ORGANOGENESIS

DEVELOPMENT OF THE: CENTRAL NERVOUS SYSTEM; PERIPHERAL NERVOUS SYTEM; EYE and EAR

Central Nervous System

 Composed of the brain and spinal cord.
 Functional cells of the CNS are the neurons; the connective tissue cells of the CNS are the gliocytes
(oligodendrocytes, astrocytes, microgliocytes and ependymal cells)
 Develops from the neural tube

Adult Fates of Neural Tube and Neural Crest

 Enlarged cranial portion of neural tube becomes
the brain
 The slender middle and caudal portions
becomes the spinal cord
 Neural canal becomes the ventricles of the
brain and the central canal of the spinal cord.
 Neural crest cells becomes neurons of the
peripheral nervous system (PNS) that have
their cell bodies in ganglia, neurolemmocytes (schwann cells) of the PNS, pigment cells in skin, adrenal
medulla cells, in the head forms mesenchyme (ectomesenchyme) which becomes meninges, bone, fascia, and
teeth.

Formation of Neurons and Gliocytes from Neuroepithelium of the Neural Tube

 Neuroepithelium give rise to neurons, gliocytes (astrocytes and
oligodendrocytes) and ependymal cells of the CNS
 Additionally, the CNS contains blood vessels and a gliocyte
(microgliocyte) derived from mesoderm
 Neuroepithelial cells have processes that contact the inner and
outer surfaces of the neural tube; they undergo mitosis: the
nucleus moves towards the neural canal and the cell becomes
spherical and loses its connection to the outer surface of the
neural tube.
 Some cell division are differential, producing neuroblast that give rise neurons, or glioblast (spongioblast) that
give rise to gliocytes.
 Microgliocytes are derived from mesenchymal mesodermal cells
 Generally, neurons are incapable of cell division, so all neurons must be formed during nervous system
development
 However, in hippocampus and olfactory bulb, some stem cells or neuroblast persist and can give rise to a small
number of neurons postnatally
 Neuroblast and glioblasts lose contact with surfaces of the neural tube and migrate towards the center of the
neural tube wall.

Layers of the Neural Tube Wall

✓ Germinal layer or ventricular zone
➢ Composed of neuroepithelial cells that remain lining the
central canal and designated as ependymal cells
✓ Mantle layer or intermediate zone
➢ Zone high cell density, formed by accumulation of neuroblast and glioblast
➢ Becomes the gray matter of CNS which contains cell bodies of neurons and several gliocytes
✓ Marginal layer
➢ Cell-sparsed zone where axons of neurons and some gliocytes are present
➢ Becomes the white matter of the CNS which contains mainly myelinated axons of neurons.

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 Plates of Neural Tube Wall
 The lateral wall of the neural tube is divided into two plates, alar or dorsal plate and
basal or ventral plate, by the sulcus limitans, a bilateral indentation in the neural
cavity that serves as a landmark to divide the wall
o Basal plates contains cells bodies of efferent or motor neurons that send
axons into the PNS
o Alar plate contains neurons that receive afferent or sensory input from the
PNS
 Midline region of the wall dorsal to the neural canal constitutes the roof plate;
counterpart wall ventral to the neural canal is the floor plate.

Positional changes of the spinal cord: ascensus medullae spinalis

 Initially, the spinal cord runs the entire length of the embryo with spinal nerves passing
through the intervertebral foramina at the levels of their origin.
 Later, however, the vertebral column and the dura grow more rapidly than the
spinal cord, leaving the posterior end of the cord terminating at a gradually
higher level in the vertebral column (asensus medullae spinalis)
 The disproportionate growth also forces the spinal nerves to run obliquely from
the spinal cord to their corresponding vertebral foramina.

Avian Spinal Cord Development

 By 27 hrs of incubation, the neural tube that will become the spinal cord
presents an elliptical neural canal
 Later on the lateral wall of the neural tube becomes greatly thickened while the
dorsal and ventral walls remain thin; the neural canal is compressed laterally to
appear like a vertical slit

Table showing the parts of the neural tube that give rise to the adult spinal cord:

Development of the Brain

 The anterior two-thirds of the neural tube develop into the brain.
 Fusion of the neural folds in the anterior region, and closure of the anterior neuropore, result in the formation
of the three primary brain vesicles from which the brain develops.

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Avian Brain Development
 By 27 hours of incubation of chick embryo the enlarged cranial end of the neural tube forms three brain
vesicles:
◦ Prosencephalon (Forebrain)
 Occupies the rostral part of the head.
Neural canal is prosocoele
 Presents a depression on its floor, the
infundibulum
 The notochord extends up the level of
infundibulum
◦ Mesencephalon (Midbrain)
 Located posterior to and marked off from
the procencephalon by a constriction. Neural canal is mesocoele
◦ Rhobencephalon (Hindbrain)
 Located posterior to and marked off from the mesencephalon by a slight constriction.
Neural canal is rhombocoele
 Continuous posteriorly with spinal cord.
 The cranial neural tube that will develop into brain presents 11 enlargements called neuromeres separated
by constrictions
◦ Neuromeres are a sign of metamerism or segmentation in the embryo
◦ Neuromeres I-III make up the prosencephalon, IV-V the mesencephalon and VI to XI, the
rhombencephalon
 By about 29-30 hours of incubation:
◦ The lateral walls of prosencephalon evaginate to form two primary optic vesicles
◦ The rostral end of prosencephalon presents the anterior neuropore that is almost closed by 33
hours of incubation
 During development, the brain undergoes three flexures which generally disappear (straighten out) in
domestic animals
 Differential growth of the five secondary brain vesicles (telencephalon, diencephalon, mesencephalon,
metencephalon and myelencephalon) gives rise to flexures.

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 ◦ As head folding occurs, the mesencephalon bends ventrally to produce the midbrain flexure
(cephalic flexure).
◦ A second more gradual ventral bend between the hindbrain and the spinal cord is termed the
cervical flexure.
◦ In the rhombencephalon a slight dorsal bending, the pontine flexure, occurs. The pontine flexure is
located in the future pontine region and causes a thinning of the roof of the hindbrain.
 At first, the developing brain shows the same basic structure as the spinal cord, but the flexures produce
considerable variations in the outline of transverse sections at different levels of the brain and in the relative
positions of white and grey matter.
 The sulcus limitans extends anteriorly only to the junction of the mesencephalon and diencephalon.
 Alar and basal plates, which are separated by the sulcus limitans, are therefore only recognizable posterior to
this junction.
 In the diencephalon and telencephalon, however, the alar plates become accentuated and the basal plates
regress.
 By 38 hours of incubation, the three brain vesicles become four due to division of the rhombencephalon
into metencephalon (VI-VII) and myelencephalon (VIII-XI). Rhombocoele of metencephalon is the
metacoele; that of myelencephalon is the myecoele
 By 55 hours of incubation the prosencephalon shows evidence of division into the cephalic telencephalon
and caudal diencephalon, the latter being composed of the optic vesicle and part of prosencephalon lying
between the optic vesicles
 Other changes at 55 hrs of Incubation
◦ Complete torsion of the head region; the left side of the head now lies on the yolk while the right side
is free
◦ Epiphysis (pineal gland) has evaginated from the mid-dorsal wall of the future diencephalon
◦ The optic vesicle has become a double-walled optic cup with a ventral opening, the choroid
fissure.
 By 72 hrs of incubation, the telencephalon presents a median portion called the median telencephalon
and 2 lateral evaginations called lateral telencephalic vesicles.
 Prosocoele of median telencephalon is the median telocoele while prosocoeles of the lateral telencephalic
vesicles are the lateral telocoeles
 By 96 hrs of incubation the infundibulum at the floor of diencephalon has deepened and contacts with the
Rathke’s pocket. The paired ganglia of cranial nerves with sensory components have developed from
cephalic neural crest. These are the:
◦ Semilunar (gasserian) ganglion of 5th (trigeminal) cranial nerve (ganglion V) and the opthalmic and
mandibulomaxillary divisions of trigeminal nerve
◦ Geniculate ganglion of CN VII and acoustic ganglion of CN VIII
◦ Superior ganglion of CN IX
◦ Jugular ganglion of CN X

Specific Development of the Cerebrum

 Right and left lateral telencephalic vesicle becomes right
and left cerebral hemispheres
 Lateral telocoeles expand to become 1st and 2nd ventricles;
a choroid plexus develops that is continuous with a choroid
plexus of the 3rd ventricle via the interventricular foramen
 Mantle layer surrounding the lateral ventricles = basal nuclei
and cerebral cortex = neuroblasts migrate radially to the
surface of the cerebral hemisphere
 Migration occurs in waves: 1st wave = deepest layer,
succeeding waves become succeeding layers to the
surface

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Specific Development of Thalamus and Associated Structures
 Diocoele expands dorsoventrally and becomes the narrow 3rd ventricle; the roof plate is stretched and
choroid plexuses develop bilaterally in the roof of the third ventricle and secrete CSF
 Floor of 3rd ventricle gives rise to the neurohypophysis of pituitary gland
 Mantle layer of diencephalon= thalamus, hypothalamus and epithalamus; the thalamus enlarges to the
point where the right and left sides meet at the middle and obliterate the center of the 3rd ventricle
 Optic nerve and retina develops from an outgrowth of the wall of the diencephalon.

Specific Development of the Corpora Quadrigemina (Mesecephalon)

 Mesocoele becomes mesencephalic aqueduct (which is not a ventricle because it is completely
surrounded by brain tissue and lacks a choroid plexus)
 Alar plate of mesencephalon forms two pairs of dorsal bulges which become rostral (anterior or superior)
colliculi for visual reflexes and caudal (posterior or inferior) colliculi for auditory reflexes
 Basal plate of mesencephalic aqueduct gives rise to oculomotor (III) and trochlear (IV) nerves that
innervate muscles of the eye.

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Specific Development of the Cerebellum
 Arises from bilateral alar plate expansions, called rhombic
lips.
 Rhombic lips, which merge across the midline, contain
three populations of cerebellar neuroblasts.
 Neurons derived from the neuroblast populations make
synaptic connections as they migrate past one another.
 The synaptic connections establish cerebellar circuits
(granule cell neurons to Purkinje cell neurons to
cerebellar nuclei neurons).

Specific Development of the Medulla Oblongata

 Alar plates of the myelencephalon move laterally and the myelocoele expands dorsally to form the 4th ventricle
 Roof of 4th ventricle (roof plate of myelencephalon) is stretched and reduced to a layer of ependymal cells
covered by pia mater (meningus); a choroid plexus develops bilaterally in the roof of the ventricle and secretes
cerebrospinal fluid (CSF)
 Basal plate (with efferent neurons of cranial nerves) is positioned medial to the alar plate and ventral to the 4th
ventricle
 White and gray matters (marginal and mantle layers) become intermixed.

CONGENITAL MALFORMATIONS OF THE NERVOUS SYSTEM

The central nervous system (CNS) is very susceptible to congenital malformations because of the nature of its
complex development. In both humans and domestic animals, CNS anomalies are a comparatively common birth
defect, surpassed in frequency only by congenital cardiovascular abnormalities.

Neural tube defects\

o Failures of the neural tube to close occur most commonly in the cranial and caudal neuropores, but failure to
close at other locations is not uncommon.
o A closure defect of the brain is called cranioschisis, whereas a closure defect of the spinal cord is termed
rachioschisis.
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 o Neural tube defects comprise a group of heterogeneous and complex congenital malformations of the CNS.
 They range from severe structural anomalies resulting from incomplete closure of the neural tube to functional
defects without any obvious structural basis.
 Commonly included in this group are spina bifida, anencephaly, and encephalocoeles.
 Spina bifida
 The most severe of the neural tube defects, includes all
abnormalities in which the vertebral arches fail to close
dorsal to the spinal cord to form the vertebral canal.
 In its simplest form the defect is called spina bifida
occulta referring to a ‘hidden’ occurrence of the
phenomenon.
 The spinal cord and meninges remain in place, but the
vertebral arch of one or more vertebrae is incomplete.
 The meninges may also be distended by fluid
(myelomeningocoele), in which case the phenomenon is referred to as a spina bifida
cystica.
 In many such cases, the spinal cord bulges or is entirely dislocated into the protruding
subarachnoidal space.
 The dislocation causes problems associated with displaced spinal roots; consequently,
neurological symptoms are commonly associated with this condition.
 Meningocele
 The next most severe category of neurotubal defects of the spinal cord
 The meninges herniate and become distended by fluid accumulation.
 The dura mater may be missing in the area of the defect, and the arachnoidea bulges
prominently beneath the skin.
 The spinal cord, however, remains in place and neurological symptoms are usually minor.

Congenital Malformations of the Spinal Cord

o Myelodysplasia
o The general term for a malformation of the spinal cord.
o Such conditions can be classified into the following major categories:
▪ Aplasia is the absence of the development of one or more segments of the spinal cord.
▪ Hypoplasia is the reduced development of segments of the spinal cord.
▪ Hydromyelia is characterized by a dilatation of the central canal due to an excess
accumulation of cerebrospinal fluid.
▪ Syringomyelia means abnormal cavitation of several segments of the spinal cord. It is
considered as a specialized condition of spina bifida, because an occult form of spina bifida
often accompanies syringomyelia. Syringomyelia is generally rare, but it is inherited in
Weimaraner dogs and in tailless Manx cats.
▪ Diplomyelia describes the situation where two spinal cords exist beside each other. Usually
they are covered by only one set of meninges and are contained within one vertebral canal.

Congenital Malformations of the Brain

o Dysraphia of the anterior neural tube
o The cranial neuropore fails to fuse properly or even remains open.
▪ Cranial meningocoele – the meninges protrude through a small defect in the skull.
▪ Meningoencephalocoele – the protrusion involves both meningeal and brain tissue or brain
tissue containing part of the ventricular system and combined with the existence of a larger
opening in the skull.
▪ Exencephaly – characterized by a complete failure of the cephalic part of the neural tube to
close. The vault of the skull does not form, leaving the malformed brain exposed.
▪ Anencephaly – The most extreme cases of cranial dysraphia which results in the absence
of the telencephalon and much of the diencephalon, although the brainstem remains intact.
Anencephaly is rare, but it is most commonly reported in ruminants.

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 o Hydrocephalus
o Is an abnormal accumulation of cerebrospinal fluid within
the ventricular system of the brain.
o In newborn domestic animals is due to an obstruction of the
aquaeductus mesencephali. This prevents the cerebrospinal
fluid of the lateral and third ventricles from passing into the
fourth ventricle and from there into the subarachnoid space,
where it is normally resorbed.
o This type of hydrocephalus is called ‘non-communicating’.
o Other common areas of blockage are the interventricular
foramina and the lateral foramina of the fourth ventricles.
o Hydrocephalus can also result from excessive
cerebrospinal fluid production or impaired fluid resorption
from the subarachnoid space.
o Hydrocephalus is probably one of the most common
congenital anomalies of the nervous system.
o It has been described most often in dogs and cattle, but it
is found in all domestic species.
o In dogs it is most common in small brachycephalic breeds, where the stenosis of the aquaeductus
mesencephali is correlated to the retarded develop ment of the cartilaginous base of the skull.
o In cattle, achondroplastic dwarfs are frequently hydrocephalic.
o Arnold-Chiari Malformation
o Arnold-Chiari malformation is a condition characterized by caudal displacement and herniation
of cerebellar structures through the foramen magnum into the cranial cervical vertebral canal.
o Arnold-Chiari malformation is often accompanied by spina bifida, meningomyelocoele, and
hydrocephalus.
o Microcephaly
o Microcephaly, an abnormally small brain, has been reported in calves, lambs, and piglets.
o Since the size of the cranium depends on the development of the brain, the cranial vault is
considerably smaller than normal.
o External features of microcephaly include a narrow and flattened frontal area of the cranium.
o Cranial bones appear thicker than normal.
o The cause of the condition may be genetic or a prenatal insult such as an infection or exposure to a
teratogen.

o Hydranencephaly
o The cerebral hemispheres are replaced by two fluid-filled sacs.
o The brain stem is usually not affected, but some degree of cerebellar hypoplasia may be observed.
o In domestic animals this condition is most commonly caused by an in utero viral infection (blue tongue
virus in ruminants and panleukopaenia in cats) or, more rarely, by an interruption of the blood supply
to the telencephalon.

o Holoprosencepaly
o Holoprosencepaly comprises a group of congenital malformations of the brain and face that is
characterized by impaired or incomplete midline division of the prosencephalon.
o Associated with chromosomal abnormalities, such as trisomy, various deletions, and other
chromosomal rearrangements, but environmental causes have also been proposed.
o In calves, it represent a failure of the neural tube to separate from the surface ectoderm at its most
rostral extent (rostral neuroporus) that prevents the normal development of the telencephalic vesicles.
o The brain stem and the cerebellum are present although their shape is altered.

o Cerebellar hypoplasia
o An insufficiency of neurons causes hypoplasia of the granular layer of the cerebellum.
o In severe cases, the Purkinje neurons will also be destroyed.
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 o The condition is most frequently found in kittens and calves, most commonly caused by prenatal or
perinatal viral infections.
o The specific viruses are the feline panleukopaenia virus and the bovine virus diarrhoea (BVD) virus.
These viruses exert their maximal effect if the infection occurs during the time of rapid cerebral growth
and differentiationdifferentiation of the external germinal layer at midgestation (from the end of first
trimester to the beginning of last trimester).

o Cerebellar abiotrophy
o Cerebellar abiotrophy is characterized by the degeneration of the Purkinje cells in the already
formed cerebellar cortex.
o It occurs postnatally or, occasionally, prenatally and has been observed in many domestic species.
o The primary defect that causes the degeneration is not known.

Peripheral Nervous System

▪ Consists of cranial and spinal nerves and their ganglia
▪ The cranial and spinal nerves are composed of axons (nerve fibers) of neurons whose cell bodies are
located within the CNS (for cranial nerves) or in ganglia (for spinal nerves)
▪ The connective tissue cells (gliocytes) of the PNS are the neurolemmocytes (schwann cells) that arise from
neural crest and migrates throughout the PNS ensheathing and myelinating axons; and amphicytes that form
satellite cells of nerve cell bodies within ganglia
▪ The Peripheral Nervous System includes all the structures derived from the neural crest cells.
▪ The Neural Crest Cells migrate from the midline and disperse along several pathways.
Fates of the neural crest cells:
o Dorsolateral to the aorta – neural crest cells are grouped segmentally and become the ganglia of the
sympathetic trunk.
o Ventrolateral to aorta – neural crest cells become the abdominal sympathetic ganglia.
o Ventral to aorta – neural crest cells becomes the secretory cells of the adrenal medulla
o Dorsomedial to the somite – neural crest cells becomes segmented and form the spinal ganglia which
contains the nerve cell bodies of sensory neurons.
o In additional, the neural crest cells also form the neuroglial cells in the ganglia and all the Schwann cells that
ensheath the peripheral nerve.
o Beneath the presumptive epidermis (surface ectoderm) – neural crest cells become melanocytes (pigment
cell) of the skin.

Neuron/Nerve cell
– structural and functional unit of the Nervous System
– receive stimuli from environment and transmit impulses to the spinal cord and brain where these impulses are
interpreted; and then to the effector organs for responses
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 • Parts of a neuron
– nerve cell body
– processes
• axon
• dendrites

Neuron Classification Based on the Number of Processes

• Unipolar neuron
• Has only one process, the axon. All afferent neurons are unipolar and have their cell bodies in sensory
ganglia (spinal or dorsal root ganglia and cranial ganglia)
• Pseudounipolar neuron
• Has two processes that arise separately from the cell body, fuse for a while to form 1 process and
divides shortly thereafter into axon and dendrite.
• Bipolar neuron
• Has two processes (one axon and one dendrite) that originate at opposite poles of the soma.
• Multipolar neuron
• Has many processes (one axon and several dendrites) that arise from an oval, pyramidal or stellate
soma.

Functional Classification of Neuron in the PNS

• Based on the direction they conduct impulses
• Sensory or afferent neurons
• Conduct impulses from the receptor
organs to the CNS
• Originate from neural crest and bipolar
cells that subsequently become
unipolar; in case of cranial nerves,
afferent neurons also originates from
placodes.
• Motor or efferent neurons
• Conduct impulses from the CNS to target organs
• Association neurons or interneurons
• Transfer sensory impulses from sensory neurons to efferent neurons.
• Based on the action or origin of the target organs they innervate
• Somatic Neurons
• Innervate voluntary muscles and structures with epithelia derived from ectoderm like skin
• Visceral Neurons
• Innervate involuntary muscles and structures derived from splanchnopleurae like blood
vessels and visceral organs
• Combination of the two
• Somatic afferent neurons
• Somatic efferent neurons
• Cell bodies are in the CNS but their axons extend into the PNS
• Visceral afferent neurons
• Visceral efferent neurons
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 • Constitute the autonomic nervous system (ANS)

Autonomic Nervous System

• Involves two motor neurons
– A preganglionic neuron with cell
body in the CNS (nucleus) but the
axon extends into the PNS
– A postganglionic neuron located
entirely in the PNS (cell body is in
the autonomic ganglion) at positions within the head, or beside vertebrae, or near the aorta or in the
gut wall (parasympathetic and come from sacral and hindbrain regions).

Nerves or Nerve Fibers

• A nerve fiber is an axon of neuron and its sheath that connects
peripheral structures with the brain or spinal cord
• The cell bodies of the nerves are either buried in the CNS to
form nuclei; or massed at some point along the nerve to form
ganglia
• Several nerves from a nerve bundle or nerve fascicle; several
nerve fascicles constitute a nerve trunk
• Majority of the nerves in the PNS end and begin in the spinal
cord and are therefore called spinal nerve. A few (12) end and
begin with the brain and are called cranial nerves.

Spinal Nerves
Spinal nerves are myelinated nerves segmentally arranged along the
spinal cord

Development of Spinal Nerve

• By 96 hours of incubation, the establishment of spinal nerve roots has begun. The adult spinal nerve connects
with the spinal cord by two (2) roots: dorsal and ventral root.
– The dorsal root is the pathway for sensory (afferent) nerve fibers and the ventral root for motor
(efferent) nerve fibers. The dorsal and ventral roots unite outside and lateral to the spinal cord to form
a spinal nerve.
• The neural crest at the level of the spinal cord become the spinal ganglia or dorsal root ganglia. They are
located on the dorsal roots between the spinal cord and the point of union of ventral and dorsal roots. Distal to
the union is a branch, ramus communicans, which extends ventrad to the sympathetic ganglion.

Four Functional Types of Spinal Nerves

• General Somatic Afferent (GSA)
• Arise from cell bodies of pseudounipolar neurons in the spinal ganglion and connect the receptor
organ and the spinal cord.
• Upon exit from the spinal ganglion the axons split into 2 segments:
a longer segment that extends to the receptor organ; and a shorter
segment that forms the dorsal root of the spinal nerve.
• The dorsal root enters the dorsal gray horn of spinal cord where it
establishes synapses with the telodendritic zones of association
neurons.
• Two types of GSA
• Exteroceptive GSA spinal nerves conduct impulses
from afferent nerve endings for touch, pain and
temperature in the skin to the spinal cord.
• Proprioceptive GSA spinal nerves conduct impulses for
positional sense from joints, tendons and muscles to the
spinal cord.

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 • General Visceral Afferent (GVA)
• Arise from cell bodies of pseudounipolar neurons in the spinal
ganglion and connect the receptor organ and the spinal cord.
• Carry impulses from the gut and other visceral organs through
the sympathetic ganglion, with ramus communicans and dorsal
root, to the spinal cord.
• General Somatic Efferent (GSE)
• Arise from cell bodies of large multipolar neurons in the ventral
gray horn of spinal cord.
• Upon exit from spinal cord, the efferent nerves form the ventral
root, which joins with the dorsal root to form a nerve trunk.
• Conduct impulses from the ventral gray horn of the spinal cord to the motor end plates in skeletal
muscles.
• General Visceral Efferent (GVE) together with the GVA, form the
Autonomic Nervous System (ANS)
• Arise from a chain of 2 neurons between the spinal cord and
the target or effector organs.
• The 1st neurons or preganglionic neurons have cell bodies in
the ventral gray horn of the spinal cord that send out
preganglionic axons to the ventral roots and into a separate
autonomic nerve branches, the ramus communicantes, to the
wall of the visceral organs to synapse with the cells bodies of
2nd neurons in the autonomic ganglia.
• The 2nd neurons or postganglionic neurons have cell bodies
in the autonomic ganglia in the visceral wall that send short
postganglionic axons into the viscera.
Cranial Nerves
Although they are arranged according to the same fundamental plan as the spinal nerves, the cranial nerves
have lost the regular segmental arrangement and have become highly specialized. By convention, roman
numerals are used to designate the cranial nerves, with cranial nerve I being the most rostral and cranial nerve
XII the most caudal.
Origin of cranial nerves and their composition
o One of the major differences between cranial and spinal nerves is the tendency of many cranial nerves
to be either afferent or efferent rather than mixed.
o Cranial nerves can be classified into three categories according to their embryonic origin and their
future:
▪ Special sensory function (special somatic afferent or special visceral afferent fibres),
• Cranial nerves I (olfactory) and II (optic)
o Are often regarded more as extensions of brain tracts than as true nerves.
• Cranial nerve VIII (vestibulocochlear)
▪ Mixed nerves that innervate pharyngeal arch derivatives (special visceral efferent and
afferent fibres)
• Cranial nerves V (trigeminal), VII (facial), IX (glossopharyngeal), X (vagus)
▪ General somatic efferent fibres
• Cranial nerves III (oculomotor), IV (trochlear), VI (abducent), XI (accessory)
and XII (hypoglossal)
All of the cranial nerves except the olfactory (I) and optic nerves (II), arise from the brain stem and only the
oculomotor nerve (III) arises outside the region of the rhombencephalon.
Rhombomeres give rise to the efferent nuclei of cranial nerves IV, V, VI, VII, IX, X, XI and XII.
The efferent neurons of the cranial nerves are located within the brainstem, sensory ganglia harbouring the
afferent neurons are situated outside of the brain. The sensory ganglia of cranial nerves originate from
ectodermal placodes or neural crest cells.

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 Functional Types of Cranial Nerves
➢ General Somatic Afferent (GSA). The GSA cranial nerves are similar to the GSA spinal nerves
except that the point of termination is the alar plate of the brain.
➢ General Visceral Afferent (GVA). The GVA cranial nerves are similar to the GVA spinal nerves
except that the point of termination is the alar plate of the brain.
➢ General Somatic Efferent (GSE). The GSE cranial nerves are similar to the GSE spinal nerves
except that the point of origin is the basal plate of the brain.
➢ General Visceral Efferent (GVE). The GVE cranial nerves are similar to the GVE spinal nerves
except that the point of origin is the basal plate of the brain.
➢ Special Somatic Afferent (SSA). The SSA cranial nerves carry impulses from the eye and ear
(involved in the relationship of the animal to external environment) to the alar plate of the brain.
➢ Special Visceral Afferent (SVA). The SVA cranial nerves carry impulses from the
chemoreceptors in the olfactory epithelium and taste buds to the alar plate of brain.
➢ Special Visceral Efferent (SVE). The SVE cranial nerves conduct impulses from the basal plate
of the brain to skeletal muscles with visceral functions.

Special Sensory Cranial Nerves

• Olfactory Nerve (CN I) SVA
– Nonmyelinated; lacks a ganglion
– Arise from olfactory hair cells and terminate in the rhinencephalon
– Develop from the nasal placode.
– These nerve fibres end in the olfactory bulb, where they form special synapses (olfactory glomeruli)
with the mitral cells of the olfactory bulb.
• Optic Nerve (II) (SSA)
– is formed by nerve fibres that are derived from ganglion cells of the primitive retina and terminate
in the superior colliculus of corpora quadrigemina in the brainstem
– Because the optic nerve develops from the evaginated wall of the diencephalon, it is usually
considered as a fibre tract of the brain.
• Vestibulocochlear Nerve (VIII) (SSA)
– Presents an acoustic ganglion beneath the auditory vesicle, caudal to the geniculate ganglion of CN
VII and divided into two kinds of sensory nerve fibres running in two bundles, the vestibular ganglion
and the spiral ganglion.

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 – The vestibular nerve takes its origin from the bipolar neurons of the vestibular ganglion. Supplies the
semicircular canal, utricle and saccule and receive stimuli for the sense of equilibrium.
– The cochlear nerve is formed by axons of the bipolar neurons of the spiral ganglion. Their dendrites
innervate the organ of Corti (spiral organ), and their axons end in the ventral and dorsal cochlear
nuclei in the medulla oblongata, and receive the stimuli for the sense of hearing.

General Somatic Efferent Cranial Nerves

• The corresponding neurons are located in general efferent (motor) nuclei of the brain stem. Their efferent axons
supply the muscles derived from the preotic and occipital myotomes.
• Oculomotor Nerve (III)
• Arise from the basal plate of mesencephalon
• Supplies muscles of the eyeball that are derived from the first preotic myotomes (the dorsal, ventral,
and medial recti, the ventral oblique, and medial portion of retractor muscles).
• Trochlear Nerve (IV)
• Arises from the basal plate of mesencephalon
• The only cranial nerve that leaves the brain stem dorsally and innervates the dorsal oblique muscle.
• Abducens Nerve (VI)
• Arises from the basal plate of myelencephalon
• It supplies the lateral rectus and the lateral portion of the retractor muscle of the eye, both of which
are derived from the most posterior of the three preotic myotomes.

• Accessory Nerve (CN XI)

• It has commisural ganglion, which is a continuation of jugular ganglion
• Motor fibers arise from the posterior myelencephalon and the first 5 or 6 segments of the spinal
cord and extend to the smooth muscles of viscera.
• The special visceral efferent fibres of the cranial root join the vagus nerve and supply the muscles
of the soft palate and the intrinsic muscles of the larynx.
• The general somatic efferent fibres of the spinal roots innervate the sternocephalicus,
cleidomastoideus and trapezius muscles.
• Hypoglossal Nerve (CN XII)
• Predominantly motor nerve fibers that arise separately from the posterior myelencephalon and form
a single trunk to supply the skeletal muscles of the tongue.

Mixed Cranial Nerves

• The cranial trigeminal (V), facial (VII), glossopharyngeal (IX), and vagus (X) nerves supply derivatives of
the pharyngeal arches.
• The cranial root of the accessory nerve (XI) is an extension of the vagus nerve. The spinal roots arise from
the spinal cord (from the five or six most cranial cervical segments).
• Trigeminal Nerve (CN V)
• Innervates the first pharyngeal arch.
• It is mainly general somatic afferent and is the principal afferent nerve of the head.
• Its large trigeminal ganglion is situated beside the rostral end of the pons and its neurons originate
from the most anterior part of the neural crest.
• The centrally running processes from this ganglion form the large sensory root of the trigeminal nerve,
which enters the lateral portion of the pons.
• The peripheral processes separate into three large divisions, the ophthalmic, maxillary and
mandibular nerves.
• Their afferent fibres innervate the skin of the face as well as the mucosa of the mouth and nose.
• The efferent fibres of this cranial nerve arise from neurons in the most anterior part of the special
visceral efferent column in the metencephalon and form the special visceral efferent nucleus of the
trigeminal nerve, which lies at the midlevel of the pons.
• Facial Nerve (CN VII)
• Supplies derivatives of the second pharyngeal arch.
• The special visceral efferent nuclei are located in the special visceral efferent column in the caudal
part of the pons. The efferent fibres of these neurons are distributed to the muscles of facial expression
and to other muscles developing from the mesenchyme of the second arch.
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 • A small general visceral efferent portion of the facial nerve ends in the peripheral autonomic ganglia
of the head.
• The geniculate ganglion provides the special visceral afferent fibres of the facial nerve. The
peripheral processes pass to the greater superficial petrosal nerve and, via the chorda tympani, to the
taste buds of the anterior two-thirds of the tongue. The central processes of the geniculate ganglion
enter the pons.
• Glossopharyngeal Nerve (CN IX)
• Innervates the third pharyngeal arch. It forms several rootlets which arise from the medulla
oblongata, just caudal to the otic vesicle, the primordium of the inner ear.
• The special visceral efferent fibres arise from nuclei in the myelencephalon and innervate muscles
of the third pharyngeal arch located in the region of the pharynx.
• The general visceral efferent fibres run to the otic ganglion. The postganglionic axons pass to the
parotid gland and to the posterior lingual glands.
• Special visceral afferent fibres innervate the taste buds of the posterior part of the tongue.
• Vagus Nerve (CN X)
• Results from the fusion of nerves of the fourth to sixth pharyngeal arches. Its large general visceral
efferent and general visceral afferent components innervate the heart, the foregut and its
derivatives, and a large part of the midgut.
• Special visceral efferent fibres originating from the fourth pharyngeal nerve (cranial laryngeal nerve)
innervate the cricothyroid muscle, whereas analogous fibres from the sixth arch gives rise to the
recurrent laryngeal nerve, which supplies the remaining intrinsic laryngeal muscles.

Differences Between Cranial and Spinal Nerves

• Ventral and dorsal roots are primitive in cranial nerves and do not unite outside the brain.
• Cranial nerves are not segmentally arranged.
• Cranial nerves are more highly specialized.
• Peripheral nerves establish contact early with the nearest somite, somitomere, placode or pharyngeal arc and
innervate derivatives of these embryonic structures.
• Innervation continuity is retained even when the derivatives are considerably displaced or when other structures
have obstructed the pathway.
• The early establishment of an innervation connection
explains why some nerves travel extended distances
and make detours to reach distant inaccessible targets.

Formation of Meninges
• Meninges surround the CNS and the roots of spinal
and cranial nerves.
• Meninges originate as two layers of mesenchyme
surrounding the embryonic neural tube.
• Three meningeal layers (dura mater, arachnoid and pia
mater) are formed as follows:
– The outer layer forms dura mater.
– Cavities develop and coalesce within the inner
layer dividing it into arachnoid and pia mater;
the cavity becomes the subarachnoid space
with contains CSF

Congenital Malformations Of The Peripheral Nervous System

Aganglionic Large Intestine

• Absence of enteric ganglia in the wall of the affected segments of the colon.
• Aganglionic large intestine arises from both dominant and recessive mutations.
• Manifested by great dilatation of certain segments of the colon,
• Has been reported in several species including horses and mice.
• In humans it is called ‘Hirschsprung’s disease’.
• Reported in white foals, born from crossings of Overo horses.

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 • Overo is a spotting pattern in Painted and Pinto ponies in which melanocyte-free, white skin areas appear
preferentially in the ventral midline of the trunk and the distal parts of the limbs and snout.
• The condition starts with signs of colic within one day of birth and the affected foals die shortly thereafter.

Congenital Defects Affecting Schwann Cells

• Abnormalities affecting the myelin production by Schwann cells in the peripheral nervous system
• Results in a hypertrophic neuropathy that has been found in Tibetan Mastiff dogs.
• Clinical signs of dysphagia and chronic ruminal bloat developed after weaning, which were attributable to
bilateral vagus nerve degeneration.
• Trunks of the sciatic nerves and brachial plexuses were similarly affected resulting in a weak shuffling gait.

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