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Neuroanatomy Notes 2010 4 PDF Free
Neuroanatomy Notes 2010 4 PDF Free
Neurons are polar and generally carry impulses from dendrites past the neuronal cell
body to an effector axon. They are thus part of either afferent / sensory or
efferent / motor pathways.
Most functional pathways involve chains of neurons (typically three) with one or more
interneurons, creating potential for filtering or modification of transmitted messages.
Grey matter of the CNS is the site of aggregated cell bodies; white matter is
predominantly bundled tracts of myelinated axons.
Grey matter is therefore more metabolically active and susceptible to insults eg. anoxia.
On the external surface of the forebrain and cerebellum, grey matter forms an external
cortex [L. = bark]; deeper in the brain grey matter exists as isolated nuclei less visible to
the naked eye. In the spinal cord, grey matter is organized into continuous longitudinal
columns, displaying a butterfly shape with dorsal and ventral horns in cross-section.
A bundle of parallel axons is called a nerve in the PNS, but in the CNS can be variously
called a tract, fasciculus, peduncle, or lemniscus.
Most pathways in the CNS cross over or decussate, thus relate to the opposite or
contralateral side of the body (same side = ipsilateral).
Knowledge of the level at which decussation occurs is often clinically important in
localizing the site of neurological lesions.
Pathways may be designated as either somatic (‘of the body’; broadly, relating to
interactions with outside environment) or visceral (related to internal organs). In general,
somatic pathways are more likely to have conscious awareness and voluntary control.
SOMATIC VISCERAL
Motor to Skeletal muscle Smooth & cardiac muscle; glands
Thoracic & abdominal organs; visceral
Tendons/ligaments,
activity; stretch receptors; blood
Sensory to joints (proprioceptive)
pressure; O2 & CO2 tension;
skin (exteroceptive)
temperature
When including the cranial nerves, these are sometimes divided into special vs general
pathways to recognize those of restricted distribution associated with the special senses:
These divisions have an anatomic basis - in the spinal cord afferent neurons lie in the
dorsal horn, efferents in the ventral horn. The white matter columns (funiculi) show the
same pattern (though less strict) where:
dorsal column - entirely afferent, except in a few species.
ventral column – generally efferent
lateral column – mixed afferent & efferent
Afferent DRG
GSA
GVA
Mixed Afferent
R•F
& Efferent
GVE
GSE
Efferent
Visceral and somatic components are also separated with visceral pathways generally
closer to the lateral base of each horn. The area around the central canal is the site of
the primitive reticular formation.
The neuronal bodies of afferent neurons lie outside the spinal cord in a dorsal root
ganglion that sits in the region of each intervertebral foramen. DRG are cells of neural
crest origin that grow back into the spinal cord.
In the brainstem, the basic SA-VA-VE-SE pattern is preserved but is „flattened out‟
(afferents = lateral, efferents = ventral/medial) as the central canal dilates and „peels
open‟ to form the 4th ventricle. In the brainstem the longitudinal grey columns („horns‟)
become fragmented into elongated islands, now called nuclei.
The pattern of these brainstem nuclei can be useful since their dysfunction has distinct
functional consequences that can locate brainstem lesions very precisely.
4th ventricle
GSA
SSA
Afferent
SVA
GVA
GVE
Efferent
GSE
The bodies of somatic efferent neurons - lower motor neurons - lie in the ends of the
ventral horns and extend effector axons all the way to the muscles they supply. All brain
and spinal cord motor pathways above the LMN (and its associated reflex arcs) are
termed upper motor neurons. The distinct symptoms associated with UMN vs LMN
dysfunction are clinically very important in locating neurological lesions:
LMN UMN
Can be excitatory or inhibitory of LMNs;
Always excitatory; essential for muscle
predominant net effect is usually
contraction
inhibition of extensors
Damage causes: Damage causes:
hypotonia / flaccidity hypertonia / spasticity
weak/absent reflexes exaggerated reflexes
severe weakness / paralysis slight weakness / paresis
rapid & profound muscle atrophy slow, modest muscle atrophy
(neurogenic atrophy) (disuse atrophy)
The LMNs of limb muscles lie in the slightly swollen regions of the brachial and
lumbosacral intumescences of the spinal cord.
eg. An injury of the region of the brachial intumescence will cause a mix of LMN
symptoms in the forelimb, and UMN signs in the hindlimb.
Robinson & Huxtable Clinicopathologic Principles for Veterinary Medicine Cambridge UP 1988
The spinal cord is shorter than the vertebral canal; spinal nerves therefore originate
rostral to their associated intervertebral foramen, particularly in the lumbosacral region.
Sympathetic Parasympathetic
Physiological response „Fight or flight‟ „Relaxed‟
Short; ganglia close to spine
Pre-ganglionic fibres Long; ganglia close to organs
(sympathetic chain)
Post-ganglionic fibres Long; adrenergic Short; cholinergic
Localisation More diffuse More discrete
Craniosacral
Outflow Thoracolumbar
(mostly vagus nerve)
Almost all organs have dual sympathetic/parasympathetic innervation (Exceptions:
adrenal, peripheral blood vessels (controlled by „sympathetic tone‟), sweat glands).
The sympathetic chain runs parallel to the spinal cord (in the roof of the thorax and
abdomen) and is linked to spinal nerve roots by the ramus communicans. It is absent in
the neck – the ascending sympathetic nerves combine with the descending vagus nerve
(parasympathetic) to form the bi-directional vagosympathetic trunk.
Afferent DRG
GSA
GVA Spinal
Nerve
GVE ramus communicans
GSE
Sympathetic chain
Efferent ganglion
Autonomic ganglion
Visceral
Nerve
Segmentation
The nervous system is segmented; fibres at each spinal level are embryologically
dedicated to a specific myotome (segmental muscle block) and dermatome
(dermal/connective tissue block).
The fidelity of mature nerves to these blocks allows mapping of sensory fields usually
termed dermatomes. Loss of sensation to a particular area can be tracked back to
known spinal level, particularly on the trunk.
Cranial nerves
Are numbered from rostral to caudal in Roman numerals (I to XII), with a few anomalies:
CN II (optic „nerve‟) is really a direct outgrowth of the brain
CN V (trigeminal) really should be three nerves (as its name suggests)
CN XI (accessory) is really a compound, rostrally-directed spinal nerve
Unlike spinal nerves, these are not regularly spaced and segmentation is not
obvious in the adult, but is clear is their derivation from the embryologic branchial
arches. From the branchial segments, dorsally-derived CNs are typically mixed,
while ventrally-derived CNs are motor.
Branchial Derived CN – Dorsal root CN –Ventral root
arch muscles derivative (mixed) derivative (motor)
Oculomotor (III)
1 (Mandibular) Masticatory Trigeminal (V)
Trochlear (IV)
2 (Hyoid) Facial Facial (VII) Abducens (VI)
3 Pharyngeal Glossopharyngeal (IX)
4-6 Laryngeal Vagus (X)
Hypoglossal (XII)
7 Neck Accessory (XI)
* CN I, II, VIII are specially-derived pure sensory nerves, not segmental nerves. Per Kardong 2009
INTRACRANIAL ANATOMY
Arterial Supply
The blood supply to the CNS is very rich: it is 2% of body weight but receives 16% of
cardiac output.
As a rule the CNS has few large arteries within its substance, probably because
pulsating vessels are not „tolerated‟ by delicate nervous tissue.
Arterial supply to the brain is via the cerebral arterial circle (Circle of Willis) which
surrounds the pituitary.
Potential supply channels into the CAC:
1. internal carotid a.
a. directly; or Dog, man, most mammals
b. via anastomosis with internal maxillary a. Sheep, Cow, Cat
c. via anastomosis with vertebral a. Cow
2. vertebral aa. basilar a.
Caudal parts of the brain eg. occipital lobes & cerebellum are mostly reliant on the
vertebral supply.
All species have vertebral artery supply but differ in how much of the cerebrum this
supplies. This can have implications for the effectiveness of severing carotids in
euthanasia, eg. in ritual slaughter (very effective in sheep but less so in cattle)
Radiating from the CAC are 3 (rostral, middle, and caudal) cerebral arteries and the
rostral cerebellar artery. Penetrating branches are generally tiny and are usually
functional end arteries.
The pattern of cerebral blood supply is very important in humans due to our
predisposition to ‘strokes’, but these are much less common (and less debilitating) in
veterinary species.
Rete mirabile [L. ‘marvelous net’] occur on the anastomosis of the internal carotid and
internal maxillary (ie. in cat, sheep, cattle). Function is unknown: eliminates pulse?
countercurrent thermoregulation? In all species, the last part of the internal carotid
passes through the cavernous sinus which may also allow some heat exchange.
Arterial supply to the spinal cord is via a pair of dorsal spinal aa. and a single
ventral spinal a. These tend to anastomose with each other, and with the supplying
dorsal and and ventral root arteries of the spinal branches (from the aorta or
vertebral a.) that penetrate at each intervertebral segment.
From these the inner zone of the spinal cord is supplied by the ventral vertical artery,
the outer zone by smaller penetrating radial arteries.
Venous Drainage
The CNS is drained by an extensive system of valveless venous sinuses, all extradural.
Unlike arteries, some of the cerebral veins sit deeper in the folds of the brain. Veins
within the brain itself are very thin-walled and valveless.
The spinal cord drains to the internal vertebral venous plexus, a pair of longitudinal
ventral epidural veins anastomosing at each segment (like train tracks). The IVVP drains
to the azygous (thorax) or caudal vena cava (abdomen).
Despite valves protecting retrograde entry of blood into the vertebral sinuses, this can
occasionally occur during abdominal straining, providing a route (termed ‘paradoxical
embolism’) for bacterial or tumour seeding from abdominal organs into the CNS.
Meninges
These are three membranes surrounding the brain and spinal cord:
1. Dura (outermost)
A tough fibrous layer. Adherent to the periosteum in the cranium but separate in
spine, creating an epidural space containing semi-fluid fat.
[Below the dura is the subdural space – normally only a potential space
but can fill with haematoma from a ruptured vessel etc.]
2. Arachnoid
A thin membrane pressed against the dura by CSF pressure. From it spider‟s
web-like projections (trabeculae) connect to the pia.
[Beneath arachnoid is the subarachnoid space – filled with CSF]
3. Pia mater (innermost)
A very thin membrane covering (& semi-fused to) the brain and spinal cord.
Contains numerous small blood vessels which makes it visible.
Dural specializations:
There are two large „baffles‟ projecting inwards within brain fissures:
1. Falx cerebri – midsagittal. Contains the dorsal sagittal sinus.
2. Tentorium cerebelli – within the transverse fissure separating cerebrum and
cerebellum. Often ossifies with age.
Dural sheaths also extend along special sensory nerves, ie. olfactory, optic &
vestibulocochlear nerves.
These dural sheaths provide portals for tracking of extracranial infection into the brain.
In most species the spinal cords ends approximately the lumbosacral junction (dog L7;
cow/horse S1; cat/pig/sheep S2). Much shorter in man – ends at L1.
At the termination of the spinal cord (the conus medullaris) the meningeal tubes taper
and fuse into the filum terminale which anchors the spinal cord to the coccygeal
vertebrae and prevents rostral movement of the spinal cord.
Infections at the root of the tail (eg. tail-biting vice in pigs) can cause epidural abscess or
ascending infection into neural canal.
Epidural anaesthesia:
Injection of anaesthetic or analgesic into the semi-liquid epidural fat allows local
diffusion to anaesthetize nerve roots. However access is harder in animals than in
humans.
The most common veterinary application is in cows (ie. to prevent straining in calving)
where epidural injection is performed between1st & 2nd coccygeal vertebra. Can be done
at L6-S1 but at this site it is possible to enter the dural tube and lumbar cistern (or
damage nerves forming cauda equina).
The subarachnoid space is expanded at certain points into several cisterns where CSF
can be collected by cisternal puncture:
Cisterna magna (aka cerebromedullary cistern) between caudal cerebellum
and medulla
Lumbar cistern which can be accessed via the lumbosacral interarcuate space
in some species only, eg. difficult to obtain fluid in dog but useful in cow &
human.
Cerebrospinal Fluid
CSF is a thin, colourless fluid formed from:
1. selective leakage (dialysis) from vessels suspended in the pia mater
2. secretion by ependymal cells lining the central canal & ventricles
3. the choroid plexuses, which are leaky tufts of arterioles, pia, and ependymal
epithelium within the four brain ventricles [=major source]
CSF (about 150ml total in man) is replaced several times a day (~ 30ml per hour in dog),
acting to flush the system. Its potential to carry hormones is suspected but not well
understood.
CSF drainage is via subarachnoid venules, or via arachnoid villi (aka arachnoid
granules), tuft-like evaginations of the arachnoid into the dorsal sagittal sinus. CSF
flows from production in the ventricles, out through lateral apertures of 4th ventricle
(below cerebullum), to the subarachnoid space, to resorption by arachnoid villi.
CSF flow is interior to exterior - failure of intracerebral flow produces internal
hydrocephalus (dilated ventricles), failure of drainage causes external hydrocephalus
(dilation of subarachnoid space).
Blood-brain barrier
Brain capillaries are made selectively permeable by (i) tight junctions between
endothelial cells and (ii) an extra coat from astrocyte „feet‟. A similar system restricts flux
into the CSF.
The BBB restricts passage of large molecules such as proteins and some drugs, eg.
antibiotics into the CNS.
THE BRAIN
The brain develops from infolding of the neural plate to form the neural tube; the tiny
central canal and brain vesicles are derived from the lumen of this tube. Pinching of the
tube forms three primary vesicles corresponding to the fore, mid and hindbrain. The
telencephalon froms from bilateral vesicles that ballon out to eventually cover most of
the caudal parts dorsally.
Cortex
Basal nuclei Lateral
F Telencephalon
Limbic system ventricles
O
Rhinencephalon
R Prosencephalon
Epithalamus
E
Thalamus
Diencephalon 3rd ventricle
Hypothalamus
[Retina]
M
Tectum (colliculi) Cerebral
I Mesencephalon
Tegmentum aqueduct
D
H Pons
Metencephalon
I Cerebellum
Rhombencephalon 4th ventricle
N
D Myelencephalon Medulla oblongata
Central
Caudal neural tube Spinal cord
canal
TELENCEPHALON
Associated with:
higher processing and association („intelligence‟) Cortex
regulation of motor function Basal nuclei
smell Rhinencephalon
memory; emotions Limbic system
NB. The term cerebrum describes the telencephalon generally, but is often used in
reference to the neocortex only.
Cortex
The cortex is heavily folded to increase the surface area of its external grey matter
(convex folds = gyri, grooves = sulci). It is loosely divided into four regions based on the
overlying bones: frontal, parietal, temporal and occipital lobes. The cruciate sulcus
divides the frontal and parietal lobes.
The primary projection areas of the cortex are associated with specific somatic tracts
projecting via the thalamus. These are surrounded by association areas that function in
processing and cognition of stimuli (as much as 85% of the cortex in man, but only
around 20% in animals).
The primary somatic motor area and primary somatic sensory area are always adjacent,
usually around the cruciate sulcus.
Lesions of cerebral hemispheres tend to produce circling and turning of head (towards
the side of the lesion), but without a head tilt as seen in vestibular disease. Other signs
include compulsive walking, ‘head pressing’, a change of ‘personality’, or epileptic fits.
Depending on the projection area affected, there may be UMN signs ie. exaggerated
reflexes (though in the dog the effect of motor cortex destruction is actually very
subtle), proprioceptive deficits, or blindness on one side of visual fields.
Limbic system
This includes the cortical gyri lining the longitudinal fissure (cingulate gyrus), the
hippocampus, fornix and amygdala [L. = almond]; also links to the hypothalamus.
The hippocampus [L. seahorse] is phylogenetically a very ancient part of the
rhinencephalon and was once part of the primitive motor cortex. It remains closely linked
to the piriform lobe thus olfaction; also richly connected to the hypothalamus therefore
the ANS.
Functions of the limbic system:
emotions – fear, etc.; emotionally conditioned behaviour
spatial and short-term memory. The hippocampus seems to be essential for
temporary retention of memory, particularly spatial, before it is „hard-wired‟ into
the cortex.
the amygdala is active in aggressive behaviour or fear
cingulate gyrus damage causes disorder of complex behaviours
The limbic system can be the site of unusual seizures known as psychomotor seizures,
sensory (eg. taste) hallucinations that can be isolated but usually precede more
generalized seizures.
Cingulate gyrus
Hippocampus
For nix
Septal
nuclei
Mammillar y
body
Olfactory input Hypothalamus
Amygdala
(piriform lobe)
Commissures
The left & right cortex interconnect at commissures. The major commissure is the
corpus callosum, formed by transverse fibres exchanging between hemispheres.
Surprisingly cutting of the corpus callosum induces subtle lesions only detected by
careful testing, eg. tasks requiring transfer of learning from one hemisphere to the
other.
Minor commissures occur at the ventral anterior commissure (between olfactory &
piriform cortex), and at the commissure of the fornix of the hippocampi.
Basal nuclei
Sometimes called „basal ganglia‟ though they are not true ganglia; these include the
caudate nucleus, putamen & globus pallidus (pallidum). These sit on either side of the
internal capsule which is the radiation of fibres projecting to/from the thalamus. Part of
the so-called extrapyramidal motor afferent system; lesions should therefore cause UMN
signs but in practice are often difficult to detect.
DIENCEPHALON
Epithalamus
The thin roof plate of the 3rd ventricle – part of which is only closed by pia. Mainly
consists of the pineal gland (epiphysis), which releases melatonin in response to
photoperiod, thus regulating „circadian rhythms‟, hormonal & reproductive cycles etc..
Thalamus
Two large expansions of the lateral walls of the 3rd ventricle that bulge inwards to meet
as the interthalamic adhesion (thus making the 3rd ventricle a ring shape). Mostly grey
matter in the form of many close-packed but distinct thalamic nuclei related to specific
pathways.
Functions of the thalamus:
All afferent tracts (except olfactory) synapse in the thalamus, making it the
principal relay centre for sensory information projecting to the cortex.
Projects stimuli directly via specific point-to-point projections, but also diffusely to
association areas.
Some processing or filtering of sensory inputs.
Completes and somewhat participates in motor circuits, eg. from the basal nuclei
and cerebellum
The caudolateral part of the thalamus (metathalamus) has two small bulges representing
the medial and lateral geniculate bodies, nuclei relaying acoustic and visual pathways
respectively.
Hypothalamus
This is the portion of the thalamus visible ventrally in an intact brain. It functions in the
autonomic nervous system, in
directs both parasympathetic and sympathetic ANS, eg. heart rate, vasomotor,
iris dilation etc.
integrative control of homeostasis (temperature, thirst, appetite, gut motility, etc.)
emotion (rage, aggression, etc.) and „animal‟ behavioural states –
sleep/wakefulness, sexual function, etc.
neuroendocrine control:
directly (ADH, oxytocin)
via the hypophysis (pituitary gland) to which it is attached by the infundibulum.
The hypothalamus and hypophysis pars nervosa are in direct continuity.
Two small mammillary bodies on the underside of the hypothalamus act as relays
between the hippocampus and the thalamus (mammillothalamic tract)
MESENCEPHALON
The midbrain; a short segment which re-forms the typical tube-like, spinal cord-type
architecture around the mesencephalic aqueduct. Has a dorsal sensory tectum and
ventral motor tegmentum:
The tectum has two pairs of swellings the rostral and caudal colliculi (aka superior
and inferior), which link with the associated geniculate bodies and act as relays and
integration centres for reflex pathways of the visual and auditory inputs respectively.
The tegmentum contains the nuclei of CN III-V, the red nucleus (origin of the motor
rubrospinal tract), the substantia nigra (a motor-associated nucleus that projects back to
the basal nuclei), and the periaqueductal reticular formation.
The crus cerebri on the underside of the tegmentum are longitudinal fascicles
connecting the cerebrum and pons.
METENCEPHALON
Here the central canal „peels open‟ to form the 4th ventricles, enclose by the paper thin
velum and overlying cerebellum.
Cerebellum
A compact mass with a tree-like internal architecture (folia) creating many small but
deep fissures on the surface. It sits suspended over the roof of the 4th ventricle, bridging
across cerebellar peduncles. It has two hemispheres connected by a mid-line vermis.
Pons
The externally visible pons is a transverse enlargement [L. = bridge] primarily formed by
the pontine nuclei (relays for cerebral motor information to cerebellum). Deeper parts of
the pons are continuous with the medulla (together, the hindbrain), and contain the
„fragmented columns‟ of the nuclei of the cranial nerves, notably the motor nuclei of V
and VII.
MYELENCEPHALON
Medulla
The most primitive part of the brain, the medulla operates primarily at reflex level. It has
many crucial functions:
Homeostatic and autonomic control centres eg. respiratory, cardiovascular,
vasomotor, alimentary „centres‟ (though the so-called „centres‟ in the medulla are
really small ill-defined zones in the medullary reticular formation)
Mediates „vegetative‟ reflex activity eg. salivating, vomiting, swallowing,
coughing, gag reflex, urination
Site of primary nuclei of cranial nerves VII-XII
Has a large part of the reticular formation at its core
Major relay centre and thoroughfare for ascending and descending pathways,
eg.:
o the ascending cuneate and gracile funiculi pass through like-named
nuclei in the dorsal medulla, before dropping to the midline as the
ascending medial lemniscus.
o the olivary nucleus sits just beside the pyramids and relays descending
motor information to the cerebellum
The corticospinal tracts decussate ventrally over the medulla as the pyramids.
Decussation of the auditory nerve creates the transverse trapezoidal body.
Damage to the medulla is typically life-threatening due to the effect on basal vegetative
functions, ie breathing, cardiac function.
Medullary damage can occur accidentally from cisternal puncture, or herniation through
foramen magnum eg secondary to cerebral oedema.
SPINAL PATHWAYS
AFFERENTS
The general pattern of afferent pathways involves a chain of three neurons:
1st neuronal body is in the dorsal root ganglion (spinal or cranial)
2nd neuron as a rule decussates (except for some special senses)
3rd neuron generally has a cell body in the thalamus projecting to the cortex,
typically in defined point-to-point fashion.
Most afferents (except proprioception) also connect to the very primitive ascending
reticular formation.
Conscious afferents
These detect touch, pressure, joint movement, proprioception, pain, heat/cold.
All project onto the primary somatic sensory area of the cortex.
Afferent information is crucial to normal posture, and coordination of effective movement.
Oliver Lorenz & Kornegay Handbook of Veterinary Neurology 3rd ed. WB Saunders 1997, p8
Unconscious afferents
There are two substantial ascending afferent systems that remain beneath conscious
perception:
1. Spinocerebellar system
These are proprioceptive neurons that project directly onto the cerebellum and
never reach the cortex, so remain completely unconscious. Provide feedback
about musculoskeletal activity, allowing the cerebellum to adjust accordingly.
Reticular neurons sit around central canal and its derivatives (eg mesencephalic
aqueduct) and its axons form the indistinct spinoreticular tract deep in the
lateral funiculus. It continues through all parts of the brainstem as far as
thalamus; from here its projection into the cortex is very diffuse and indistinct
(subconscious).
Functions:
- arousal of cortex (hence “Reticular Activating System”, RAS) to alert it
to engage its more specific sensory systems („awareness‟). Inhibition of
the RAS results in sleep or coma (hence a role in general anaesthesia)
- non-specific deep or severe pain transmission (hence a role in analgesia).
SOMATIC EFFERENTS
Initiated from the primary somatic motor cortex, which lies adjacent to the somatic
sensory and has a similar somatotopic arrangement. These are long axons extending
from the motor cortex all the way to spinal LMNs. The bulk of axons decussate in the
pyramids to become the lateral corticospinal tract. An ipsilateral ventral
corticospinal tract (which in any case decussates just before its destination) is minor in
primates and in negligible in domestic species.
Damage in to the corticospinal tract in dogs has only mild effects on motor function
(mild ataxia or contralateral paresis).
The equivalent system for head muscles (via cranial nerves) is termed the
corticobulbar tract. It too is mostly contralateral.
2. Extrapyramidal system
This is a diverse but interconnected system of motor neurons in nuclei scattered
throughout all levels of the brain – a multisynaptic pathway.
Its effect on LMNs is mixed inhibitory-facilitatory, with the net effect (particularly from the
cortex itself) typically being mostly inhibitory.
Therefore disease of the extrapyramidal system mainly causes symptoms of hypertonus
and hyperreflexia.
The caudate nucleus lies in the rostral floor of the lateral ventricles; it is separated from
the related lentiform nucleus by the internal capsule (a curious name for the fibre
radiation from the thalamus into the cortex).
The globus pallidus (=pallidum) is a converging point for all basal nuclei, effectively the
top of the descending reticular formation.
Most basal ganglia probably exert effect through thalamus; their main role has been
described as collaboration with cortex via thalamus.
The extrapyramidal system has two descending pathways, both decussing in the
hindbrain. Most extrapyramidal efferent tracts lie around the tip of ventral horn.
1. Red nucleus rubrospinal tract
Thought to be involved in semi-skilled motor movement, though damage is
typically subtle and difficult to detect.
The vestibulospinal tract is concerned with reflex postural (extensor) tone rather than
voluntary movement.
The vestibulospinal tract originates in the vestibular nucleus and is strongly facilitatory of
ipsilateral extensors, but is normally dampened by cerebellar inhibition.
Cerebellar damage releases uninhibited activity of this system, therefore also causes
marked opisthotonus in this case termed decerebellate rigidity.
The tectospinal tract is a minor motor pathway for neck muscles, initiated from the
tectum (ie. rostral and caudal colliculi). It is involved only in unconscious reflex activity,
ie. reflexively turning the head towards sudden auditory or visual stimuli.
Feedback on intended movement is relayed from the brainstem to the cerebellum via the
pontine nuclei (from primary somatic motor cortex) and olivary nucleus (from red
nucleus, reticular formation).
Primar y
motor Cortex
Basal nuclei:
Caudate nucleus
Globus pallidus
Thalamus
FORE
Descending
Tectum (vision)
Formation
Reticular
Red nucleus
de
cus
s MID
PY dec
RA
MI uss Pontine RF
Vestibular DS Olivar y nucleus
nucleus Pontine decu Cerebellum
nuclei
ss Medullar y RF
HIND
de
cus
Reticulospinal tracts
Vestibulospinal tract
Spinocerebellar tracts
s
Corticospinal tract
Rubrospinal tract
+ +
+ - + - - Extensor activation
Gait generation
Lower motor neurons or alpha skeletomotor neurons sit in the ventral horn (column) of
the spinal cord. LMNs are typically set to „auto-correct‟ small changes in muscle tension
(resulting from changes to joint angle), creating the reflex arcs tested clinically.
Rate of firing of LMNs (which are always excitatory for skeletal muscle) determines the
power of contraction of the supplied motor unit. Firing of LMNs is modified by summation
of inhibitory and excitatory effects from UMNs.
Damage to LMNs always results in flaccid paralysis, loss of reflexes, and rapid muscle
wasting.
The extrapyramidal pathways act mostly through fusimotor gamma neurons, thus
recruits spinal reflexes to effect voluntary movement.
UMN
Me
Po
dul
nt
Rubrospinal
ine
Vestibulospinal
lary
Re
tic
Ret
ulo
icul
sp
ina
osp
l
ina
l
‘Central Pattern Generator’ Interneurons
LMN
MUSCLE CONTRACTION
THE ‘GENERAL’ CRANIAL NERVES
NB. Cranial nerves I (olfactory), II (optic) , III, IV, VI (motor to eye muscles), and VIII
(vestibulocochlear) are described with Special Senses.
V - Trigeminal:
A composite somatic nerve:
Somatic sensory to the eye, facial skin and deeper parts of the face
Somatic motor to the muscles of mastication.
The trigeminal has three major branches loosely corresponding to the embryonic
processes forming the face. Each has a defined area of autonomous cutaneous
innervation: between the eyes (ophthalmic), muzzle (maxillary), and chin
(mandibular).
Nerve bodies of the afferent neurons form the large trigeminal ganglion.
The trigeminal motor neurons are in the pons, while sensory afferents track caudally to
the medulla.
Medullary lesions can therefore potentially cause sensory loss without masticatory
muscle involvement.
Deficits of the ophthalmic branch will affect the sensory component of the corneal and
palpebral reflexes. This differentiates trigeminal lesions from those of the
somatosensory cortex.
Unilateral motor lesions are difficult to detect initially until they become obvious as the
masticatory muscles atrophy.
VII - Facial
A mixed nerve, mostly due to what is termed the intermediate nerve („intermediofacial’):
Somatic motor to muscles of facial expression
Parasympathetic motor to salivary glands, lacrimal glands
Special sensory to mucous membranes, taste
Somatic sensory to skin & hairs inside ear
These two components part ways within the petrous temporal bone, thus the part exiting
the skull from the stylomastoid foramen is the facial nerve proper.
Smaller peripheral branches often combine with the trigeminal (V) to form mixed somatic
sensory/motor nerves.
The facial nerve is colosely associated with the vestibulocochlear (VIII) in both the
brainstem and the petrous temporal bone, which they enter through the internal
acoustic meatus within the same meningeal sheath.
Lesions of the facial nerve cause ipsilateral facial paralysis, +/- parasympathetic motor
loss (ie. dry lacrimal and salivary glands).
Facial paralysis typically causes drooping lip, deviated nose, or dropped ear. Food may
collect in the oral vestibule (lips).
Facial nerve damage is often associated with vestibular signs due to their close
association both centrally and peripherally – which must be differentiated due to the
different prognosis (ie for central vs peripheral vestibular disease).
Part of the facial nerve lies exposed within the middle ear thus are often affected by
otitis media.
IX – Glossopharyngeal:
A mixed nerve:
Somatic motor to some pharyngeal muscles.
Parasymathetic motor to parotid, zygomatic salivary glands
Sensory to mucosa of caudal 1/3rd of tongue (including taste buds), palate,
pharynx
Sensory to carotid body baroreceptor.
Nerves to the pharynx intermingle with those of the vagus via the pharyngeal plexus.
The glossopharyngeal contribution is mostly sensory, with the motor component mostly
but probably not entirely limited to the stylopharyngeus.
The nerve runs through the guttural pouch in horses and can be affected by infections of
the pouch.
Damage to the glossopharyngeal normally manifests as difficulty swallowing (dysphagia)
and reduced gag reflex.
X – Vagus:
[L. = ‘wandering’ nerve]
The vagus exits the skull through jugular foramen, with the glossopharyngeal.
The recurrent laryngeal nerves become „trapped‟ by the 4th branchial arteries which
become the right subclavian and (left) aortic arch, explaining its strange path and length.
The left recurrent laryngeal is therefore most prone to damage from stretching,
especially in horses – causing laryngeal hemiplegia (‘roaring’)
XI – Accessory:
Not really a single nerve but a brief amalgamation (emerging via foramen lacerum) of a
spinal root bundle which heads rostrally to meet a cranial root, which quickly joins the
vagus of which it is really part.
CN IX, X and XI emerge from same medullary nucleus, the nucleus ambiguus.
Dysphagia most common sign of cranial nucleus ambiguous lesion (glossopharyngeal,
vagus).
Laryngeal paralysis is the most common sign of a caudal nucleus ambiguous lesion
(vagus, accessory).
XII – Hypoglossal:
Arises from ventral roots of the caudal medulla and emerges via the small hypoglossal
canal. It is:
somatic motor to the intrinsic and extrinsic muscles of the tongue.
Damage causes paresis/paralysis of tongue, causing difficulties prehending food. The
tongue is deviated and eventually atrophies on side of lesion.
PERIPHERAL AUTONOMIC NERVOUS SYSTEM
The autonomic nervous system operates under the higher influence of the hypothalamus
and medulla, mostly projecting downwards via the reticular formation and reticulospinal
tracts. A ganglion is always interposed between the spinal „LMN‟ and the target organ.
Damage to the reticulospinal tracts can cause classic symptoms of autonomic
disturbance, eg. in man - vasodilation, anhydrosis;
A horse with a broken spine sweats behind the lesion due to apocrine sweat glands.
Parasympathetic
Pre-ganglionic neurons are located in the brain and lumbosacral intumescence.
Parasympathetic oculomotor (CN III) nucleus in the midbrain, aka Edinger-
Westphal nucleus – constricts pupil.
Parasympathetic nuclei of medulla – to CNs VII, IX-XI.
Sacral outflow to pelvic nerves (above rectum) to innervate pelvic viscera, eg.
bladder.
Post-ganglionic fibres typically originate from tiny plexuses in or directly adjacent to the
target organs, eg. cardiac, gastric, hepatic, mesenteric plexus, where they join incoming
sympathetic nerves.
Sympathetic
Pre-ganglionic are in the thoracolumbar spinal cord; post-ganglionic fibres originate from
either (1) ganglia of the sympathetic chain or its extension up the neck:
or (2) some pre-ganglionic fibres bypass the sympathetic chain ganglia and continue as
the splanchnic nerves to major abdominal ganglia:
Coeliac ganglia and cranial mesenteric ganglion at base of like-named
arteries – supply most abdominal viscera (via gastric, splenic, renal plexus etc.)
Caudal mesenteric ganglion – forms hypogastric nerves to pelvic viscera
including bladder (forming the pelvic plexus with parasympathetic pelvic nerves)
Control of micturition
Innervation of bladder is placed far caudally, ie S1-3 in dog & cat. Sensory (stretch)
neurons and parasympathetic outflow ( detrusor muscle contraction) are in the pelvic
nerve; sympathetic (contraction of neck & sphincter) is L1-L4 via hypogastric nerve.
The urethra is striated muscle under somatic control (pudendal nerve).
Spinal reflexes can effect emptying, therefore higher-level lesions cause loss of
voluntary control but not always complete, flaccid bladder distension. However lesions
of the lumbosacral segment itself interrupts reflex emptying and causes severe
distension and overflow dribbling.
Robinson & Huxtable Clinicopathologic Principles for Veterinary Medicine Cambridge UP 1988
Horner’s syndrome:
A distinctive set of facial autonomic symptoms indicating disruption of the cervical
sympathetic pathway:
constriction of pupil (miosis)
drooping of the eyelid (ptosis)
protrusion of 3rd eyelid, secondary to retraction of the eyeball (enophthalmos)
facial vasodilation/hyperaemia (eg. ear pinna, nasal mucosa); sweating in horses
Lesions can be anywhere along the pathway from the midbrain to the eye:
- brachial plexus (eg. brachial plexus avulsion)
- cranial mediastinum (eg. neoplasm - thymoma)
- vagosympathetic trunk (eg. needle laceration during misguided jugular injection)
- petrous temporal bone (eg. otitis media/interna)
- retrobulbar region (eg. haematoma)
Horner’s syndrome accompanying brachial plexus trauma indicates very proximal (nerve
root) trauma thus a poor prognosis.
THE SPECIAL SENSES – NEURAL PATHWAYS
OLFACTORY PATHWAY
From specialized olfactory epithelium high in back of nasal passages (ethmoidal
conchae) and the vomeronasal organ (pheromone receptor).
1st neuron is the neuroepithelial cell; these pass bundled axons through the cribriform
plate of the ethmoid.
2nd neuron is in the olfactory bulb, axons form olfactory tract to olfactory tubercle.
3rd neuron is in the olfactory tubercle and projects directly onto the olfactory cortex
(piriform lobe) without thalamic relay (indicating a phylogenetically ancient pathway).
VISUAL PATHWAYS
Rods & cones Bipolar cells (1st neuron) ganglion cell of retina (2nd neuron) optic
disc (via lamina cribrosa) optic nerve (CN II) via optic foramen optic chiasm
optic tracts (1) lateral geniculate body (3rd neuron) primary visual cortex
(occipital lobe)
(2) pretectal nucleus ( PLR)
and
(3) rostral collicullus of tectum ( visual reflexes & coordination of
eye movement)
Decussation at the optic chiasm varies between species; almost complete in species
with laterally facing eyes, since visual fields don‟t overlap. (NB. Decussation is the
standard, reduced decussation is an advanced characteristic of carnivores and
particularly primates).
The rostral colliculus coordinates eye movement, and mediates visual reflexes,
projecting:
bilaterally to eye muscles (ie. motor nuclei CN III, IV, VI) to reflexively move both
eyes towards stimulus (eg. movement or a flash of light)
contralaterally to tectospinal tract, to reflexively move head and neck towards
stimulus.
The adjacent pretectal nucleus is the relay for the pupillary light reflex. It projects
bilaterally to the Edinger-Westphal nucleus (=parasympathetic nucleus of the
oculomotor n.) to constrict pupils consensually.
The menace response is not a tectal reflex but is instead a primitive learned response
involving the contralateral visual & motor cortex, plus the ipsilateral pontine nuclei and
cerebellum.
IV – Trochlear:
Unusual, small nerves emerging dorsally over rostral medullary velum from midbrain.
Somatic motor to dorsal oblique muscle (which hooks through a trochlea, hence
the name)
VI – Abducens:
Nucleus in caudal brainstem.
Somatic motor to lateral rectus and retractor bulbi muscles.
BALANCE
There are 5 inputs to the vestibular ganglion (1st neuron):
cristae of the ampulla of each semicircular duct (dynamic movement of head)
macula of the utricle (static position of head relative to ground)
macula of the saccule (static position of head relative to vertical)
vestibular nuclei (2nd neuron)
1. via medial longitudinal fasculus medial geniculate body (3rd neuron) cortex.
The cortex receives balance information but does control balance – this is the job
of the hindbrain.
2. ipsilateral vestibulospinal tract
3. ipsilateral cerebellar cortex.
The cerebellum provides feedback to vestibular nuclei, strongly influencing their
activity particularly the vestibulospinal tract. Cerebellar influence is greatest
during movement rather than when still, ie. effects of cerebellar disease are seen
when trying to move.
4. reticular formation – this is source of visceral responses such as vomiting.
Variations to these typical clinical signs that might indicate central (brainstem,
cerebellum) rather than peripheral (middle or inner ear) disease include:
- Vertical or variable nystagmus
- Marked ipsilateral hypertonus
ear pinna
tragus External vs middle vs inner „ear‟.
external auditory canal, acoustic meatus Vestibular vs cochlear components of
auricular / annular / scutular cartilage inner ear & vestibulocochlear nerve.
cerumen Cartilage vs bony portions of ear canal.
tympanic membrane Semi-sealed, air-filled, mucous-
auditory (Eustachian) tube membrane-lined nature of middle ear.
auditory ossicles: incus, malleus, stapes Anatomical principles of sound
tympanic bulla (vibration) amplification.
vestibule; oval window, round window Passage of adjacent nerves through
the middle ear, eg facial n., chorda
utricle, saccule, macula tympani
cochlea; cochlear duct Bony vs membranous labyrinth.
scala vestibuli / tympani Perilymph vs endolymph.
semicircular canals; ampulla, cupola Fluid dynamics in cochlea, ie. why the
spiral organ of Corti; cilia round window?
endolymph, perilymph Kinetic (ampullae) vs static (maculae)
detection of position & movement.
Vestibular & auditory neural pathways,
conscious vs unconscious.
Typical signs of vestibular disease.