DEVELOPMENTAL BIOLOGY (BIO 304)

Part I: Animal Development

I. Introduction: Brief history of developmental ideas
II. Embryo Development
0. Fertilization
1. Cleavage and blastulation
2. Gastrulation
3. Organogenesis
III. Post-Embryonic development
1. Sexual maturation
2. Gametogenesis
Part II: Plant growth and development
I. Plant life cycles
II. Plant embryo development
III. Germination and seedling development
IV. Apical meristems and development
V. Flower development
VI. Plant growth regulators
VII. Plant propagation
VIII. Plant Senescence.
 PART I: ANIMAL DEVELOPMENT

I. INTRODUCTION

1. Brief history of developmental ideas

In the past there was a controversy over whether the organs of the embryo are formed de novo at
each generation, or whether the organs are present, in miniature form, within the egg (or sperm).
The first view was called epigenesis and it was supported by Aristotle (384–322 B.C.) and
Harvey (1578–1657). Aristotle believed that the embryo essentially formed by coagulation in the
uterus immediately after mating when the form-building principle of the male acted on the
material substance provided by the female. The idea was that an animal (or a plant) forms
gradually over time from a relatively formless egg or spore through a sequence of steps in which
organs are formed.
The second view, called preformation considered that organs were prefigured and embryonic
development merely required growth of existing structures, not formation of new ones. Adult
organism was prefigured in the germs cells, another generation already existed in a prefigured
state within the germ cells of the first prefigured generation.
The embryological case for epigenesis was proved by carefully observing developing embryos
and thus demonstrating that the embryonic parts develop from tissues that have no counterpart in
the adult organism. Preformationism was supported by many “embryologists” until the end of
the 1820s, when a combination of staining techniques and improved microscopes created a
revolution in descriptive embryology.
Epigenesis theory is now fully supported and in current biological usage, it refers primarily to the
origin of new structures during embryonic development.

2. What is Developmental Biology?

Developmental Biology is a large field of investigation that includes the study of all changes
associated with an organism as it progresses through the life cycle (Fig. 1.1). In its earliest
stages of development, a plant or animal is called an embryo. Sometimes the embryo is
contained within a protective structure, such as a seed coat, an eggshell, or a uterus. An embryo
does not photosynthesize or feed actively; instead, it obtains its food from its mother directly or
indirectly (by way of nutrients stored in the seed or egg).
The life cycles of all multicellular organisms exhibit many similarities. That is, as an organism
(plant or animal) progresses from one generation to the next there is a series of common
processes: for example, gametogenesis, fertilization, embryogenesis, cell differentiation, tissue
differentiation, organogenesis, maturation, growth, reproduction, senescence, and death.
Embryologists consider development from a zygote to a multicellular organism. In the particular
case of humans, development does not even stop at birth. Note for example that teeth continue to
develop, sex glands with sexual differentiation mature long after birth and in particular,
enormous developmental changes occur in the brain in the years between birth and adolescence.
Especially in the early years, there is a great deal of plasticity in the organization of the nervous
system as patterns of connection between neurons develop.

Figure 1.1 Stages of Development: Stages of development from embryo to adult are shown for a plant and an
animal. Growth, differentiation, and morphogenesis are all part of the complex process of development.

II. EMBRYO DEVELOPMENT

0. Fertilization
In most living organisms sexual reproduction takes place. It involves fertilization (Fig.1.2) which
the union of the male and female gametes. The haploid male gamete (sperm cell), which carries
the paternal genetic information, unites with the haploid female gamete (ovum or egg), which
carries the genetic information of the female parent, to form a diploid zygote. The entry of a
sperm into an egg activates the egg metabolically and initiates the rapid series of cell divisions.
Once the zygote is formed, replication of DNA begins, and the first division soon follows.
Figure1.2: Fertilization of a mammalian egg. This egg is encased in a jellylike substance (zona pellucida). The
first step in fertilization is a loose attachment of sperm to the surface of the zona pellucida. A mature sperm is an
elongated cell with 3 distinct parts: a head, a mid-piece, and a tail, all of which are enclosed by a cell membrane.

The zygote produces a diploid multicellular organism by several repeated and organized mitotic
divisions and cellular differentiation. It is the first step in the development of embryo formation.
Each cell in an animal (except reproductive cells) has the same set of genes that are used to build
the animal, yet animals have many different kinds of cells. From the fertilized egg come large
numbers of cells (trillions in humans) that consistently give rise to structural features of the
animal body.
Although the details of fertilization vary from species to species, conception generally consists of
four major events:
- Contact and recognition between sperm and egg. In most cases, this ensures that the sperm and
egg are the same species. The head of the sperm cell contains digestive enzymes which break
down an egg’s outer layer and enable fusion of the head of the sperm and the egg’s cell
membrane
- Regulation of sperm entry into the egg. Only one sperm can ultimately fertilize the egg. This is
usually accomplished by allowing only one sperm to enter the egg and inhibiting any others from
entering. Electrical changes that occur in an egg’s cell membrane help to keep other sperm cells
from penetrating the egg.
- Fusion of the genetic material of sperm and egg;
- Activation of egg metabolism to start development.
Fertilization can occur inside the body of the mother (internal fertilization like in mammals) or
outside the mother's body in water (external fertilization like in fishes, frogs etc.). Initially during
evolution, when life was only in water, there was no problem of leaving eggs without the fear of
them getting dried out. Internal fertilization developed in evolution when the animals started to
migrate onto the land from water.
1. Cleavage and blastulation
1.1. Cleavage
Fertilization is followed by the transformation of the diploid zygote into a mass of cells through a
rapid succession of cell divisions called cleavage or segmentation. It is now clear that cleavage
starts only after the egg is stimulated to begin its division. Cleavage produces a solid ball of
smaller and smaller cells, called a morula which is not larger than the zygote. The variety of egg
types result in different patterns of development so that variation is also seen in the
developmental steps of cleavage and subsequent stages (blastula, gastrula, neurula and
organogenesis).
The cleavage phase arbitrarily ends when this ball forms a central fluid-filled cavity called a
blastocoel, at which point the embryo is called a blastula. Its individual cells are called
blastomeres; the building blocks of future organs of developing embryo. As there is no growth
phase during cleavage (in other words, there is no G 1 or G2 phase of the cell cycle in this case),
so the size of the blastomeres decreases.

 Patterns of cleavage
Based on the potency or capability of blastomeres two main types of cleavage are classified as:
- Determinate cleavage: where the fate of each blastomere is predetermined as in most
protostomes (nematodes, annelids, molluscs, ascidian and cephalochordates). Damage to or
destruction of any of these cells results in malformation of an organism. These eggs are also
called mosaic eggs or plastic eggs.
- Indeterminate cleavage: where the blastomeres have similar developmental potential and, if
isolated, can each give rise to a complete individual embryo as in echinoderm and all the
vertebrate eggs. This type of cleavage occurs in the eggs which are without predetermined
regions. These types of eggs are called regulative eggs.
During cleavage or early blastula formation in mammals, for example, if the blastomeres are
physically separated into two groups, both groups can produce complete embryos (Fig.1.3).
Since the two embryos come from the same zygote, they will be monozygotic twins—genetically
identical. Non-identical twins occur when two separate eggs are fertilized by two separate sperm.
Thus, while identical twins are always of the same sex, non-identical twins have a 50 percent
chance of being the same sex.
 Figure 1.3. Twinning in Humans Because humans have regulative development, remaining cells can compensate
when cells are lost in early cleavages. Monozygotic (identical) twins can result when cells in the early blastula
become physically separated and each group of cells goes on to produce a separate embryo.

Based on the amount and distribution of yolk (=Stored food substances in the egg cell or in
developing embryo) there are different patterns of cleavage as:
 Holoblastic cleavage is total cleavage of the entire egg. It can be equal holoblastic when
blastomeres are of equal size (Amphioxus, mammals) or unequal holoblastic when blastomeres
are of unequal size (amphibians).
 Meroblastic cleavage is the incomplete cleavage due to inert dense inactive yolk in the
telolecithal in which the cleavage furrow stops short (or fails to form) and the ‘'daughter cells'’
retain cytoplasmic continuity.
It is of two types discoidal (fishes, reptiles, birds and monotremata) where the cleavage is
restricted to a small discoidal area over yolk free cytoplasm and superficial meroblastic cleavage
is in centrolecithal eggs as of insects where the cleavage is confined to peripheral cytoplasm of
egg and does not extend to central yolk (see types of blastula, Fig. 1.4).
Similarities in cleavage patterns in different animal groups provide evidence for the evolutionary
relationships as the cleavage patterns in reptiles, birds and later development similarities in
mammals point to their close ancestral relationships.

Table 1: Examples of cleavage patterns

Type of cleavage Type of egg Examples Illustration

Holoblastic cleavage [Link] Radial
(complete)
(sparce,evenly distributed (echinoderms,
yolk) amphioxus)

Rotational
(Mammals,
nematodes)

2. Mesolecithal Displaced radial

cleavage
(moderate vegetal yolk
disposition) (Amphibians)

Meroblastic cleavage 1. Telolecithal Bilateral cleavage

(incomplete) (dense yolk in most cells) (Cephalopod
molluscs)

Discoidal cleavage
(Fish, reptiles, birds)

2. Centrolecithal Superficial cleavage

(yolk in center of egg) (ex: most insects)

1.2 Blastula
Morula is succeeded by blastula where the blastomeres arrange themselves into epithelium,
called blastoderm. A cavity also appears underlying blastoderm which is called blastocoel or
segmentation cavity. It is important as it provides space for the morphogenetic movements.
Blastula is normally spherical in shape. As variety of egg types result in variation in cleavage
patterns so the different forms of blastula are also expected. Whatever be the variation in the
gross form, all have a common cause of constructing a stratified embryo called gastrula (gaster-
stomach) which is the next step in development process.
Types of blastula (Fig.1-4):
The types of blastulae greatly vary among different animals depending upon various factors such
as the egg size, the amount and distribution of yolk, and the rate and pattern of cleavage. The
various types of blastula can be classified into six types (see Fig.1-4):
1. Stereoblastula: Solid blastula characterized by the absence of a blastocoel as in annelids,
molluscs, nemerteans and Planaria.
2. Coeloblastula: Hollow spherical blastula surrounded by single layer of blastoderm
(echinoderm, Amphioxus)
3. Amphiblastula: Blastoderm is of unequal thickness made of micromeres and
macromeres with an eccentric blastocoel as in amphibia.
 4. Discoblastula: Multi layered flat disc separated from yolk by subgerminal cavity in
macrolecithal egg of fishes, reptiles and birds. Due to the presence of yolk, blastoderm
becomes arched and the floor is flattened.
5. Periblastula: Superficially cleaving egg of insect with peripheral blastomeres and yolk
filled central blastocoel.
6. Blastocyst: In mammals, the blastula has the blastoderm which draws food for the
yolkless embryo from the uterine wall. The blastula is called a blastocyst. It has a group
of embryo formative cells called inner cell mass with small fluid filled spaces
representing a place for food in the form of yolk in the ancestors. Blastoderm is called
trophoderm or trophoblast as it helps in embedding of embryo into uterus. Blastoderm is
the embryo formative region in invertebrates and in most of the chordates, but it is the
inner cell mass in mammals.

Figure 1.4: Types of blastula. (a) Stereoblastula (b) Coeloblastula (c) Amphiblastula (d) Discoblastula (e)
Periblastula (f) Blastocyst.

1.1. Cleavage and blastulation in Amphibia

In the case of the frog and most amphibians, the development of the embryo is complicated by
the fact that the egg contains an appreciable amount of yolk material that constrains the
divisions. Eggs with a small amount of yolk material evenly distributed throughout, as in starfish
eggs, are termed isolecithal eggs (from the Greek words isos meaning “equal, uniform’’ plus
lekirhos meaning “egg yolk”).
Amphibians, which have an appreciable amount of yolk which is unevenly distributed, produce
eggs referred to as moderately telolecithal (from the Greek word telos meaning “consummate,
complete”).
In the extreme telolecithal egg of birds, the mass of the yolk is so great that embryogenesis
involves only a small disk of cytoplasm at one end of the original egg cell.
In moderately telolecithal organisms such as frogs, a considerable degree of differentiation
marks the unfertilized egg. The upper, animal hemisphere is heavily pigmented, but the lower,
vegetal hemisphere is much denser and yolk-laden. Shortly after the attachment of the sperm, the
eggs orient themselves with the animal poles uppermost. Meiosis, arrested in metaphase II, is
activated by the sperm entry and proceeds to its completion.
Syngamy (union of the nuclei) will occur, and the first cleavage will be completed about two to
three hours after sperm entry. Early cleavage sections are complete, but the vegetal hemisphere
cells cleave at a slower rate. The cleavage of a fertilized and telolecithal frog egg is of
holoblastic and inequal type. The first cleavage is plane is meridional and results in in two equal
sized blastomeres. The second set of divisions is likewise holoblastic and equal, and the planes
are meridional, but at right angle to the first plane. But with the third set of divisions the
inequally distributed yolk makes it influence felt.
Egg yolk reduces the rate of cleavage therefore; vegetal blastomeres divide slowly and with
lesser frequency. At about fourth or fifth cleavage, a small space called blastocoel appears which
increases rapidly. As cleavage proceeds, the adhesion of the blastomeres to each other increases
and they arrange themselves into a true epithelium called blastoderm. This blastoderm remains
two-celled thick toward the animal pole of the egg and here, it forms a sort of fluid filled cavity
formed in the animal hemisphere: the blastocoel. This cavity will be important for allowing the
cell movements to occur during gastrulation. The sides and the floor of the blastocoel are
occupied by multilayered bastoderm of large yolky blastomeres. Thus, there is formation of
blastula which is a hollow ball, with a thin bi-layered roof of animal cells and floor of large
yolky blastomeres so called micromeres and macromeres respectively.

1.2. Cleavage and blastulation in Birds (chick)

Fertilization of the chick egg occurs in the oviduct, before the albumen and the shell are secreted
upon it. Blastula and gastrula stages occur while the egg is still located within the oviduct. The
egg is telolecithal (like that of the fish), with a small disc of cytoplasm sitting on top of the yolk
mass. Like fish eggs, the yolky eggs of birds undergo discoidal meroblastic cleavage. Cleavage
occurs only in the blastodisc, a small disc of cytoplasm 2-3mm in diameter at the animal pole of
the egg cell, the cleavage furrows do not penetrate the yolk. The first cleavage furrow appears
centrally in the blastodisc, and other cleavages follow to create a single layered blastoderm. As
in fish embryo, these cleavages do not extend into the yolky cytoplasm. This type of incomplete
cleavage, called discoidal cleavage, is common in fishes, reptiles, and birds.
 Figure 1-5: Discoidal meroblastic cleavage in a chick egg. (A-D): Four stages viewed from the animal pole (the
future dorsal side of the embryo); (E): an early cleavage cleavage embryo viewed from the side stages.
Thereafter, equatorial and vertical cleavages divide the blastoderm into a tissue five to six layer
thick. Between the blastoderm and the yolk is a space called the subgerminal cavity created at
the time the blastoderm cells absorb water from the albumen and secrete the fluid between
themselves and the yolk. At this stage, the deep cells in the center of the blastoderm are shed and
die, leaving behind one-cell-thick area pellucida. This part of blastoderm forms most of the
actual embryo. The peripheral ring of blastoderm cells that have not shed their deep cells
constitutes the area opaca. Between the two areas is a thin layer cells called the marginal zone
(see also Fig.1-5).

Figure 1-6: Discoidal meroblastic cleavage in a chick egg. (a-c): view from the animal pole; (d-f): section through the
blastoderm at different cleavage stages.
As the fertilized egg passes downward inside the oviduct two events occur simultaneously, firstly, it
undergoes the process of cleavage; blastulation and even gastrulation; and secondly various accessory
coverings (= vittelline membrane, albumen or white of the egg, outer and inner shell membrane,
calcareous membrane) are added to the developing egg by the oviduct wall.

Figure 1-7: Diagram of a newly-laid hen egg.

1.3. Cleavage and blastulation in mammals

Mammalian eggs are among the smallest in the animal kingdom, making them hard to
manipulate. The human zygote for instance, is only 100µm in diameter (less than one-thousandth
the volume of a frog egg). It is also difficult to obtain enough material for studies as the
mammalian zygotes are not produced in numbers compared to sea urchin or frog.
It is difficult to obtain eggs and embryos in man for experimentation though it can be obtained
by sacrificing pregnant females in other mammals for in vitro studies.
New techniques have developed for in vitro studies that have made astonishing advancements in
mammalian embryology. Heavy doses of gonadotrophins are given to volunteer females that
cause ripening of ovarian follicles. By a small incision, an aspirator and laparoscope is
introduced into the abdomen and follicles are sucked up. They are grown and fertilized in vitro
for developmental studies. After in vitro fertilization, the egg has to be introduced into the uterus
of female because of embryo’s dependence on it. Reimplantation is done in pseudo pregnant
uterus of female. This method allows production of test tube babies.

Cleavage in mammals is unique; several features of mammalian cleavage are very different from
those seen in other animal groups.

Cleavage in mammals is very slow; cell divisions are 12–24 hours apart, compared with tens of
minutes to a few hours in non-mammalian species. The oocyte is released from the ovary and
swept by the fimbriae into the oviduct and fertilization occurs in the ampulla of the oviduct.
Meiosis is completed at this time, and first cleavage begins about a day later. Meanwhile, the
cilia in the oviduct push the embryo toward the uterus; the first cleavage occurs along this
journey.
The pattern of cleavage in mammals is rotational: The 1st cleavage is normal meridional
division, yielding two blastomeres. ; however, in the second cleavage, one of the two
blastomeres divides meridionally and the other divides equatorially. (Fig. 1.8).

Figure 1.8: Mammals have rotational cleavage, in Figure 1.9: Comparison of early cleavage in (A)
which the plane of the first cleavage is parallel to the echonoderms and amphibians (radial cleavage)
animal–vegetal (A,V) axis, but the planes of the and (B) mammals (rotational cleavage)
second cell division (shown in beige) are at right
angles to each other.

The cell divisions of mammalian blastomeres are not in synchrony with each other. Because the
blastomeres do not undergo mitosis at the same time, the number of cells in the embryo does not
progress in the regular (2, 4, 8, 16, 32, etc.) progression typical of other species, but frequently
contain odd numbers of cells.

Another unique feature of the slow mammalian cleavage is that the products of genes expressed
at this time play roles in cleavage. In species such as sea urchins and frogs, gene expression does
not occur in the blastomeres, and cleavage is directed exclusively by molecules that were present
in the egg prior to fertilization.
  Formation of the blastocyst (mammal blastula):

As in other animals that have complete cleavage, the early cell divisions in a mammalian zygote
produce a loosely associated ball of cells. However, at about the 8-cell stage, the behavior of the
mammalian blastomeres changes. They change shape to maximize their surface contact with one
another, form tight junctions, and become a very compact mass of cells (Fig.1-10).

At the transition from the 16-cell to the 32-cell stage, the cells separate into two groups. The
inner cell mass will become the embryo, while the surrounding cells become an encompassing
sac called the trophoblast, which will become part of the placenta. Trophoblast cells secrete
fluid, creating a cavity (blastocoel) with the inner cell mass at one end (Fig.1-10). At this stage,
the mammalian embryo is called a blastocyst to distinguish it from the blastulas of other
animals.

Figure 1.10. The Mammalian Zygote Becomes a Blastocyst : Starting late in the 8-cell stage, the
mammalian embryo undergoes compaction of its cells, resulting in a blastocyst—a dense inner cell mass
on top of a hollow blastocoel, completely surrounded by trophoblast cells.

Examples:
The ovum of placental mammals is alecithal (i.e possess only a negligible amount of yolk) and undergoes
holoblastic cleavage.

(A) (B) (C) (D) (E) (F)

Figure 1.11: The cleavage of a single mouse embryo in vitro. (A) 2-cell stage (B) 4-cell stage. (C) 8-cell stage. (D) Compacted
8-cell stage. (E) Morula (F) Blastocyst.
In man, the first three days are spent by the fertilized egg in descending through the uterine tube
during which time the cleavage starts. Fertilization occurs in the upper reaches of the mother’s
oviduct, and cleavage occurs as the zygote travels down the oviduct to the uterus.
 Figure 1.12. Development of a human embryo from fertilization to implantation (0-6 days). The embryo hatches
from the zona pellucida upon reaching the uterus. During its migration to the uterus, the zona prevents the embryo
from prematurely adhering to the oviduct rather than traveling to the uterus.

By 3rd day morula is formed. At the eight cell stage, the blastomeres develop differential
adhesion properties so, the outer cell surface becomes convex and inner surface becomes
concave. This reorganization is called compaction which involves the activity of cytoskeletal
components and adhesion of blastomeres.
The development of differential adhesion between blastomeres results in segregation of cells as
inner cell mass and outer cell mass or embryonic knob. Inner cell mass is also called
embryoblast as it gives rise to the embryo proper.
By 4th day, the cells begin to absorb fluid which is collected in the blastula so it is the called
blastocyst. As the blastocyst moves down the oviduct to the uterus, it must not embed itself in
the oviduct wall. Early implantation is normally prevented by an external proteinaceous layer
called the zona pellucida, which surrounds the egg and remains around the cleaving ball of cells.
By the 5th day the blastula hatches from the zona pellucida and later on embeds in uterine wall
(this is called implantation). As long as zona pellucida is present the embryo cannot attach to
the uterine wall. When the blastocyst arrives in the uterus, the trophoblast adheres to the
endometrium (the uterine wall). This event begins the process of implantation that embeds the
embryo in the wall of the uterus ( Fig. 1.12). In humans, implantation begins on about the sixth
day after fertilization.
 Figure 1.13: Tissue formation at 7 and 8 days after. A–B: Human blastocyst immediately prior to gastrulation. The inner
cell mass delaminates hypoblast cells that line the blastocoel, forming the extraembryonic endoderm of the primitive yolk
sac and a two-layered (epiblast and hypoblast) blastodisc similar to that observed in avian embryo. The trophoblast
divides into the cytotrophoblast which will form the villi and the synciotrophblast which will ingress in the uterine tissue.

The blastocyst is composed of: the cells of inner cell mass which give rise to the three germ
layers so they are the formative cells and the trophoblast or trophoderm, a layer of cells
surrounding the inner cell mass and the blastocyst cavity. The trophoblast combines with the
maternal endometrium to form the placenta in eutherian mammals. The cells of trophoblast
multiply rapidly and push into the uterine wall. The cytolytic action of trophoblast breaks the
uterine tissue which is used by the embryo as food till it develops placenta. Sooner, a cavity
appears in between inner cell mass and trophoblast. The fluid is imbibed in this cavity and
trophoblast is lifted off the egg but it remains attached from one side (called Rauber’s layer)
which corresponds to the dorsal side of embryo. Hypoblast layer separates from inner cell mass
and lines the cavity.

Occasionally the blastocyst also implants in the peritoneal cavity or on the surface of ovary or on
the other abnormal site. The epithelium at that region responds with the increased vascularity so
that the blastocyt starts its development. This is called ectopic or tubal pregnancy which may
sometimes threaten the life of mother as blood vessels at these sites are ruptured due to the
growth of embryo.
When inner cell mass is isolated and grown under certain conditions, the cells remain
undifferentiated and divide in culture (Martin, 1981). They are called embryonic stem cells,
which are now days exploited for therapeutic use. They are called embryonic stem cells as each
of them can generate all the cells of embryo when separated (see indeterminate cleavage).

3. Gastrulation
3.1. Establishment and significance of gastrulation

Gastrulation is the process whereby the blastula is transformed by massive movements of cells
into a gastrula with multiple tissue layers (also called cell layers or germ layers) and visible
body axes.
Gastrulation begins as soon as blastulation is accomplished. The mechanism of this process
depends on the type of the blastula type. In most phyla, the gastrula does not remain a two-layer
structure. As development progresses, a third layer, the mesoderm, forms between the endoderm
and the ectoderm. Gastrulation thus results in the formation of a three-layered embryonic
structure with an established longitudinal axis and a general outline of the broad features of the
adult. Each of the three primary germ layers (ectoderm, endoderm, and mesoderm) will soon
produce a variety of differentiated structures. The gastrulation has two- fold significances, which
are rearrangement of cells which later on help in neurogenesis and the formation of different
organs and systems.
3.2. General processes involved in gastrulation

Gastrulation is an integrated, dynamic process, controlled largely by intrinsic forces bound in the
specific physico-chemical conditions of the various presumptive organ forming areas of the late
blastula and early gastrula. These forces, in turn, are correlated with external conditions. In
vertebrates, it includes different kinds of morphogenetic movements.

 Morphogenetic movements

Gastrulation is the compilation of different types of cellular movements which creates a

trilaminar gastrula exhibiting variation in blastulation. These formative or morphogenetic
movements may express themselves in various ways. The basic movement is of spreading and
stretching of cells and most of the technical terms used are related to variations in these two
movements. The net result of all these movements is the formation of gastrula.

• Epiboly means (in the case of frog gastrulation) spreading of micromeres over macromeres or
movement of presumptive ectodermal cells by stretching and spreading.
• Emboly is inward movement of cells which is of various types. Embolic movements result in
relocation of the cells of the external surfaces to the interior and expansion of remaining cells to
replace the parts moved to interior.
- Invagination means infolding or insinking of egg surface followed by migration of cells.
- Involution is the inturning or rotation of cellular materials upon itself so that it is directed
inside.
- Infiltration is movement of cells to form a new layer, hypoblast.
- Delamination is separation of cells by splitting of layer.
- Convergence is movement of cells towards midline to form axial structures.
- Divergence : as the cells after involution do not remain as heap of cells but move to future
position so it is called divergence
- Extension is the elongation of gastrula along anterior posterior axis.
- Ingression is movement of individual cells or small group of cells into the blastocoel.
- Constriction is reduction in size of blastopore.

Figure 1.14: Illustration of some morphogenetic movements

3.3. Gastrulation in sea urchin (Echinoderm)

Invagination of the vegetal pole characterizes gastrulation in the sea urchin. In forms with little yolk like
echinoderm (with coeloblastula) gastrulation is an inpocketing of the blastula (see Fig.1.15). Gastrulation
is marked by changes in the shape of cells and the way the cells interact with each other. As the inward
folding continues, the now cup-shaped embryo enlarges, and a deep cavity, called the archenteron, or
primitive gut, develops. The open end of the archenteron is called blastopore. Forming the outer layer of
the gastrula is the outer germ layer, the ectoderm. The inner germ layer is the endoderm.
Figure 1.15: Gastrulation in sea urchin (echinoderm). The blastula reorganizes and forms the cup-shaped gastrula. (Note other
phyla have somewhat different patterns of gastrulation).

Invagination of the vegetal plate in the single-layered blastula forms the blastopore, which will become
the anus. Further differentiation yields a gastrula with three embryonic tissue layers.

 Endoderm: Infolding forms the archenteron, which later develops into the digestive tube with a
second opening that becomes a mouth.
 Mesoderm: Mesenchyme cells migrate into the blastocoel, and will later form the skeleton.
 Ectoderm.
Finally, the gut, ectoderm, and the skeleton undergo further obvious differentiation to make the pluteus
larva (see fig. 1.16).
 Figure 1. 16: Gastrulation in sea urchin embryo

3.4. Gastrulation in Amphibia (Frog)

Mature egg of frog is 2.0-3.0 mm in diameter and is mesolecithal. It is radially symmetrical prior
to fertilization but later on develops dorsoventral symmetry due to internal cytoplasmic
rearrangement that forms gray crescent (dorsal side) a forerunner of notochord. The cleavage is
holoblastic and for the first two divisions it is complete but later on becomes unequal due to
yolk. Egg yolk reduces the rate of cleavage therefore; vegetal blastomeres divide slowly and with
lesser frequency.
At about fourth or fifth cleavage, a small space called blastocoel appears which increases rapidly.
Thus, there is formation of blastula which is a hollow ball, with a thin multi-layered roof of
animal cells and floor of large yolky blastomeres so called micromeres and macromeres
respectively.

Figure 1.17: Frog blastula stage

During pregastrulation, all material for different organs disappears from the surface of blastula
and come inside to take their final position in the embryo where different organs develop from
their respective prospective areas.
In many animals, these movements of the blastomeres are so regular and well orchestrated that it
is possible to label a specific blastomere with a dye and identify the tissues and organs that form
from its progeny. Such labeling experiments produce fate maps of the blastula (Fig.1-18). The
fate map of frog blastula has been prepared by vital dyes which gives the description of what the
particular area is destined to become later.
Figure 1.18: Fate Map of a Frog Blastula: The colors indicate the portions of the blastula that will form
the three germ layers, and subsequently the frog’s tissues and organs. Frog blastula-stage fate map
Amphibian gastrulation begins when certain cells in the gray crescent change their shape and
their cell adhesion properties. The main bodies of these cells bulge inward toward the blastocoel
while they remain attached to the outer surface of the blastula by slender necks. Because of their
shape, these cells are called bottle cells. The bottle cells mark the spot where the dorsal lip of the
blastopore will form (Fig.1.19). As the bottle cells move inward, they create this lip, over which
successive sheets of cells will move into the blastocoel in a process called involution.
The first involuting cells are those of the prospective endoderm, and they form the primitive gut,
or archenteron.
Closely following are the cells that will form the mesoderm.

As gastrulation proceeds, cells from the animal hemisphere, the future ectoderm, change shape,
move over the outer surface toward the site of involution in a process called epiboly. The
blastopore lip widens on both sides of the embryo and eventually forms a complete circle
surrounding a “plug” of yolk-rich cells. When the sides of the lip meet, the blastopore forms a
circle that becomes smaller as ectoderm spreads downward over the surface.

As cells continue to move inward through the blastopore, the archenteron grows, gradually
displacing the blastocoel. As gastrulation comes to an end, the amphibian embryo consists of
three germ layers: ectoderm on the outside, endoderm on the inside, and mesoderm in the
middle. The embryo also has a dorsal– ventral and anterior–posterior organization. Most
importantly, however, the fates of specific regions of the endoderm, mesoderm, and ectoderm
have been determined.
At the beginning, the blastopore lies in the pigment free prospective endoderm and by the time it
forms a complete circle, dark, pigmented cells of animal hemisphere moves into the interior
except for the large group of yolk cells crowded at the blastopore called yolk plug. The size of
the archenteron increases obliterating the original blastocoel. The lips of blastopore or rim of
blastopore are not an anatomical entity but only constantly changing structural expression of
turning of surface cells inside. The cells of the blastopore do not simply pile up but move
through the blastopore lips to the interior and are redistributed. Thus, the cells at the rim of
blastopore undergo invagination. Once inside, the cells take different routes such as the
notochordal and somatic mesoderm cells stream upward and stretch out along the longitudinal
axis; and the lateral and ventral mesoderm spreads out in forward and downward direction
between ectoderm and endoderm.
The dorsal lip of blastopore is the most important and powerful self-differentiating group of cells
or organiser in the amphibian gastrula.
Figure 1.19: Gastrulation in a frog embryo. In the frog blastula, the blastocoel is displaced toward the
animal pole and is surrounded by a wall several cells thick. The cell movements that begin gastrulation
occur on the dorsal side of the blastula, where the gray crescent was located in the zygote. Although still
visible as gastrulation begins, the gray crescent is not shown here.

Throughout the gastrulation, the embryo retains its uniform size and shape because as soon as the
prospective endoderm and mesodermal cells move from the surface, their place is taken over by
ectoderm cells which undergo epiboly or stretching. So there is pronounced thinning and
stretching of ectodermal cells and to compensate for the increased surface area the blastoderm
changes from thick layer to a thin layer. Because of rapid epiboly by prospective ectoderm cells
and NIMZ (Non Involuting Marginal Zone) blastopore is shifted posteriorly. The contracting
blastopore withdraws the yolk plug completely into the interior which marks the end of
gastrulation movement in frog.

The steps in the amphibian gastrulation are:

• Invagination by change in shape of the cells which is apparent as a small slit in beginning
below the gray crescent area.
• It is accompanied by involution.
• Blastopore changes shape and its final form is a circle.
• Epiboly occurs simultaneously with emboly.
• The blastocoel is obliterated by archenteron. The chordamesodermal cells move as a mass
called chordamesodermal mantle.
• The termination of gastrulation is by contraction of blastopore and its subsequent shift to
posterior position.
3.5. Gastrulation in Birds (Chick)

The eggs of reptiles and birds contain a mass of yolk, and therefore the blastulas of these species
develop as a disc of cells on top of the yolk (Fig.1. 20). We will use the chicken egg to show
how gastrulation proceeds in a flat disc of cells rather than in a ball of cells (Fig.1. 21).

Cleavage in the chick results in a flat, circular layer of cells called a blastodisc. Between the
blastodisc and the yolk mass is a fluid-filled space. Some cells from the blastodisc break free and
move into this space. Other cells grow into this space from the posterior margin of the blastodisc.
These cells come together to form a continuous layer called the hypoblast, which will later give
rise to extraembryonic membranes that will support and nourish the developing embryo. The
overlying cells make up the epiblast, which will form the embryo proper. Thus, the avian
blastula is a flattened structure consisting of an upper epiblast and a lower hypoblast, which are
joined at the margins of the blastodisc. The blastocoel is the fluid-filled space between the
epiblast and hypoblast.

Gastrulation begins with a thickening in the posterior region of the epiblast caused by the
movement of cells toward the midline and then forward along the midline (Fig. 1.21). The result
is a midline ridge called the primitive streak.

A depression called the primitive groove forms along the length of the primitive streak. The
primitive groove functions as the blastopore and epiblast cells migrate through it downward into
the blastocoels to form mesoderm and contribute to endoderm formation together with hypoblast
cells. The hypoblast will also form the yolk sac, an extra-embryonic compartment containing
food material.
 In the chick embryo, no archenteron forms, but the endoderm and mesoderm migrate forward to
form the gut and other structures. At the anterior end of the primitive groove is a thickening
called Hensen’s node, which is the equivalent of the dorsal lip of the amphibian blastopore. In
fact, many signaling molecules that have been identified in the frog organizer are also expressed
in Hensen’s node.

Cells that pass over Hensen’s node become determined by the time they reach their final
destination, where they differentiate into certain tissues and structures of the head and dorsal
midline (but not the nervous system).

Figure 1.20: The chicken egg with a disc of cells on top of the yolk
Figure 1.21: Gastrulation in Birds Because of the large yolk mass in bird and reptile eggs, these
embryos display a pattern of gastrulation very different from that of sea urchins and amphibians.

Figure 1.22: Longitudinal section showing formation of mesoderm

(A) (B)
Figure 1.23: Cell movements in the primitive streak of the chick embryo. (A-C) Dorsal view of the formation and
elongation of the primitive streak 3-4 hrs (A), 7-8hrs (B) and 15-16hrs (C) after fertilization. (D) 19-22hrs:
Formation of notochord and mesodermal somites as primitive streak regresses. Fate map of the chick epiblast is
shown at the definitive streak stage (the neurulation stage is not shown in this figure)
Figure 1.24: Chick embryo after 24
hours. Development of neural tube
and mesoderm; formation of neural
tube following regression of the
primitive streak. The cephalic (head)
region has undergone neurulation,
while the caudal (tail) regions are still
undergoing gastrulation.
Figure 1.25: Chick gastrulation 24-26hrs after fertilization (A) the primitive streak at full extension (24hrs). The
head process (anterior notochord) can be seen extending from Hensen’s node. (B) Two-somite stage (25hrs).
Pharyngial endoderm is seen anteriorly, while the anterior notochord pushes up the head process beneath it. The
primitive streak is regressing. (C) Four-somite stage (27hrs). (D) 28hrs the primitive streak has regressed to the
caudal portion of the embryo.
3.6. Gastrulation in mammal (man)
The inner cell mass splits into an upper layer called the epiblast and a lower layer called the
hypoblast, with a fluid-filled cavity between them. The embryo will form from the epiblast, and
the hypoblast will contribute to the extraembryonic membranes (Fig. 1.26). The epiblast also
contributes to the extraembryonic membranes; specifically, it splits off an upper layer of cells
that will form the amnion. The amnion will grow to surround the developing embryo as a
membranous sac filled with amniotic fluid. Gastrulation occurs in the mammalian epiblast just as
it does in the avian epiblast. A primitive groove forms, and epiblast cells migrate through the
groove to become layers of endoderm and mesoderm.

Figure 1.26: A Human Blastocyst at Implantation Adehesion molecules and proteolytic enzymes secreted by
trophoblast cells allow the blastocyst to burrow into the endometrium. Once implanted within the wall of the uterus,
the trophoblast cells send out numerous projections—the chorionic villi—which increase the embyro’s area of
contact with the mother’s bloodstream.

As the blastocyst completes implantation during the second week development, the embryoblast
undergoes marked differentiation. A slitlike space called the amniotic cavity forms between the
embryoblast and the trophoblast. The embryoblast flattens into the embryonic disc which
consists of two layers: an upper ectoderm, which is closer to the amniotic cavity, and a lower
endoderm, which borders the blastocyst cavity. A short time later, a third layer, the mesoderm
forms between the endoderm and the ectoderm. These three layers constitute the primary germ
layers.
Figure 1.27: Tissue formation between days 9 and 11. (C) The inner cell mass delaminates hypoblast cells thet line the
blastocoel, forming the extraembryonic endoderm of primitive yolk sac and a two-layered (epiblast and hypoblast) blastodisc
similar to that of avian embryos. The trophoblast divides into the cytotrophoblast which will for, the villi, and the
synciotrophoblast, which will ingress into the uterine tissue. The epiblast splits into the amnionic ectoderm a the embryonic
epiblast. (D) The extraembryonic endoderm forms the yolk sac.

Figure 1.28: Amnion formation and cell movement during human gastrulation. (A, B): human embryo and uterine connection at
day 15 of gestation (A) sagittal section through the midline. (B) view looking down on the dorsal surface of the embryo. (C) The
movement of the epiblast cells through the primitive streak and Hensen’s node and underneath the epiblast are superimposed on
the dorsal surface view.
Figure 1.29: Schematic diagram showing the derivation of tissues in human and rhesus monkey embryo.

Comparative study of gastrulation in different animals enables us to understand thoroughly the

complicated process. These movements depend on the type of egg, the type of blastula and mode
of development. The similar development process in birds and mammals, though the differences
exist in the egg types, throws light on the fact that both have descended from the common
reptilian ancestors.

Mammalian gastrulation has some peculiar features:

1. It follows the similar gastrulation process as that of bird and reptile though it is having
negligible amount of yolk in the egg. Small primitive streak is formed as compared to that of
bird and reptile.
2. The blastoderm differentiates into trophoblast and inner cell mass.
3. Inner cell mass forms the complete embryo so the gastrulation is restricted in this region
4. Foetal membranes, a characteristic feature of amniotes, start forming before gastrulation
because of the immediate requirement of food by the growing embryo. This is a prerequisite
for the formation of placenta in viviparous mammals.
3.7. Extra-embryonic membranes and fetal membranes
The extraembryonic membranes function in nutrition, gas exchange, and waste removal.

 In birds and most reptiles, the embryo with its extra-embryonic membranes develops within a
shelled egg.

Figure 1.30: Schematic drawings of the extraembryonic membranes of the chick. The embryo is cut longitudinally and the
albumen and shell are not shown.

Figure 1.31: The extra-embryonic

structures and circulation of the chick
embryo

 The amnion surrounds the embryo, forming the amniotic cavity. The amnion secretes
fluid called amniotic fluid into the cavity, providing a protective environment for the
embryo. (against chock, injury and desiccation)
  The yolk sac contains yolk - the sole source of food until hatching. Yolk is a mixture of
proteins and lipoproteins. The yolk sac is the first extraembryonic membrane to form.
The yolk sac grows to encloses the entire body of yolk in the egg

 The chorion forms a continuous membrane just under the eggshell. It limits water loss
from the egg and also works with the enlarged allantoic membrane to exchange
respiratory gases (O2 and CO2) between the embryo and the outside world.

 The allantois stores metabolic wastes (chiefly uric acid) of the embryo and, as it grows
larger, also participates in gas exchange.

In mammalians the four extra-embryonic membranes or fetal membranes develop to cover and
protect the embryo. The structure, development and function of these membranes are more or
less similar to those of birds and reptilians with slight modifications.
The membranous structure that surrounds the developing fetus and forms the amniotic cavity is
derived from fetal tissue and is composed of two layers: the amnion (inner layer) and the chorion
(outer layer). The amnion is a translucent structure adjacent to the amniotic fluid, which provides
necessary nutrients to the amnion cells. The chorion is a more opaque membrane that is attached
to the decidua (i.e. maternal tissue that lines the uterus during pregnancy). The amnion and
chorion are separated by the exocelomic cavity until approximately three months gestation, when
they become fused. Intact, healthy fetal membranes are required for an optimal pregnancy
outcome.

The allantois of mammals is composed splanchnopleure (endoderm+splanchnic mesoderm). It

arises from the hind gut as a small sac which gradually increases in the extraembryonic coelom
i.e space between the amnion and chorion. Soon, it grows over the embryo outside the amnion
where it enlarges. Later on, mesodermal component of allantoic wall give rise to the blood vessel
system of the allantois and fuses with mesoderm of chorion to form of allantoic chorion.
 Figure 1.32: Formation of fetal membranes and umbilical cord in mammals

The narrow connecting stalk of extra-embryonic mesoderm that links the embryo to the
trophoblast eventually forms the vessel of the umbilical cord.
The fully developed extra embryonic organ, consisting of trophoblast tissue and blood vessel-
containing mesoderm, is called the chorion, and it fuses with the uterine wall to create the
placenta. Thus, the placenta has both the maternal portion (the uterine endometrium, which is
modified during pregnancy) and a fetal component (the chorion); the chorion may be closely
opposed to maternal tissues while still being readily separable from them (ex. in pig) or it may be
so intimately integrated with maternal tissues that the two cannot be separated without damage to
both the mother and the developing fetus (as in the deciduous placenta of most mammals,
including humans).
The placental functions are nutritional, endocrine and immunological. The placentas provide
hormones that enable the uterus to retain the pregnancy and also accelerate mammary gland
development. Placentas also block the potential immune response of the mother against the
developing fetus. The umbilical cord is a composite structure formed by contributions from:
Fetal connecting (body) stalk, yolk sac and amnion. The embryo is attached to the chorionic
placenta. It is through the blood vessels of the umbilical cord that nutrients and oxygen from the
mother reach the developing fetus and wastes, including carbon dioxide and urea, are removed.

(A) (B)
Figure 1.33: (A) Relationship of the chorionic villi to the maternal blood supply in the uterus. In the umbilicus,
there are two arteries (the umbilical arteries) and a single vein (the umbilical vein), buried within Wharton's jelly It
contains the right and left umbilical arteries, the left umbilical vein, and mucous connective tissue. The umbilical
vein supplies the fetus with oxygenated, nutrient-rich blood from the placenta. Conversely, the fetal heart pumps
deoxygenated, nutrient-depleted blood through the umbilical arteries back to the placenta. (B). Placenta. The
placenta is an organ that connects the developing fetus to the uterine wall to allow nutrient uptake, waste
elimination, and gas exchange via the mother's blood supply.
With the four fetal membranes, the developing embryo is able to carry on essential metabolism
while sealed within the egg. Surrounded by amniotic fluid, the embryo is kept as moist as a fish
embryo in a pond.
Although (most) mammals do not make a shelled egg, they do also enclose their embryo in an
amnion. For this reason, the reptiles, birds, and mammals are collectively referred to as the
amniota.

Figure .1.34: Extra-embryonic membranes in amniota

Mammals fall into three groups that differ in the way they use the amniotic egg.
 Monotremes [Prototheria eg. spiny anteater (echnid) and duckbill platypus]: lay eggs
 Marsupials (Metatheria eg. Kangaroo) the young are born in a very immature state as
the interface between the tissues of the uterus and the extra-embryonic membranes never
become developed; the placenta of marsupials is very short-lived and does not make as
much of a contribution to fetal nourishment as in other placental mammalians.
 Placental mammals (Eutheria, eg. humans): the extra-embryonic membranes form a
placenta and umbilical cord, which connect the embryo to the mother's uterus in a more
elaborate and efficient way.

3.8. Gastrulation: protostomes versus deuterostomes

In amphibia the blastopore is reduced to a small slit called proctodeum which joins the
endodermal outgrowth to form a cloacal aperture. It is called anus in higher vertebrates. The
archenteron forms pharynx and oral evagination extends from it towards epidermis. An ingrowth
of epidermis referred to stomodeum fuses with the oral evagination to form oral plate. This oral
plate is perforated to form mouth. The fate of blastoderm into anal or oral apertures is also one of
the criteria for classifying the animals into protostomes (mostly invertebrates) and
deuterostomes (echinoderms, chordates.)
After gastrulation, the blastopore of protostomes (first mouth) divides to form both mouth and
anal opening whereas in deuterostomes, the mouth arises at a different place away from site of
blastopore and the anus forms at the site of blastopore.
The protostomes division of metazoans embraces flat worms, annelids, molluscs and arthropods.
The deuterostome division of metazoans includes echinoderms and chordates.

Figure1. 35: Protostomic versus deuterostomic development.

Note: The Deuterostomes also show indeterminate development, in which each of the cells of the eight-
cell embryo if separated remain capable of developing as complete organisms, in contrast to determinate
development, in which the development fate of each cell in the adult organism has already been
determined.

4. Post gastrulation or organogenesis: Differentiation of

primary germ layers into tissues and organs

Gastrulation produces an embryo with three germ layers that are positioned to influence one
another through inductive interactions. During the next phase of development, called
organogenesis, many organs and organ systems develop simultaneously and in coordination
with one another. An early process of organogenesis that is directly related to gastrulation is
neurulation, the initiation of the nervous system in vertebrates. Neurulation involves the
formation of an internal neural tube from an external sheet of cells.
4.1 The formation of the neural tube
Neural tube is developed from prospective neuroectoderm which during pregastrular processes
takes its position somewhere in the mid dorsal region of the gastrula. The first signs of
neurulation are flattening and thickening of the ectoderm overlying the notochord; this thickened
area forms the neural plate. The neural plate is formed by the thickening of neuroectodermal
cells.

The ectoderm on the sides of the ventral plate rises as a pair of neural folds. The edges of neural
folds are developed by a process of differential growth and migration of cells called the
convergence. The plate between the folds sinks downwards and forms the neural groove. The
elevation of neural folds increases dorsally and ultimately fuses over the neural groove to form
the neural tube and a continuous overlying layer of ectoderm. The closure of the neural tube
begins just in front of the mid region and proceeds both interiorly and posteriorly. The neural
tube develops bulges at the anterior end, which become the major divisions of the brain; the rest
of the tube becomes the spinal cord.
The presumptive neural ectoderm region becomes distinct from the non-nervous ectoderm when
inductive signals are given by the underlying mesodermal cells on the ectoderm. Following these
instructions, the cells become columnar neural plate cells. The shaping of neural plate involves
the forces intrinsic to neural plate cells. The midline neural plate cells also known as Median
Hinge Point (MHP) cells, are the most important in shaping the neural plate. The bending of
neural plate is by both the intrinsic and extrinsic factors. MHP cells are anchored to the
notochord cells below and form a hinge which results in furrowing in the middorsal line of the
neural plate. This is mainly due to the induction by the notochordal cells on the overlying cells to
reduce the height of cells by changing them to wedge shape. The furrowing of neural plate
facilitates the formation of neural folds which fuse at the dorsal midline to form the neural tube.
The neural tube encloses a space called neural canal or neurocoel which opens anteriorly and by
neuropore but the opening is closed posteriorly due to the complete fusion of medullary folds.
In certain vertebrates it may communicate with the archenteron as neurenteric canal. The cells
which do not participate in the formation of neural tube, form neural crest cells. The formation
of neural tube does not occur simultaneously throughout the ectoderm e.g. in the bird’s
neurulation starts forming at the anterior end earlier than that at caudal end.
 a. Neural plate stage

b. Neural fold stage

c. Neural crests

d. Neural tube stage

Figure 1.36: Neural tube formation through four stages (a-d)

Unlike the neural tube closure in chicks, in mammals it is initiated at several places along the anterior
posterior axis. In humans, failure of the neural tube to develop normally can result in serious birth defects.
Failure to close the neural tube in posterior region results in abnormal condition of spine called spina
bifida or if it fails to close at anterior region it result in anencephaly. (The neural tube closure which is
very important step of neurulation is a result of genetic and environmental factors).
 Figure1.37: Failure of neural tube closure in human

4.2 Fate of germ layers in different animal groups

Diploblastic organisms
Gastrulation is important in laying down the primary germ layers of the embryo. The simple organisation
of embryo when it changes from monolayered to bilayered and trilayered is reminiscent of ancestral adult
conditions that occur in primitive groups of invertebrates. Cnidarians and ctenophores have only two
germ layers that are outer ectoderm and inner endoderm so they are called diploblastic but the germ
layers are called primary germ layers.

Figure 1. 38: General organization of a cnidarian.

Triploblastic organisms

In Bilateria (all animals having a bilateral symmetry) which includes few invertebrates starting
from platyhelminthes to all the chordates, three germ layers are laid and are called secondary
germ layers.
The relationship of these layers is also evident from their names, the outer layer ectoderm, the
middle layer mesoderm and the endoderm as the inner layer. The germ layers are the starting
point of variation which different classes of animals have developed but with the common lay
out plan of general body structure of vertebrate group as a whole. It also marks the transition
from simple increase in number of cells to developmental differentiation as within the layer
localized group of cells with different development potentials become apparent. This is evident
from the fact that gastrulation is followed by neurulation, notogenesis and mesogenesis. The
development of notochord from chordamesoderm is called notogenesis and the separation of
mesoderm is called mesogenesis.
Ectodermal cells in front of the blastopore or in front of the primitive streak in chick and
mammals begin to divide rapidly to form thick plate of ectoderm followed by neural fold and
neural tube. Neural tube differentiates anteriorly as brain and posteriorly as spinal cord. Neural
crest cells form few ganglion cells or pigment cells. The cells of roof of archenteron form the
notochord below the neural tube.

Figure 1. 39: Cross section in the embryo showing the neural tube and somite formation. (A) Beginning of somite
formation; (B) Seven-somite embryo.
 Figure 1.40: Diagram showing the
neural tube and somite formation

On the sides of the notochord, the mesoderm spreads out in the form of sheet. It grows down on
either side of gut and meets in the midventral line. The mesodermal sheet splits into two layers,
the outer layer near the ectoderm as somatic mesoderm and the inner layer close to endoderm is
the splanchnic mesoderm. The space between the two is coelom. At the same time, the sheet
gets sub divided horizontally into three distinct regions.
1. Dorsal segment is epimere
2. Middle segment mesomere
3. Ventral segment is hypomere

The lateral wall of the epimere is called dermatome. It forms dermis and dermal derivatives, its
dorsal median wall is called sclerotome. It forms the units of vertebral column. The ventro
median wall is myotome that forms the skeletal muscles and appendicular skeleton.
Somatopleure is the term for the somatic mesoderm and ectoderm whereas splanchnic
mesoderm and endoderm form splanchnopleure.
Mesodermal tissue is important as besides forming different organs it also forms the body cavity
or coelom. The coelom is the cavity which encloses various visceral organs. These organs are
held in position by the mesodermal fold called peritoneum.
Coelom formation protostomes is by the splitting of mesodermal tissue which creates a space or
cavity in which organs are placed. Lower deuterostomes on the other hand have different method
of coelom formation. The archenteron evaginates to form enterocoelic pouches. The cavity of
pouch becomes the coelom and the pouch moves down to meet mid ventrally and the wall forms
the mesoderm. Similar coelom development in protochordates and the echinoderms explains the
origin of this group from echinodermata. The flatworms lack body cavity and have solid body
construction so they are called acoelomates. Rotifers and roundworms do not have true coelomic
cavity as it is not lined by mesoderm. It is called pseudocoel. It is formed form the persistent
blastocoel. Since it is not a true cavity so the viscera is not held by peritoneal cover but lies free
`in the cavity or space.

Figure 1.41: General structure of triploblastic organisms; Animal Body Cavities. There are three major
types of body cavities among the animals. (a) Acoelomates do not have enclosed body cavities. Animals
that lack an enclosed body cavity are called acoelomates. In these animals, the space between the gut and
the body wall is filled with masses of cells called mesenchyme (Figure 1.41a). (b) Pseudocoelomates
have only one layer of muscle, and it lies outside the body cavity. Pseudocoelomate animals have a body
cavity called a pseudocoel, a liquid-filled space in which many of the internal organs are suspended. Their
control over body shape is crude because the pseudocoel has muscles only on its outside; there is no inner
layer of muscle surrounding the organs (Figure 1.41b). (c) Coelomates have a peritoneum surrounding
the internal organs. The body cavities of some coelomates, such as this earthworm, are segmented.
Coelomate animals have a coelom, a body cavity that develops within the mesoderm. It is lined with a
special structure called the peritoneum and is enclosed on both the inside and the outside by muscles
(Figure 1.41 c).

The most significant aspect of differentiation in early embryogenesis is the creation of three
primary germ layers.
The ectoderm, the outer covering of the embryo, gives rise to the outer layer of skin, the hair
and nails, and the secretory cells of the sweat glands.
From the nerve tube induced by the notochord, the ectoderm also produces the entire nervous
system-brain, spinal cord, and peripheral nerves, as well as the specialized end receptors of the
sense organs.
Some of the lining of the mouth and anus, as well as the enamel of the teeth, also derives from
the ectoderm.
The endoderm provides the linings of the digestive tract and the major respiratory ducts, most
of the liver and pancreatic cells, the lining of the urinary bladder and the inner layer of the
urethra, and the thyroid and parathyroid glands.
The mesoderm is the third of the germ layers to be formed, but it is the source of the greatest
bulk of living material within the organism. All muscles, solid connective tissues (bone,
cartilage, and fiber), blood and its vessels, and thin mesenteries that connect most visceral organs
to the body wall are derived from the mesoderm.
The dermis, the primary functional stratum of the skin, also arises from the mesoderm, as do the
kidneys and the reproductive organs. The segmentation that characterizes many multicellular
groups is a result of the division of mesodermal tissue into a sequence of repeating blocks early
in development.
Later structures, such as the vertebrae and major muscle systems of the body, are arranged into
somites, or regular repeating units running from front to back, because of this prior segmentation
of their precursor mesodermal structures.
Figure 1. 42: General significance of germ layers in vertebrates (embryoblast tree).
4. 3. Embryo and fetal development in human
Human development can be divided into embryonic development (months1 and 2) and fetal development
(months 3-9). Organ development begins with neural tube and heart formation. There follows a steady
progression of organ formation during embryonic development. During the fetal development, refinement
of features occurs, and the fetus adds weight. Fetal development includes the third through ninth months
of development. At that time the embryo looks human. The period of time between conception and birth
during which the fetus grows and develops inside the mother's womb is called gestation. In humans, the
length of pregnancy or gestational age is the time measured from the first day of the woman's last
menstrual cycle to the current date. It is measured in weeks.

The following list describes specific changes that occur in the womb during human embryo and fetal
development:

 Week 5 of pregnancy (gestational age)

o The brain, spinal cord, and heart begin to develop.
o The gastrointestinal tract begins to develop.
 Weeks 6 - 7 of pregnancy (gestational age)
o Arm and leg buds become visible.
o The brain develops into five areas and some cranial nerves are visible.
o The eyes and ear structures begin to form.
o Tissue forms that develops into the vertebra and some other bones.
o The heart continues to develop and now beats at a regular rhythm.
o Rudimentary blood moves through the main vessels.
 Week 8 of pregnancy (gestational age)
o The arms and legs have grown longer, and foot and hand areas can be distinguished.
o The hands and feet have fingers and toes (digits), but may still be webbed.
o The brain continues to form.
o The lungs begin to form.
 Week 9 of pregnancy (gestational age)
o Nipples and hair follicles form.
o Elbows and toes are visible.
o All essential organs have begun to form.
 Week 10 of pregnancy (gestational age)
o The eyelids are more developed.
o External features of the ear begin to take their final shape.
o Facial features continue to develop.
o The intestines rotate.

The end of the 10th week of pregnancy marks the end of the "embryonic period" and the beginning of the "fetal period."

 Weeks 11 to 14 of pregnancy (gestational age)

o Eyelids close and will not reopen until about the 28th week.
o The face is well formed.
o Limbs are long and thin.
o Genitals appear well differentiated.
o Red blood cells are produced in the liver.
 o The head makes up nearly half of the baby's size.
o The baby can make a fist with its fingers.
o Tooth buds appear for the baby teeth.
 Weeks 15 to 18 of pregnancy (gestational age)
o The skin is almost transparent.
o Fine hair called lanugo develops on the head.
o Meconium is made in the intestinal tract.
o More muscle tissue and bones have developed, and the bones become harder.
o The baby begins to make active movements.
o The liver and pancreas produce fluid secretions.
o Sucking motions are made with the mouth.
 Weeks 19 to 21 of pregnancy (gestational age)
o The baby can hear.
o The baby makes more movements.
o The mother may feel a fluttering in the lower abdomen.
 Week 22 of pregnancy (gestational age)
o Lanugo hair covers entire body.
o Eyebrows and lashes appear.
o Nails appear on the fingers and toes.
o The baby is more active with increased muscle development.
o The mother can feel the baby moving.
o The fetal heartbeat can be heard with a stethoscope.
 Weeks 23 to 25 of pregnancy (gestational age)
o Bone marrow begins to make blood cells.
o The lower airways of the baby's lungs develop but still do not produce surfactant (a substance that allows the
alveoli to open for gas exchange).
o The baby begins to store fat.
 Week 26 of pregnancy (gestational age)
o Eyebrows and eyelashes are well formed.
o All eye parts are developed.
o The baby has a hand and startle reflex.
o Footprints and fingerprints are forming.
o Air sacs form in lungs.
 Weeks 27 to 30 of pregnancy (gestational age)
o Rapid brain development occurs.
o The nervous system is developed enough to control some body functions.
o The eyelids open and close.
o The respiratory system, while immature, has developed to the point where gas exchange is possible.
 Weeks 31 to 34 of pregnancy (gestational age)
o A rapid increase in the amount of body fat occurs.
o Rhythmic breathing movements occur, but the lungs are not fully mature.
o The bones are fully developed, but still soft and pliable.
o The baby's body begins storing iron, calcium, and phosphorus.
 Week 38 of pregnancy (gestational age)
o Lanugo begins to disappear.
o Body fat increases.
o Fingernails reach the end of the fingertips.
 Weeks 39 to 42 of pregnancy (gestational age)
o Lanugo is gone except for on the upper arms and shoulders.
o Fingernails extend beyond fingertips.
o Small breast buds are present on both sexes.
o Head hair is now coarse and thicker.

Source: [Link]
III. Post-Embryonic development
In general, upon transition to the postembryonic state the organism either immediately possesses the
principal pubertal morphological characteristics (direct development) or essentially differs from the
pubertal form, in which case the larva that hatches out of the egg must undergo a metamorphosis before it
reaches its adult state(indirect development). Growth continues during the period of postembryonic
development and further organogenesis and histogenesis occur. The functions of the developing organism
become more complex; establishment of the final proportions of the body is especially characteristic.

1. Metamorphosis
1.1. Metamorphosis in Amphibians
In frogs (and other anurans), the fertilized egg is a single cell that rapidly divides again and
again, producing new cells that quickly differentiate into the organs of the frog embryo. Within a
few days after fertilization, depending on water temperature, the eggs hatch into tadpoles. A
newly hatched tadpole lives off yolk stored in in its body. It gradually grows larger and develops
three pairs of gills that allow it to breathe efficiently underwater. Eventually, the tadpole’s mouth
opens, allowing it to feed. The tadpole continues to swim, eat and grow for several weeks before
it matures to the next stage. The first sign of further development is the appearance of hind legs.
During this process of changes called metamorphosis, the front legs develop and the tail becomes
shorter as it is resorbed. The mouth broadens, developing teeth and jaws. Internally, the tadpole's
gills are replaced with lungs until finally the tadpole has become a frog. The young frog grows and
matures to adulthood over a period of 2-4 years.

Figure 1.43: Developmental history of

Rana pipiens. This figure also shows the
presumptive zones of the germline cells in
the developing embryo (dark spot) and the
location of the gonads.
 Figure 1.44: In and Out of the
Water Most stages in the life
cycle of temperatezone frogs
take place in [Link] aquatic
tadpole is transformed into a
terrestrial adult through
metamorphosis.

The adult frogs then lay their eggs and begin the cycle again. Metamorphosis in anurans mimics
mammalian postembryonic development. This process involves distinct change in different
organs and tissues. The larval specific organs, such as the tail and gills, are totally resorbed
during metamorphosis, while the adult specific ones, such as the limbs, develop de novo. Most of
the organs/tissues are present in both tadpoles and frog but are drastically remodeled during
metamorphosis. Interestingly all such diverse changes during amphibian metamorphosis are
totally dependent on a hormone called thyroxine which is produced by the thyroid gland and
circulates throughout the blood stream to stimulate metamorphosis.

1.2. Metamorphosis in other groups

Metamorphosis is also observed in other animal groups. Insects may undergo gradual
metamorphosis, where transformation is subtle, or complete metamorphosis, where each stage of
the life cycle appears quite different from the others. Although all insects molt and change body
shape through their life, some of them undergo only minor changes while most undergo extreme
changes in structure and function. In some insects, there may be no true metamorphosis at all.
With regard to metamorphosis, entomologists divide insects into three groups: ametabolous,
hemimetabolous, and holometabolous (see BIO 303)
 A B

Figure 1-45: Metamorphosis in insects. A: hemimetabolous insect (migratory locust); B: holometabolous insect
(fly)

Note: Metamorphosis can also be observed in other invertebrate like in trematoda (flatworms).

2. Sexual maturity in vertebrates (Mammalia case)

After birth (or hatching) animals continue growth and development. One of the most important
developmental stages is characterized by changes that allow the animal to perpetuate the species.
Sexual maturity is the age or stage when an organism can reproduce. It is sometimes considered
synonymous with adulthood, though the two are distinct. In humans, the process of sexually
maturing is termed puberty.

Most multicellular organisms are unable to sexually reproduce at birth (or germination), and
depending on the species, it may be days, weeks, or years until their bodies are able to do so.
Also, certain cues may cause the organism to become sexually mature. They may be external,
such as drought, or internal, such as percentage of body fat (such internal cues are not to be
confused with hormones which directly produce sexual maturity). Sexual maturity is brought
about by a maturing of the reproductive organs and the production of gametes. It may also be
accompanied by a growth spurt or other physical changes which distinguish the immature
organism from its adult form. These are termed secondary sex characteristics, and often represent
an increase in sexual dimorphism. For example, before puberty, human children have flat chests,
but adult females have breasts while adult males generally do not. However, there are exceptions
such as obesity and hormone imbalances such as gynecomastia.
After sexual maturity is achieved, it is possible for some organisms to become infertile, or even
to change their sex. Some organisms are hermaphrodites and may or may not be able to produce
viable offspring. Also, while in many organisms sexual maturity is strongly linked to age, many
other factors are involved, and it is possible for some to display most or all of the characteristics
of the adult form without being sexually mature. Conversely, it is also possible for the
"immature" form to reproduce (see progenesis).

3. Gametogenesis
Gametogenesis is the production of gametes, either eggs by the female or sperm by the male,
through a process involving meiosis. In animals, the cells which will ultimately differentiate into
eggs and sperm arise from primordial germ cells set aside from the potential somatic cells very
early in the formation of the embryo. The final products of gametogenesis are the large,
sedentary egg cells, and the smaller, motile sperm cells. Each type of gamete is haploid; that is, it
contains half the chromosomal complement and thus half as much deoxyribonucleic acid (DNA)
as the somatic cells, which are diploid. Reduction of the DNA content is accomplished by
meiosis, which is characterized by one cycle of DNA replication followed by two cycles of cell
division.

In higher animals the ovum differs from the sperm in that it is larger and is non-motile, a smooth
sphere or oval lacking the flagellum of the sperm. Like that of the sperm, its nucleus contains the
chromosomes, which bear the hereditary material of the parent. A gamete, ovum or sperm,
contains half the number of chromosomes found in the body cells of the parent, i.e., the gamete
is haploid. In animals, ova contain stored food called the yolk, the amount of which varies in
different species, depending on the length of time required for the embryo to become self-
sufficient in obtaining nourishment outside the egg. In this ovum all the yolk from the original
cell is collected; the three other, yolkless, cells are called polar bodies and never develop further.

3.1. Spermatogenesis
Spermatogenesis appears to be a fairly conserved process throughout the vertebrate series. Thus,
spermatogonia develop into spermatocytes that undergo meiosis to produce spermatids which
enter spermiogenesis where they undergo a morphological transformation into spermatozoa.
There is, however, variation amongst the vertebrates in how germ cell development and
maturation is accomplished. This difference can be broadly divided into two distinct patterns,
one present in anamniotes (fish, amphibia) and the other in amniotes (reptiles, birds, mammals).
For anamniotes, spermatogenesis occurs in spermatocysts (cysts) which for most species develop
within seminiferous lobules. Cysts are produced when a Sertoli cell becomes associated with a
primary spermatocyte. Mitotic divisions of the primary spermatocytes produce a cohort of
secondary spermatocytes that are enclosed by the Sertoli cell which forms the wall of the cyst.
With spermatogenic progression a clone of isogeneic spermatozoa is produced which are
released, by rupture of the cyst, into the lumen of the seminiferous lobule. Following
spermiation, the Sertoli cell degenerates. For anamniotes, therefore, there is no permanent
germinal epithelium since spermatocysts have to be replaced during successive breeding seasons.
The morphology of sperm is highly specialized, with distinctive organelles forming both the
posterior motile apparatus and the anterior acrosome, which assists in penetration of the oocyte
at fertilization.

3.2. Oogenesis

Formation of the ovum most often involves substantial increases in cell volume as well as the
acquisition of organellar structures that adapt the egg for reception of the sperm nucleus, and
support of the early embryo. Among lower vertebrates and invertebrates, mitotic divisions of the
precursor cells, the oogonia, continue throughout the reproductive life of the adult; thus
extremely large numbers of ova are produced. In the fetal ovary of mammals, the oogonia
undergo mitotic divisions until the birth of the fetus, but a process involving the destruction of
the majority of the developing ova by the seventh month of gestation reduces the number of
oocytes from millions to a few hundred. Around the time of birth, the mitotic divisions cease
altogether, and the infant female ovary contains its full complement of potential ova. At puberty,
the pituitary hormones, follicle stimulating hormone (FSH), and luteinizing hormone (LH)
stimulate the growth and differentiation of the ova and surrounding cells. One important feature
of oocyte differentiation is the reduction of the chromosome complement from the diploid state
of the somatic cells to the haploid state of gametes. Unlike the formation of sperm, in which the
two divisions of meiosis produce four equivalent daughter cells, the cytoplasm of the oocyte is
divided unequally, so that three polar bodies with reduced cytoplasm and one oocyte are the final
products. In this ovum all the yolk from the original cell is collected; the three other, yolkless
polar bodies never develop further. Generally, each fertilized oocyte produces a single embryo,
but there are exceptions. Identical twins, for example, arise from the same fertilized egg.

Figure 1-46: Gametogenesis in male and female

humans. Spermatogenesis (A): two divisions of
meiosis produce four equivalent daughter cells;
Oogenesis (B): three polar bodies with reduced
cytoplasm and one oocyte are the final products.
Gametogenesis in humans begins with 46 pair of
chromosomes, splits off to 23 then combines for a
unique new 46 pair of chromosomes (C).
The provision of nutrients for the embryo is a major function of the egg, and this is accomplished
by the storage of yolk in the cytoplasm. Yolk consists of complex mixtures of proteins (vitellins),
lipids, and carbohydrates in platelets, which are membrane-surrounded packets dispersed
throughout the egg cytoplasm (ooplasm). The amount of yolk in an egg correlates with the
nutritional needs of the embryo. Although the eggs of mammals are extremely small as
compared to the fetus, the bulk of the nutrition is supplied by the placenta; yolk is required only
until implantation in the uterine wall.