
Donald Kennedy (Stamford
University) suggested the following lines (they may be sung to the
tune of "It's a long way to Tipperary"):
Goodbye, fins
and gill-slits
Hello, teeth and hair
It's a long way from Amphioxus
But we all came from there ...
It is a reminder that we
still use embryonic processes that first arose in early chordate
ancestors.

pharyngeal or branchial?
During the 4th and 5th weeks
of development, the human embryo closely resembles a fish embryo. In
particular, the series of parallel ridges seen on the side of the
future head and neck region (in relation to the pharynx) resemble
the developing gill arches of the fish embryo. This resemblance
prompted the earlier embryologists to refer to the corresponding
arches in the human embryo as branchial arches, where ‘branchium'
means ‘gill'. (Not to be confused with ‘brachium', which means
‘arm'!) However, at no stage do perforations occur between the
arches of the human embryo - as they do in fish embryos to form the
gill-slits - and it has now become more acceptable to refer to these
structures as pharyngeal arches, rather than branchial arches. Be
prepared to hear both terms - clinicians especially will tend to
refer to branchial arches, branchial cysts, etc - but try to use the
term ‘pharyngeal arches' when you can.
pharyngeal arches,
pouches, and clefts
A total of six pairs of
pharyngeal arches develop in cranio-caudal sequence. The first arch
is complex, and contributes to both upper jaw and lower lower jaw.
The 5th arch is very rudimentary in the human embryo, and difficult
to define.
The external surface of the
embryonic pharyngeal region is covered with ectoderm, and the
internal aspect lined with endoderm. Between these layers is a core
of mesoderm.
Between successive arches,
there are indentations - both on the external (ectodermal) aspect
and the internal (endodermal) aspect. Those on the outside are
called clefts, while those on the inside are called pouches.
Become familiar with the
terms:
 |
pharyngeal arches |
 |
pharyngeal clefts |
 |
pharyngeal
pouches |
because they will be used
frequently as we study development of this region.

Pharyngeal region of the embryo

pharyngeal arch
derivatives
The mesodermal core of each
pharyngeal arch differentiates into three main types of tissue, and
each arch becomes associated with a particular cranial nerve.
The tissues are:
 |
skeletal tissue (eg: cartilage, bone, ligaments) |
 |
muscle tissue (striated musculature, but not all
under voluntary control) |
 |
arterial arch (which may or may not be converted
into a definitive major artery). |

Skeletal elements within the
pharyngeal arches
The Table below summarises
these important relationships between the arches, their derivatives,
and cranial nerve supply.
|
arch |
skeletal element |
muscles |
cranial nerve |
|
1 |
(Meckel’s cartilage)
malleus
incus |
muscles of
mastication
anterior belly of
digastric
tensor tympani
tensor palati |
5 trigeminal
(mandibular division) |
|
2 |
stapes
styloid process
upper part of hyoid |
stapedius
stylohyoid
muscles of facial
expression
posterior belly of
digastric |
7 facial |
|
3 |
lower part of hyoid |
stylopharyngeus |
9 glossopharyngeal |
|
4-6 |
laryngeal cartilages |
cricothyroid
pharyngeal
constrictors
intrinsic laryngeal
muscles |
10 vagus |
You will find that a clear
understanding of these relationships between pharyngeal arches,
derived structures, and nerve supply will be of great help when
studying the arrangement of the head and neck and disorders of
development in those regions.
pharyngeal pouches
on the inside of the
embryonic pharyngeal region are endodermally-lined grooves between
adjacent pharyngeal arches. They contribute to the development of a
surprising diversity of structures, ranging from the middle ear
cavity to endocrine glands and components of the lymphatic system.
The last two pouches are difficult to distinguish clearly, and are
often considered as a single unit.
pharyngeal clefts
are ectodermally-lined
grooves on the outside of the embryonic pharynx. Only the first
cleft is important - it develops into the external auditory meatus
of the ear and provides the outer epithelium of the tympanic
membrane. It used to be believed that the remaining clefts were
‘submerged' beneath overgrowing folds and then disappeared, but more
recent work shows that the clefts simply ‘fill out' by proliferation
of the underlying mesoderm.

Derivatives of the pharyngeal
pouches and clefts
The Tables below summarise
the derivatives of the pharyngeal arches and clefts:
|
pouch |
derivatives |
|
1 |
middle ear cavity,
endodermal aspect of tympanic membrane, pharyngotympanic
tube |
|
2 |
palatine tonsil |
|
3 |
inferior parathyroid
gland, thymus |
|
4 & 5 |
superior parathyroid
gland, parafollicular cells of thyroid gland |
|
cleft |
derivatives |
|
1 |
external auditory
meatus, ectodermal aspect of tympanic membrane |
|
2 - 4 |
cervical sinus? |
tongue
You have probably noticed
that the tongue has a complex nerve supply - 5 different cranial
nerves are involved. It is however possible to make sense of this by
learning about the embryological origin of the tongue.
Note that the tongue develops
in the floor of the pharynx in relation to the ventral ends of the
pharyngeal arches.
Most of the musculature of
the tongue is derived from myoblasts which migrate in from the first
four somites - the occipital somites. The only exception is
palatoglossus, which develops from mesoderm in the facial region.
The occipital myoblasts bring with them their nerve supply: the
hypoglossal nerve (cranial nerve 12). The course of the hypoglossal
nerve as it loops down and around the great arteries of the neck
before turning up into the tongue reminds us of the migration made
by the myoblasts.
The body of the tongue is
formed by the fusion of several mesodermal swellings, and the nerve
supply to the endodermal epithelium which covers them is derived
from the neighbouring pharyngeal arches.
In addition to the general
sensations of touch, pain, pressure and temperature we must also
take into account the special sense of taste.
As you study development of
the tongue, notice how the thyroid gland originates from the
endoderm near the centre of the tongue, and then begins a long
migration down into the neck region. It remains linked with the
tongue for a while by a duct - the thyroglossal duct - but normally
this connection disappears by the end of the embryonic period,
leaving only a small depression (foramen caecum) in the surface of
the tongue.
abnormal migration
Whenever you hear about
sequences of development that involve the migration of cells or
structures, bear in mind the possibility that this migration may go
wrong, with cells going too far, or stopping short of the correct
destination, or migrating to one side or another of the correct
path. Read in your textbooks about the abnormalities of this type
affecting the thyroid, parathyroid, and thymus glands.

Questions on the
pharyngeal arches:
1 What names are given
to the main subdivisions of the first pharyngeal arch?
2 In relation to which
pharyngeal arches does the ear develop?
3 Initially, the
entrance into the mouth is closed by the buccopharyngeal membrane.
This marks the transition from the ectodermal stomodeum to the
endodermal foregut. Thus, which parts of the definitive mouth have:
 |
an ectodermal origin? |
 |
an endodermal origin? |
4 Describe normal
development of the mandible. Is it formed by endochondral
ossification or intramembranous ossification?
5 Explain why the
muscles acting on the ossicles of the ear have different nerve
supplies.
6 What is meant by the
first arch syndrome?
7 Why is the second
pharyngeal arch sometimes called the hyoid arch?
8 Which branch of the
vagus nerve supply the developing larynx? Is there anything unusual
about the course taken by these nerves?
9 What is the
embryological origin of the tympanic membrane (eardrum)?
10 What happens to
pharyngeal clefts 1 - 4?
11 Which migrate further
during development - the superior parathyroid glands or the inferior
parathyroid glands?
12 List the cranial
nerves supplying the tongue. What is the embryological explanation
for these supplies?
13 How do the palatine
tonsils develop?
14 What is a branchial
cyst? How can it be distinguished from a thyroglossal cyst?
15 What is a lingual
thyroid?
16 When does the tongue
become mobile? |