Module 7

Organogenesis
After the three germ layers are established at the gastrula stage, it leads to
organogenesis, the formation of organs. It is at this stage that process of pattern
formation continues as specific structures of the organism are formed from the
three germ layers.

The cells that remain on the outer surface of the gastrula is the ectoderm that
will form the epidermis of the skin, nervous system, sense organs and a few other
cell types. The endoderm gives rise to tissues that line the digestive tract and
organs that develop as outgrowths of the digestive tract (including the liver,
pancreas, and lungs). The remaining intermediate layer is the mesoderm that will
give rise to the skeletal, muscle tissues and circulatory, excretory and reproductive
systems.

Learning Objectives: At the end of this lesson, the student should

be able to:

1. Explain how the different organs develop by induction through chemical

signals from neighboring cells
2. Define organogenesis and summarize the fate of each of the germ layers.

Induction

We have learned that even as early as in the fertilization stage, the anteroposterior
and dorsoventral axes are already specified and established. By by the blastula
stage of the amphibian embryo, the fates of the cells in different regions of the
embryo are already specified. Then in the gastrula stage, the cells of the blastula
undergo different kinds of movements to organize and establish the three germ
layers from where the different organs of the animals will be derived from.

Genetic programs are also crucial in the formation of organs. Even before the egg
is fertilized, genes that will encode for specific proteins essential for development
are already in existence. But with the fertilization of the egg by the sperm, a new
set of genetic programs (e.g. gene expression) for new set of proteins of
development come into view. These proteins released by group of cells act as
chemical signals to adjacent cells or tissues to to set in motion a different pathway
of differentiation, leading to the morphogenesis of tissues or organs.

The response of these cells to the chemical signals released by adjacent cells is
called embryonic induction. It is the process by which the identity of certain
cells influences the developmental fate of surrounding cells
([Link]
[Link]?_ga=2.66937582.158688425.1608142260-1842665231.1608142260)

Prepared by Margaret L.C. de Guzman. 2020 Department of Biology. College of

Arts and Sciences. University of the Philippines Manila.
Two scientists are recognized for this phenomenon of induction, and they are the
German embryologist Hans Spemann and his student Hilde Mangold. They did
several experiments by transplanting the cells of the dorsal lip of the blastopore
(DLB) of salamanders to different sites of the recipient embryos. Based on the
results of their experiments, the transplanted DLB results to the formation of a
secondary axis, then to a second central nervous system (CNS), muscles and other
axial organs in the recipient. What does this tell us? The DLB contains chemical
signals that induces the neighboring cells to develop as what they are
programmed to become.

The classic experiment of Spemann and Mangold is shown below.

[Link]
Spemann/d0315ad858eaae42a8125d33cc555103e2d70561/figure/3

The transplantation experiment of Spemann and Mangold have shown that the
cells in the DLB have an extraordinary role in the formation of the dorsal
mesoderm, particularly the notochord and some pharyngeal endoderm.

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
The cells of the DLB have been referred to as the Spemann organizer since the
cells here induces the formation of the CNS. One important chemical that was
later known to be the cause of the induction of the CNS is chordin (encoded by
the gene chordin). Before the CNS is induced to form, chordin is a protein that
dorsalises early vertebrate embryonic tissues binds to bone morphogenetic
proteins (BMPs) that may be involved also in organogenesis ( Segen’s Medical
Dictionary, 2012). If you want to know more about embryonic induction, you can
go to this link : [Link] In other embryonic
vertebrates with no DLB, the organizer will be embryonic shield (fishes) and the
Hensen’s node (amniotes).

Neurulation as the first induction event in animal development

The organs first formed in the early vertebrate embryos are the notochord, brain
and spinal cord. The notochord is first formed from the mesodermal cells of the
DLB or the mesodermal cells of the Hensen’s node of amniotes called the
chordamesoderm.

But before we proceed further to induction, let us first understand more about
the Hensen’s node in amniotes, including humans. As mentioned in the previous
module, a blastopore is not formed in amniote gastrulation because of the
enormous amount of yolk. Instead a primitive streak is formed by the movement
of the cells and later on a primitive groove is formed as the functional equivalent
of the blastopore.

Further movement (by ingression) of the cells from the surface epiblast results to
a thickened structure at the anterior end of the primitive streak called the
Hensen’s node. The aggregate of cells in this thickened knot are destined to form
the notochord. Notochord, if you recall, is made of cartilage-like cells that serves
as a flexible axial support in all chordate embryos. It grows forward along the
length of the embryoas a cylindrical rod of cells. At this point, we can generalize
that where the DLB in amphibians or the Hensen’s node in amniotes, establishes
the posterior end of the embryo since the notochord develops anterior to these
structures.

How does the notochord play a role in the formation of the brain and spinal cord?
Several experiments have been done by transplanting portions of the notochord
mesoderm to other areas of the embryo. These experiments have established
where first induction event happens in chordate embryos. The notochord releases
chemical signals to the ectoderm above or dorsal to it. Note that the mesodermal
cells are not in close contact with the overlying ectoderm. This tells us that the
chemical signals, generally called as morphogens, diffuse to these neighboring
cells and induce their differentiation.

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
Results of the experiments have shown that the notochord induces/causes the
overlying ectoderm to thicken and differentiate to form the precursor of the central
nervous system, and that is the neural plate. Later on, the cells of the neural
plate undergo conformational changes that makes the the central cells of the
neural plate to move downward forming a depression called the neural groove
(NG); and the structures flanking this groove are called the neural folds (NF).
Further changes in the shape of the cells of the neural folds cause them to move
closer until the two folds meet and fuse to form the neural tube.

Click this link for a 2 minute video on neurulation: [Link]

AoA

Therefore, at the end of the neurulation event, the ectoderm that did not
differentiate as neural tube will become the outer layer of the skin, the epidermis.
Further growth and differentiation of the neural tube cause the anterior end to
become the brain and the rest of the tube develops into the spinal cord. Recall
that the neural tube encloses the former neural groove and as a result the neural
tube is said to be hollow until the adult stage; the reminder of this embryonic
beginnings as hollow strctures can be seen as the ventricles of the adult brain
and the central canal of the spinal cord.

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
Go back to the figure above on neurulation. In the section of the neural plate,
notice the small area colored green, that later increase in size at the neural tube
stage. This green area indicates the location of cells that what we call the neural
crest. So, these cells are also ectodermal cells that differentiate not as epidermis
of the skin nor the neural tube. The neural crest cells migrate downward from
their original position and form the the dorsal root ganglia of the spinal nerves
and the postganglionic sympathetic neurons.

The figure below shows the migration of the neural crest cells from the anterior
region of the body to the posterior end and what are their fates in these regions.
The link given is where this figure was copied from and can also be read by you
for mor more information and understanding about the neural crest cells.
[Link]
0

Formation of other organs

When the nervous system is developing thru neurulation, other organs also take
shape (by morphogenesis, right?). Shown below is a cross-section of the frog
neurula, the embryonic stage when the neural tube is forming and has formed

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
already. We can see that blocks of anterior mesoderm (or epimere) called
somites form on either side of the neural tube. Further differentiation of these
somites will give rise to vertebrae, muscles, and other parts of the body axis. The
cells of middle mesoderm (intermediate mesoderm or mesomere) gives rise to
the prospective kidneys and reproductive structures.

[Link]
development/[Link]

The ventrolateral mesoderm (lateral plate mesoderm or hypomere) splits into

two layers : an outer layer that becomes associated with the ectoderm the somatic
mesoderm layer and an inner layer that becomes associated with the

endoderm, the splanchnic

mesoderm. The splanchnic
mesoderm covers the endoderm
and eventually the gut that
develops from it. There is a
space that is formed between
these two layers, it is called the
coelom.

Other organs that develop very

early too in development are the
heart and blood vessels and
function already while still
developing.

[Link]

Another example of organogenesis is seen in the development of the four-

chambered heart of birds and mammals. Lifting the figure from Solomon, Berg &
Martin is the formation of heart in birds and mammals. We can see that heart
forms from the fusion of two blood vessels (a) and that at first the end that receives

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
blood from veins is at the bottom. The developing heart twists or loops in the very
small space called the pericardial cavity, bringing the atrium to position above
the ventricle, and the chambers then divide by septation to form the four-
chambered heart.

If you are interested to know what can go wrong in the the development of the
cardiovascular system, you can click this link
[Link] Any
mistake that happens in the development of organs while the mammal is still in
the mother’s womb is called congenital deformities or diseases.

The endoderm layer as we have said gives rise to organs of the digestive and
respiratory systems. The digestive tract is formed as a separate foregut and
hindgut as the body wall grows and folds. As the embryo grows, these tubes,
which are lined by endoderm, grow and become greatly elongated.

Lifted from the site [Link] the

formation of the primitive gut tube is as follows:

(a) The gut tube is formed from endoderm lining the yolk sac which is
enveloped by the developing coelom as the result of cranial and caudal
folding.
(b) During folding, somatic mesoderm is applied to the body wall to give rise to
the parietal peritoneum.
(c) Visceral (or splanchnic) mesoderm is wraps around the gut tube to form the
mesenteries that suspend the gut tube within the body cavity.
(d) The mesoderm immediately associated with the endodermal tube also
contributes to most of the wall of the gut tube.
(e) Nerves and neurons found in the wall are derived from neural crest.
(f) Summary of germ layer contributions:

• endoderm: mucosal epithelium, mucosal glands, and submucosal

glands of the GI tract.

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
 • mesoderm: lamina propria, muscularis mucosae, submucosal
connective tissue and blood vessels, muscularis externa, and
adventitia/serosa
• neural crest: neurons and nerves of the submucosal and myenteric
plexus

Basic subdivisions of the gut tube

Cranio-caudal and lateral folding cause the opening of the gut tube to the yolk
sac to draw closed (like a purse string) forming a pocket toward the head end of
the embryo called the "anterior (or cranial) intestinal portal" and a "posterior (or
caudal) intestinal portal" toward the tail of the embryo. These are the future
foregut and hindgut, respectively. The midgut remains open to the yolk sac.

Further folding and growth of the embryo causes the communication of the gut
with the yolk sac to continue to get smaller and the regions of the gut (foregut,
midgut, and hindgut) to become further refined:

The derivatives of the gut regions are as follows:

FOREGUT Trachea and respiratory tract, lungs, esophagus, stomach,

liver, gallbladder and bile ducts, pancreas and upper
duodenum
MIDGUT Lower duodenum, jejunum, ileum, cecum, appendix,
ascending colon, proximal 2/3 of descending colon
HINDGUT Distal 1/3 of the transverse colon, descending and sigmoid
colon, rectum, upper anal canal, urogenital sinus

The respiratory system develops from the ventral evagination of the foregut called
the lung buds. The trachea grows downward from it and bifurcates into two lung

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.
buds, which develop into lungs. The anterior part of the foregut becomes the
pharynx. A series of small outpocketings of the pharynx, the pharyngeal pouches
bud out laterally. These pouches meet the branchial grooves, a corresponding
set of inpocketings from the overlying ectoderm. The arches of tissue formed
between the grooves are called branchial arches. They contain the rudimentary
skeletal, neural, and vascular elements of the face, jaws and neck.

These are just some of the overview of organogenesis in vertebrate embryos. And
do you need to memorize these derivatives from the three germ layers….YES..
kalimutan niyo na si Ma’am Meggie, wag lang ang mga germ layers and the organs
derived from each 😊. Aside from these derivatives, Wilt & Hake (2004) said you
should memorize also the characteristic arrangement of these germ layers since
the basic body plan of all vertebrates have these as the foundation of the
anatomical arrangements in the subphylum of vertebrates. Any deviation from
this basic plan results to anomalies, malformations, abnormalities of the
organs and this results to malfunctioning body parts.

There are still more to know about the development of animals, but for now,
knowing and understanding the basic concepts and principles involved in
transforming a fertilized egg into a new being or individual is commendable.

Have a wonderful semester/Christmas Break.

Let’s be grateful to everything we can do and have despite the pandemic.

Thank you…Ma’am Meggie 😊

BOOK REFERENCES:

1. Carlson, B.M. 1994. Human Embryology and Developmental Biology. Mosby.

2. Carlson, B.M. 1996. Patten’s Foundation of Embryology. 6th ed. McGraw Hill
Book Company.
3. Solomon EP, Berg LR and Martin DW. 2008. Biology. 8 th ed. Thomson
Brooks/Cole.
4. Wilt, F & S. Hake. 2004. Principles of Developmental Biology. W.W. Norton &
Company.

Prepared by Margaret L.C. de Guzman. 2020. Department of Biology. College of Arts and
Sciences, University of the Philippines-Manila.