the
somites, which occupy the entire length of the trunk on either side of the middle line from the occipital
region of the head. Each segment contains a central cavity (known as a [myocoel), which, however, is
soon filled with angular and spindle-shape cells. The somites lie immediately under the ectoderm on the
lateral aspect of the neural tube and notochord, and are connected to the lateral mesoderm by the
intermediate cell mass. Those of the trunk may be arranged in the following groups, viz.: cervical 8,
thoracic 12, lumbar 5, sacral 5, and coccygeal from 5 to 8. Those of the occipital region of the head are
usually described as being four in number. In mammals, somites of the head can be recognized only in the
occipital region, but a study of the lower vertebrates leads to the belief that they are present also in the
anterior part of the head and that, altogether, nine segments are represented in the cephalic region.[26][25]

Organogenesis
At some point after the different germ layers are defined,
organogenesis begins. The first stage in vertebrates is called
neurulation, where the neural plate folds forming the neural
tube (see above).[8] Other common organs or structures that
arise at this time include the heart and somites (also above),
but from now on embryogenesis follows no common pattern
among the different taxa of the animalia.[2]

In most animals organogenesis, along with morphogenesis,

results in a larva. The hatching of the larva, which must then
Human embryo, 8–9 weeks, 38 mm
undergo metamorphosis, marks the end of embryonic
(1.5 in)
development.[2]
See also
Cdx2 gene Leech embryogenesis
Collective cell migration Parthenogenesis
Drosophila embryogenesis Plant embryogenesis
Enterocoely Schizocoely
Homeobox genes Shell-less chick embryo culture
Human embryogenesis

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External links
Cellular Darwinism ([Link]
on_in_cell_differentiation_and_embryo_development.html)
Embryogenesis & MMPs ([Link] PMAP The
Proteolysis Map-animation
Development of the embryo ([Link] (retrieved November
20, 2007)
Video ([Link] of embryogenesis of the frog
Xenopus laevis from shortly after fertilization until the hatching of the tadpole; acquired by
MRI (DOI of paper ([Link]

Retrieved from "[Link]