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19.1 Face and upper foregut

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Introduction
Development of the facial region
Oral cavity
Tongue
Development of the teeth
Development of the salivary glands
Definitive pharynx: Derivatives of the first and second pharyngeal arches
o Thyroid
o Middle ear
o Tonsils
Derivatives of the third and fourth pharyngeal arches
o Thymus
o Parathyroid glands and ultimopharyngeal bodies

Lungs

Introduction Quiz
Quiz 02

Many facial components such as the ears, parts of the nose as well as the jaw, the teeth and
salivary glands arise from the pharyngeal portions of the foregut, among others. That is why an
understanding of the development of this region is important.
The embryonic foregut (stage 8, ca. 23 days, 8 ) dead-ends in the cranial region below the
very rapidly growing brain. Together with the ectoderm, which covers the embryo, it forms the
oropharyngeal membrane that, however, is soon torn (stage 11, ca. 29 days, 11 ). A
connection between the foregut and the amniotic cavity has arisen and it is now termed the
stomodeum (see Fig. 3). During the flexions of the embryo in the following days (the strongest
occurring in stage 14, ca. 32 days, 14 ) accumulations of mesenchyma occur in the
region of the foregut on both sides, which then become the pharyngeal arches. They are
separated, one from the other, by the pharyngeal folds that, on the inside, border on the
foregut endoderm (pharyngeal pouches).

Fig. 4 - View and section through an embryo in stage 13 (ca. 32 days) Legend
 Fig. 4
Not all pharyngeal
arches are the same
size. The first two are
very prominent, whereby
the second, as an
operculum, even grows
over the smaller third
and fourth pharyngeal
arches lying just below it
(see white arrows).

1 First pharyngeal arch (mandibular arch)

2 Second pharyngeal arch (hyoid arch)
3 Third pharyngeal arch
4 Fourth pharyngeal arch
5 Pharyngeal pouches
6 Pharyngeal folds

The mesenchyma in the pharyngeal arch region arises mainly from emigrated neural crest
cells. It is also termed the mesectoderm. In addition one also finds mesenchyma from
paraxial mesoderm, which fills the remaining space between ectoderm and endoderm. This
entire portion is called the embryonic pharynx.

Fig. 5 - Overview: the structures that are involved in forming the pharynx Legend
 Fig. 5
Lateral view of an
embryo: Organization of
the head and pharynx
with the various types of
tissue depicted
separately. The various
portions of tissue of the
individual segments are
connected with each
other by vertical
columns.

More info
The morphologic segmentation in this region is based on the differing expression of certain
homebox- and other genes, their products being released from the neural tube
(mesencephalon [midbrain] and rhombencephalon [hindbrain]), whereby finally interaction with
the ectoderm also plays a decisive role.
A large aortic arch, which arises directly from Fig. 6 - The metameric arrangement Legend
the aortic root and discharges into the dorsal of the structures in the neck region
aorta, passes through each pharyngeal Fig. 6
arch, together with a cerebral nerve and Side view of an embryo:
precartilagineous mesenchyma. From the The pharyngeal arches
precartilagineous mesenchyma arise the with their associated
typical skeletal structures of the neck as found cerebral nerves (CN)
in adults. The knowledge of this basic
structure of the pharyngeal arches is
important since these are retained, even
though it appears that this metameric
arrangement is displaced by differing rates of
growth as development proceeds.

1 Trigeminal nerve
2 Facial nerve
3 Glossopharyngeal nerve
4 Vagus nerve
I First pharyngeal arch
II Second pharyngeal arch
III Third pharyngeal arch
IV Fourth pharyngeal arch

- Muscular portion of the pharyngeal arches

- Neural portion of the pharyngeal arches
- Skeletal portion of the pharyngeal arches

In the following diagram all the derivatives of the pharyngeal pouches, folds and arches are
shown.

Fig. 7 - Derivatives of the pharyngeal region

Development of the facial region

 Quiz 04

The structures for the face

stem from various primordia
around the stomodeum that
have arisen up to the tearing
of the oropharyngeal
membrane (stage 11, ca. 29
days, 11 ):
a medial sacculation in
front of the
oropharyngeal
membrane as the
primordium of the
adenohypophysis.
the medial
frontonasal process
(processus
frontonasalis) that soon
forms a medial and a
lateral nasal process
on both sides and
pushes the
adenohypophysis into
the stomodeum.
the bilateral
mandibular arch (first
pharyngeal arch) with a
maxillary and a
mandibular process.

In order to be able to study the individual facial structures in Video

detail, please use the interactive diagram (726 kB) with a Development of the
profile view of facial development. facial region, front
 view (1,1 Mb);
profile view (1.2
Mb)

Oral cavity Quiz

Quiz 26

Following the tearing of the oropharyngeal membrane in the

young embryo, the ectoderm-covered stomodeum, which gets
surrounded by the mandibular prominences, goes directly
over into the endoderm-covered foregut. It is also called a
primary oral cavity because at this time no nasal cavity yet
exists. (For the forming of the exterior nose, see facial
development).
Only through the formation of the nasal sacs are the primary
palates (= floors of the two nasal sacs) formed that divide the
primary oral cavity into oral and nasal cavities. The wall
between the two nasal sacs is called the nasal septum.
Using the two interactive diagrams the relationships can be
studied in detail.
View from the oral cavity towards its roof (1.9 Mb)
View from the oral cavity towards its roof shown as if it
were transparent
(1.7 Mb)
The adenohypophysial primordium (AH) appears in stage 10
(28 days) in front of the oropharyngeal membrane as
thickening of the ectoderm. It develops in the immediate vicinity
of the basal plate of the neural tube (prosencephalon
[forebrain]).
At the same time a sacculation arises from this part of the basal
plate, the neurohypophysis (NH). This basal plate belongs to
the future diencephalon (part of the prosencephalon). The AH
primordium forms a pouch (Rathke's pouch) in stage 13 (32
days), which remains in constant contact with the NH of the
diencephalon and later surrounds this in the front. (1)
 Fig. 8 - Rathke's pouch Legend
Fig. 8
The
adenohypophysis
forms from a small
pouch coated with
ectoderm (Rathke's
pouch) that forms in
front of the
oropharyngeal
membrane and with
the increasing
flexion is displaced
into the pharyngeal
roof.

1 Primordium of the adenohypophysis

2 Primordium of the neurohypophysis

Subsequently, the AH loses Fig. 9 - Rathke's pouch in Legend

the contact to the ectoderm
and, through further embryonic
flexion and the expansion of
the prosencephalon (=
future di- and
telencephalon) gets
displaced at the cerebral base
together with the NH.
With the facial formation
through the spreading
maxillar and mandibular
arches the AH orifice is
displaced into the pharynx
roof. The skull base (os
sphenoideum) forms between
the pharynx roof and the AH in
the later embryonic period.
stage 17
(ca. 41 days)
1 Right posterior nasal orifice
2 Orifice of the Rathke's pouch
3 Maxillary process
Fig. 9
The orifice of the
adenohypophysis
gets displaced into
the pharynx roof by
the growing facial
processes. In stage
17 (ca. 41 days) the
connection is still
open, but is
obliterated a short
time later.
More info

In the adenohypophysis mesenchymal trabeculae arise and

divide it into various lobules. The pars tuberalis surrounds the
NH stalk, whereas a pars intermedia is not identifiable in
humans. At the end of the embryonic period the first hormone
activities in the blood can be measured.

Tongue Quiz
Quiz 05

The tongue begins to form at approximately the same time as Quiz

the palates. It extends from various protuberances on the Quiz 28
pharynx floor. Already at the time of the medial fusion of the
first (mandibular) and second (hyoid) pharyngeal arches a
medial protuberance, the tuberculum impar, appears on the
lower edge of the mandibular arch. To the left and right of it two
further swellings form, the lateral lingual prominences.
These three swellings extend
from the mandibular arch and
later form the anterior 2/3 of
the tongue. Thus this part the
tongue consists of both
ectodermic and endodermic
portions. At the caudal end of
the tuberculum impar arises
the primordium (or anlage) of
the thyroid (stage 10, ca. 28
days 10 ) as an unpaired
ventral endodermic bud. In its
further development it
migrates in front of the larynx
and, at the invagination site at
the tongue base, leaves
behind the foramen cecum.
The posterior third is formed
from the material of the fusion
site of the 2nd and 3rd
pharyngeal arches, the
copula, and a small portion of
the 4th pharyngeal arch. It
only consists of endodermic
parts. The third pharyngeal
arch probably grows over the
second one so that in the end
the second one has no
mesenchymal contribution
to tongue formation. Between
the anterior two-thirds and the
posterior third the terminal
sulcus is found. The tongue is
delimited caudally by the
hypopharyngeal eminence
that marks the entrance into
the trachea (laryngeal inlet).
Lingual musculature stems Fig. 10 - Embryo in stage 14 Legend
from muscle cells that ca. 33 days
immigrate from the occipital Fig. 10
The occipital
somites into the tongue. The somites are shown
emigrating predecessor cells in green. The
follow the hypoglossal nerve tongue receives
(CN 12). material from all 4
occipital somites for
building up the
musculature. The
connective tissue
and the myoblasts
together form a
hypoglossal cord.
1 Hypoglossal tongue cord

Development of the tongue Fig. 11 - Innervation of the Legend

involves several pharyngeal tongue
arches. This is also evidenced Fig. 11
Cerebral nerves of
in their innervation that at the the pharyngeal
first moment seems relatively arches grow into the
complex. tongue and provide
it with motor,
sensitive, and
sensorial
innervation.

1 Maxillary nerve (CN V2)

2 Lingual nerve Quiz
(mandibular nerve, CN V3)
3 Chorda tympani Quiz 06
(facial nerve, CN VII)
4 Glossopharyngeal nerve (CN IX) Quiz
5 Vagus nerve (CN X)
6 Hypoglossal nerve (CN XII) Quiz 30
7 Tongue
Sensitivity: (touch)
Anterior two-thirds of the 1. Pharyngeal arch N. lingualis (CN V3)
tongue
Posterior third of the 3. Pharyngeal arch N. glossopharyngeus (CN
tongue IX)
Tongue base 4. Pharyngeal arch N. vagus (CN X)
Sensorial: (taste)
Anterior two-thirds of the 2. Pharyngeal arch Chorda tympani (CN VII)
tongue
Posterior third of the 3. Pharyngeal arch N. glossopharyngeus (CN
tongue IX)
4. Pharyngeal arch N. vagus (CN X)
Motor: (movement)
Entire musculature 6. Pharyngeal arch N. hypoglossus (CN XII)
Development of the teeth Quiz
Quiz 07

One subdivides the human denture into four jaw quadrants. In Quiz
each one 5 deciduous (or milk) teeth and later 8 permanent Quiz 08
teeth form. Quadrants 1-4 are numbered clockwise (first
number); the teeth from mesial towards distal in each of the four Quiz
quadrants (second number). Quiz 25

right mesial left

18 17 16 15 14 13 12 11 21 22 23 24 25 26 27 28
48 47 46 45 44 43 42 41 31 32 33 34 35 36 37 38

The development of the teeth begins in the late embryonic period More info
(stage 18, ca. 44 days, 18 ) with the formation of an epithelial Histologic picture
lamella parallel to the lip edge. This ridge, the labiodental of a dental ridge
(dental lamina).
lamina, later forms a furrow (the labiodental sulcus), out of
which the oral vestibule arises. Dental development also begins
through a ridge-shaped thickening on this labiodental lamina
that is oriented towards the oral cavity. A U-shaped band, the
dental lamina arises.
Through interactions between neural crest cells and ectoderm
10 roundish teeth buds soon grow in the lower and upper jaws on the
labial side of this dental lamina (early fetal period), which represent the
primordia of the deciduous teeth. Somewhat later (ca. 16 weeks) small
buds also form on the oral side. These are the early anlagen of the
permanent teeth.
Dental ridge

The dental
lamina
forms as a
thickening
that during
the course
of its further
developmen
t grows into
the
labiogingival
lamina and
is oriented
towards the
oral cavity.
Each teeth bud has thus an Fig. 12 - Development of a Legend
ectodermal origin and is also tooth
called an enamel organ. On from dental bud to dental bell
the inside, they surround a stage
solidified mesenchyma of Fig. 12
neurectodermal origin that The tooth bud
forms the dental pulp. The arises as a bud
edges of the enamel organ from the U-
grow faster than the middle shaped dental
part, thus forming a dental bell lamina. The
that arises from the tooth bud edges grow
via a cap-shaped stage. The faster than the
mesenchyma around the interior so that
1 Cap-shaped tooth bud
whole tooth bud also 2 Tooth bell soon a bell is
condenses and forms the formed.
dental follicle. Out of it arises
the periodontium and the
cement of the tooth root.
More info
Histological
picture of a dental
bell.
Tooth bell

Out of the dental

lamina a tooth bell
arises that surrounds
neural crest derived
mesenchyma tissue
that becomes the tooth
pulp.

In an enamel organ one distinguishes between an outer and an Synonyms

inner enamel epithelium. Between them is found the enamel Ameloblast
pulp. The enamel epithelium is nourished from the outside by =
Adamantoblast
a capillary network while the enamel pulp always remains
vessel free. The ameloblast layer, which forms out of the
inner enamel epithelium, produces the enamel - in the form
of enamel prisms - in the direction of the tooth pulp.
Immediately next to the inner enamel epithelium the
mesenchyma that lies below it organizes itself into an epithelial
layer, the odontoblast layer, which excretes predentin
towards the outside. Dentin then arises through deposits of
calcium salts in the predentin.

Fig. 13 - Anlage of a Fig. 14 - Structure of the Legend

deciduous tooth enamel organ
with its permanent tooth before and after the beginning
of the enamel deposition
Fig. 13
Section through the
mandible at birth.
One sees the
deciduous tooth that
has not yet broken
through as well as
the permanent
tooth.
Fig. 14
Cellular
structure of the
tooth anlage.
1 Mandible 1 Outer layer of the enamel organ
The inner layer
2 Anlage of the permanent tooth 2 Enamel pulp of ameloblasts
3 Enamel organ 3 Inner layer of the enamel organ form a stratified
4 Enamel (ameloblasts)
5 Dentin 4 Enamel
epithelium that,
6 Labiogingival sulcus 5 Dentin in turn, forms
6 Odontoblasts the dental
enamel.
 More info
Development of the
permanent teeth
from the lamina
dentalis
In both the lower and upper Formation of the molars from Legend
jaw 10 tooth buds emerge the
from the dental lamina for the distal processes of the dental
deciduous teeth. In the middle lamina
fetal period the anlagen for the The three molars
permanent teeth, which will form from distal
processes of the
replace the deciduous teeth, dental lamina
form on the oral side. For the
three molar teeth that arise in
all four quadrants there are no
predecessors. They erupt from
the tongue-shaped, distal
process of the dental lamina,
the extension of the dental 12 Dental lamina
Deciduous tooth (second
lamina. 3 premolar)
Anlage of a molar from the
extension of the dental lamina

Root formation takes place Fig. 15 - Hertwig's epithelial Legend

only after the formation of the sheath
tooth crown is largely Fig. 15
At its edges, the
completed. In the fold, enamel organ grows
between the inner and outer into the depths and
enamel epithelium no enamel forms the Hertwig's
is produced the two layers epithelial sheath,
which will close up
lie closely on one another. This the tooth below
region is also known as except for a root
Hertwig's epithelial sheath. canal (or possibly
Via proliferation the enamel two or three root
1 Outer enamel epithelium canals.
organ grows further into the 2 Inner enamel epithelium
depths in this region and 3 Hertwig's epithelial sheath
preforms the anlage of the 4 Enamel pulp
5 Tooth pulp
later dental root in that the
edges grow conically towards
each other and leave one, two
or three root canals open.

More info
Root formation takes place only after the crown formation has been completed. The transition
from outer to inner layer of the enamel organ lengthens and forms one, two or three root canals.

Root formation Legend

The root canals arise
through extension and
later fusion of the
enveloping layer
between the outer and
inner enamel epithelium.

1 Tooth with two root canals

2 Tooth with three root canals
Development of the salivary glands
The salivary glands arise only in the later embryonic period (stage 18, ca. 44 days, 18 ). More info
The first sign is a thickening of the epithelium on the side of the tongue, outside the anlage of
Histological picture of a
the dental arch (lamina dentalis).
parotid gland.
In the labiogingival sulcus, on the exterior of the labiodental lamina, arises the anlage of the
Histological picture of a
ectodermal parotid gland (serous glands).
sublingual gland.
In the linguogingival sulcus that lies towards the tongue, the sublingual (mucous) and
Histological picture of a
submandibular (sero-mucous) glands arise, which have an endodermal origin. They arise in
submandibular gland.
the hyoid arch region.

More info
Since the submandibular gland does not remain on the mouth floor, but in the course of the
further development grows below the mandibular arch and the mylohyoid muscle, its outlet
crosses over the lingual nerve, which grows into the tongue on both sides.

Parotid gland

The parotid gland is a purely

serous gland. The cell nuclei are
round. (azan staining)

Sublingual gland

The sublingual gland is a mucous gland. The

cell nuclei lie basally and are flattened. (azo
staining).
 Submandibular gland

The submandibular gland is a mixed sero-

mucous gland. In the mucous portion, the cell
nuclei lie flat on the basal membrane, while the
serous portion have round cell nuclei in the lower
third of the cell.
(HE staining)

Definitive pharynx: Derivatives of the first and Quiz

Quiz 03
second pharyngeal arches
Quiz 27

The embryonic pharynx represents the cranial section of the foregut. On the side and ventrally it
is surrounded by the pharyngeal arches. The definitive pharynx consists of three parts:
Nasopharynx
Oropharynx
Laryngopharynx
The differentiated oral cavity now no longer counts as part of the definitive pharynx. An
important property of a developing definitive pharynx is the series of pharyngeal arches and
pouches that are associated with the formation of the thyroid, parathyroid, the tonsils,
middle ear, the thymus, the epithelial bodies and the trachea. It is to be noted that, except
the tonsils and the anlage of the middle ear, which always retain a connection to the pharynx,
these structures have a close relationship with the developing heart. This explains the fact that,
with the progressing descent of the heart, all of these structures come to be located in the
lower neck or thorax region.

Thyroid
The primordium of the thyroid (stage 10, ca. 28 days, 10 ) forms as a median thickening on
the floor between the first and second pharyngeal arches. Soon a bilobular sacculation of the
endoderm develops out of it that remains connected with the mouth floor by the thyreoglossal
duct. This also gets thinner and thinner and, with the further descent, the epithelial cord
disintegrates and the thyroid loses its contact with the mouth floor. What remains is the
foramen cecum.
(see the overview about the descent of the derivatives of the third and fourth pharyngeal
pouches)
Middle ear
On the side, between the first and second pharyngeal arches the first pharyngeal pouch
forms on the inside in parallel with the first pharyngeal cleft that is visible on the outside. This
pharyngeal pouch lengthens but always remains in connection with the throat (tubotympanic
recess for the middle ear cavity and the Eustachian tube). On the one side, its dead-end gets a
close relationship thereby with the inner ear and, on the other, it forms the anlage material for
the eardrum (tympanon) with the bordering ectoderm of the first pharyngeal cleft.
(see the overview about the descent of the derivatives of the third and fourth pharyngeal
pouches)

Tonsils
At the beginning of the digestive tract, at the transition between the oral and nasal cavities in the
oropharynx, there is a large collection of lymphatic tissue. This tonsillar ring (Waldeyer's
tonsillar ring) is composed out of several lymphoreticular organs. A few lymph follicles can be
found scattered in the whole region of the pharyngeal mucosa, especially around the posterior
nasal orifice, on the isthmus of the fauces, the piriform recess and laryngeal inlet.
Accumulations of lymphatic tissue, though, are found especially in the region of the tonsils.
Four lymphoepithelial organs can be distinguished:
pharyngeal tonsil
tubal tonsil
palatine tonsil
lingual tonsil
These lymphatic organs of the tonsillar ring make possible an initial defense against infection at
the beginning of the digestion process and respiration. These lymphatic organs arise only in
the early fetal period since up till birth no defense against infection needs to take place.

The pharyngeal tonsil arises in the roof of the nasopharynx. The mucosa epithelium (here
respiratory epithelium) is opened up in folds thereby and, under it, lymphatic tissue
accumulates. The portion of the lymphatic tissue around the oral tubal pharynx is called the
tubal tonsil.

In the young fetus, the palatine tonsil forms on both sides in the tonsillar sinus as the
remainder of the 2nd pharyngeal pouch. The epithelium thickens and under it a mesenchymal
accumulation forms from material of the second pharyngeal arch. At the same time, the blood
and lymph vessel system also gets denser in this region. Endodermal epithelium cords begin to
grow into the underlying mesodermal blastema and later form the cryptae of the first and
second order.

The lingual tonsil forms on the tongue floor Fig. 16 - Base of the tongue with Legend
in the region of the terminal sulcus. As in the terminal sulcus
palatine tonsil the mucosa invades into the Fig. 16
depths with short canals and forms tonsillar The palatine tonsils and
crypts. These become bounded by lymphatic the lenticular papillae
tissue that is surrounded by a connective are part of Waldeyer's
tissue capsule. On the surface their openings pharyngeal ring.
are crater-shaped depressions in
protuberances of the mucosa, the so called
lenticular papillae of the tongue base.
(see the overview about the descent of the
derivatives of the third and fourth pharyngeal
pouches)

1 Tongue
2 Terminal sulcus
3 Arythenoid prominence
4 Palatine tonsil
5 Lenticular papillae
I Pharyngeal arch
II Pharyngeal arch
III Pharyngeal arch
IV Pharyngeal arch

Derivatives of the third and fourth pharyngeal Quiz

arches Quiz 09

The outwardly visible 3rd and 4th pharyngeal arches are never as effectual as the 1st and 2nd
ones. Normally the hyoid arch (2nd pharyngeal arch) grows over the 3rd and 4th ones
(operculum).

Thymus
The thymus arises from the 3rd and 4th pharyngeal pouches that are found on both sides at the
base of the cervical sinus. The endodermal portions of the thymus anlagen grow on both sides
as ventrally- oriented sacculations of the pharyngeal pouches in the neck mesenchyma (for
comparison, the parathyroid arises from dorsal solidifications). In the segmentation of the
pharyngeal pouches, the neighboring ectoderm of the cervical sinus is also incorporated and
surrounds the endodermal epithelial cords. Through fusion of the two anlagen of the 3rd and
4th pharyngeal pouches and the derivative of the cervical sinus the thymus arises as a
bilobular organ in the anterior upper mediastinum. (see the overview about the descent of
the derivatives of the third and fourth pharyngeal pouches)

Parathyroid glands and ultimopharyngeal

bodies
The parathyroid gland has its origin initially in one of the two More info
dorsal thickenings of the epithelium of the 3rd and 4th The anlagen of the
pharyngeal pouches. Soon afterwards arise the solid parathyroid glands
always consist of
epithelium cords, the epithelial bodies, which with the solid epithelial
descent of the heart are shifted caudally. The anlagen from the cords and lose
3rd pharyngeal pouches (larger portion of the thymus and their contact to the
inferior parathyroid gland) grow faster and get shifted further endoderm of the
pharyngeal
caudally than those of the 4th pharyngeal pouches, (smaller pouches relatively
portion of the thymus and superior parathyroid gland)]. Thus the late (epithelial
superior parathyroid gland gets only to the upper pole of the bodies of the 3rd
median-lying thyroid (stage 17, ca. 41 days, 17 ), while the pharyngeal
pouches in stage
inferior parathyroid glands of the 3rd pharyngeal pouches 19 [ca. 46 days],
reach its lower pole (stage 19, ca. 46 days, 19 ). and those of the
The ultimopharyngeal body arises from bilateral ventral 4th pharyngeal
swellings of the 4th pharyngeal pouches and gets integrated pouches only in
stage 22 [ca. 53
into the thyroid where it forms the parafollicular cells or C- days].
cells for the production of calcitonin.

Legend
Anlage of the lung in stage 10 (ca. 28 days)
Fig. 17
In this median
sagittal section
median buddings of
the intestinal tube
can be recognized:
the thyroid below
the first aortic arch,
the lungs at the
level of the heart
and the liver below
the heart at the
transition to the
umbilical vesicle.

1 Foregut
2 Midgut
3 Rectum
4 Notochord
5 Oropharyngeal membrane / stomodeum
6 Cloacal membrane/ proctodeum
7 Allantois
8 Anlage of the thyroid
9 Anlage of the lungs
10 Anlage of the liver

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