FUNCTIONAL NEUROANATOMY BASIC CONCEPTS

The nervous system consists of the: Central nervous system brain (encephalon) and spinal cord and Peripheral nervous system spinal nerves, cranial nerves, and their ganglia. Note this is a crude, gross anatomical definition only since nerve cells (axons) are continuous across anatomical junction the CNS and PNS. 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 Motor to Skeletal muscle Sensory to Tendons/ligaments, joints (proprioceptive) skin (exteroceptive) VISCERAL Smooth & cardiac muscle; glands Thoracic & abdominal organs; visceral activity; stretch receptors; blood pressure; O2 & CO2 tension; 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: General
Exteroceptive

Spinal nerves, CN V Vision, hearing Muscle & joint proprioception Vestibular - CN VIII Spinal splanchnic nerves; CN VII, IX, X Taste, olfaction Skeletal muscles Smooth & cardiac muscle; glands; autonomic NS

GSA SSA GP SP GVA SVA GSE GVE

Special Somatic General AFFERENT

Proprioceptive

Special General Visceral Special Somatic General General

EFFERENT Visceral

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
GS A GVA
Mixed Afferent & Efferent

DRG

RF GVE GS E

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 GS A
SA S S VA

Afferent

GVA GVE GS E

Efferent

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 Always excitatory; essential for muscle contraction Damage causes: hypotonia / flaccidity weak/absent reflexes severe weakness / paralysis rapid & profound muscle atrophy (neurogenic atrophy) UMN Can be excitatory or inhibitory of LMNs; predominant net effect is usually inhibition of extensors Damage causes: hypertonia / spasticity exaggerated reflexes slight weakness / paresis slow, modest muscle 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.

Autonomic nervous system

Visceral efferent pathways are distinct in always having a ganglion outside the spinal cord, and are further divided into sympathetic and parasympathetic divisions: Sympathetic Physiological response Pre-ganglionic fibres Post-ganglionic fibres Localisation Outflow Fight or flight Short; ganglia close to spine (sympathetic chain) Long; adrenergic More diffuse Thoracolumbar Parasympathetic Relaxed Long; ganglia close to organs Short; cholinergic More discrete Craniosacral (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 GS A GVA GVE GS E Efferent

DRG

S pinal Nerve
ramus communicans

S ympathetic chain 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. The brain is drained: 1. Dorsally via the dorsal sagittal sinus and deeper great cerebral vein (within dural falx cerebri). At the tentorium cerebelli this splits bilaterally into the transverse sinus and descends. The dorsal system drains via the maxillary vein or via the ventral system. 2. Ventrally via the cavernous sinus and dorsal petrosal sinuses which communicate with the IVVP. The cavernous sinus also drains the deep face, upper tooth roots, nasal cavity, and orbit. Infection eg. from a tooth abscess can potentially enter the ventral sinus system and cause intracranial infection. Despite extensive anastomoses, venous occlusion can (rarely) cause local necrosis.

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 spiders 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] The choroid plexuses develop in two regions: 1. the roof of the 3rd ventricle, extending through the interventricular foramina into each lateral ventricle. 2. the caudal roof (velum) of the 4th ventricle, extending through the lateral foramina into the subarachnoid space. 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 Limbic system Rhinencephalon Epithalamus Thalamus Hypothalamus [Retina] Tectum (colliculi) Tegmentum Pons Cerebellum Medulla oblongata Spinal cord Central canal

F O R Prosencephalon E

Telencephalon

Lateral ventricles

Diencephalon

3rd ventricle

M I D H I N D

Mesencephalon

Cerebral aqueduct

Metencephalon Rhombencephalon Myelencephalon Caudal neural tube

4th ventricle

TELENCEPHALON
Associated with: higher processing and association (intelligence) regulation of motor function smell memory; emotions Cortex Basal nuclei Rhinencephalon Limbic system

Phylogenetically, the telencephalon consists of: Pallium Neopallium = neocortex Archipallium = hippocampus & cingulated gyrus Paleopallium = piriform lobes Subpallium Striatum = basal nuclei eg. caudate nucleus, putamen Pallidum = globus pallidus 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.

Primary projection areas of a cat

King AS A Guide to the Physiological and Clinical Anatomy of the Central Nervous System 5th ed. University of Liverpool 1976

Lobe Frontal Parietal Occipital Temporal Piriform

Projection area (approx) Primary motor area Primary sensory area Visual area Auditory area Olfactory area

Function Intricate/learned motor skills (pyramidal system) Prefrontal: behaviour & emotions in man Conscious sensation (pain, touch, conscious proprioception); speech in man Vision; cognition in man Hearing; learning & memory in man (closely associated with hippocampus) Smell

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

S eptal nuclei

For nix

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. Functions of the cerebellum: The main coordinating control box of somatic motor movement, but has no power at all to initiate movement. Motor cortex informs the cerebellum of its intended movements (corticopontine tracts), and the cerebellum regulates to ensure these take place, in two ways: exerts pre-control, ie. compares current state with intended state and modifies intended movement accordingly; exerts feedback control as movements are in progress, to perfect and smooth movement. Retains memory related to motor events. Maintains posture, equilibrium and balance by integration of inputs from spinal afferents (spinocerebellar tracts), vision, vestibular system, and motor input from higher centres With cerebellar lesions there may be no abnormal effects evident when at rest, though disturbed equilibrium may give a broad-based stance. There is typically a tremor coinciding with movement (an intention tremor), and ataxia resembling drunkenness uncoordinated, exaggerated, uneven. Unilateral cerebellar lesions cause ipsilateral symptoms. 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. There are two main conscious afferent systems: 1. Gracile-cuneate system Together termed the lemniscal system. Formed by long, ascending, collateral axons occupying the dorsal funiculus of the spinal cord. The fasciculus gracilis is longer (lumbosacral origin), the fasciculus cuneatus forms lateral to it from the upper trunk upwards. These tracts show precise somatotopy, ie. the spatial arrangement (sacral-lumbarthoracic-cervical) is preserved through to their projection onto the primary sensory cortex (afferents from the cranial nerves, principally V, complete the somatotopic arrangement) The gracile-cuneate tracts transmit conscious sensation (touch/pressure) and kinaesthesia - joint position (proprioception) and movement and are therefore important in locomotion. These are relatively more advanced efferent systems, well developed in domestic mammals. 2. Spinothalamic or extralemniscal system This pathway transmits thermal sensation (heat/cold), and sharp or pricking pain (or more correctly, noxious stimuli pain is a relative perception). This is less well developed in domestic species and may not exist as a specific tract in all species (eg. cat, cow, horse said to be more reliant on the primitive ascending reticular formation). Sharp, pricking pain transmits via fast, myelinated fibres, with accurately localized projection through the thalamus. Aching or deep pain transmits via thin, unmyelinated (ie. slow) fibres. It is poorly localized and partly projects/filters through the ascending reticular formation. Visceral pain enters the system mostly with sympathetic nerves, except for pelvic viscera (with pelvic & pudendal nerves), and thoracic viscera (with vagus).

The small size and deep position of deep pain fibres means these are usually the last to be affected by spinal cord compression eg. intervertebral disc prolapse. Loss of deep pain sensation therefore indicates a poorer prognosis for recovery.

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. Forms both dorsal (ipsilateral) and ventral (contralateral) spinocerebellar tracts. The latter is a contralateral tract but decusses a second time to project ipsilaterally. The dorsal tract is mostly from muscle spindles, ventral tract is more from the tendon (Golgi) organ. Damage to these tracts (on the lateral surface of the spinal cord) causes incoordination and ataxia. 2. Ascending reticular formation This is a very primitive amorphous network running up the core of the CNS, which receives many inputs from all afferents (except proprioception) including all the senses. Although seemingly made obsolete by more modern, specific afferent pathways it still forms more than 50% of neurons in the CNS. 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 Efferent Motor Tracts

These are split into pyramidal and extrapyramidal pathways, primarily based on the situation in humans. 1. Pyramidal system = Corticospinal tract In man this is traditionally responsible for fine/skilled motor movements, though this must not be the case in all species, since the system is minimal in ungulates and ends in the cervical region. This interspecies variation indicates the pyramidal system to be phylogenetically recent. 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 extrapyramidal pathways have two higher sources: 1. The cortex outside the primary somatic motor area, ie. non-specific in origin 2. Basal nuclei (often basal ganglia though they are not ganglia) deep in the telencephalon. Most important are caudate nucleus and lentiform nucleus (=globus pallidus & putamen). 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. 2. Reticular formation Reticulospinal tracts The reticular formation, modulated by the basal ganglia and cortex, projects two efferent tracts from the hindbrain. The pontine reticular formation is more autonomous and stimulates extensor tone; the medullary reticular formation is reliant on higher input and mainly suppresses extensor tone. Severe cortical or midbrain damage therefore releases inhibitory influences on extensor muscles (rubrospinal and medullary reticulospinal tracts), causing the generalized extensor rigidity termed opisthotonus or decerebrate rigidity.

Reflex/feedback motor systems:

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 Reticular F ormation

Red nucleus
de cus s

Tectum (vision)
MID

Vestibular nucleus
V estibulospinal tract

PY dec RA MI uss DS

Pontine RF
Pontine decu ss nuclei

Olivar y nucleus

Medullar y RF
de cus s

Cerebellum
HIND

Reticulospinal tracts

+ +

+ -

Extensor activation

Ipsilateral LMNs

Contralateral LMNs

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. Local interneurons (central pattern generators) are capable of generating stereotypic/robotic motor movements, explaining the spinal walking movements sometimes seen even after complete damage of the spinal cord above.

S pinocerebellar tracts

Corticospinal tract

Rubrospinal tract

Initiation of forward movement by UMNs can be conceived as alternating from a postural phase (anti-gravity extensor tone dominant), to a limb-swinging protraction phase (flexor tone dominant). This explains why the predominant effect of UMNs is inhibitory of extensor tone.

Postural phase
+ Anti-gravity extensors -F lexors

Protraction phase
+ Flexors - Extensors

UMN

V estibulospinal

Po nt ine Re tic ulo sp ina l

l ina osp icul Ret lary dul Me

Rubrospinal

Central P attern Generator Interneurons

LMN

S pinal reex arcs

alpha neurons
(muscle spindle)

gamma neurons
(fusimotor)

MUS CLE 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. Ophthalmic
(via orbital fissure)

Lacrimal Frontal Nasociliary

Maxillary
(via round foramen)

Mandibular
(via oval foramen)

Zygomatic Lesser palatine Greater palatine Nasal Infraorbital (muscular branches) Buccal Auriculotemporal Lingual Inferior alveolar

Lateral canthus (angle) of eye Upper eyelid; forehead; frontal sinus Medial angle (infratrochlear n.); eye and cornea (ciliary n.); septal and dorsal nasal mucosa (ethmoidal n.) Lower eyelid; horn cattle (corneal n.) Soft palate Hard palate Ventral nasal mucosa, sinuses Teeth; muzzle, nose Motor to masticatory muscles: masseter, temporalis, pterygoids. Cheek Rostral parts of external ear; skin of temporal region Oropharynx; tongue mucosa (with facial n.) Motor to mylohyoid, cranial digastric; sensory to lower teeth; lower lip and chin (mental n.)

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. Intermediate [Autonomic component] - via geniculate ganglion Major petrosal n. Motor: Nasal, lacrimal glands Sensory: Palate taste buds Motor: Mandibular & lingual salivary glands Sensory: rostral 2/3rds of Tongue (with CN V) Sensory to caudal skin inside ear

Chorda tympani

Facial [proper]

Internal auricular nn. (caudal & middle) Auriculopalpebral n. Dorsal buccal br. Ventral buccal br.

Motor to upper eyelid, ear Motor to cheek, lips, nose

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). The motor component of the palpebral/corneal reflexes may be absent or reduced. This, combined with reduced lacrimal secretion, may cause exposure keratitis (dry eye). 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] A complex, mixed nerve combining the sensory component of caudal pharyngeal arches, plus the parasympathethic outflow to the cranial half of the body. Somatic motor to larynx and pharynx Parasymathetic motor to thoracic and abdominal viscera Sensory to mucosa of larynx & pharynx Sensory to external ear canal. The vagus exits the skull through jugular foramen, with the glossopharyngeal.
[Somatic component]

Auricular n. Pharyngeal n.

Sensory to skin of ear canal Motor to pharynx & cranial oesophagus (with glossopharyngeal) Motor to cricothyroid Sensory to larynx Motor to other laryngeal mm. & caudal oesophagus Parasympathetic to heart, lungs, cranial abdominal viscera

Cranial laryngeal n.

Recurrent laryngeal n.

[Visceral component]

Dorsal branch Ventral branch

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. The spinal accessory nerve is: somatic motor to the superficial neck muscles: trapezius, sternocephalic, brachiocephalic and omotransverse. 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 EdingerWestphal 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: Cervicothoracic ganglion is the enlarged, most rostral ganglion in the sympathetic chain (under head of 1st rib) Middle cervical ganglion at thoracic inlet (to heart, lungs, etc); sympathetic nerves from cervicothoracic split to entrap subclavian artery in ansa subclavia. Cranial cervical ganglion lies directly adjacent to distal vagus ganglion. Sources of efferents to all head/neck viscera eg eye (dilation of pupil), lacrimal & salivary glands, sweat glands, carotid sinus

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

Horners 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) Horners 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 dont 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.

EYE-ASSOCIATED CRANIAL NERVES

III Oculomotor: Nucleus in tectum of midbrain. Somatic motor to most eye muscles Visceral efferent parasympathetic (Edinger-Westphal nucleus) to constrictor of pupil 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.

AUDITORY / VESTIBULAR PATHWAYS

HEARING Neuroepithelial cells in organ of Corti cochlear ganglion (1st neuron) cochlear nucleus (2nd neuron) via lateral lemniscus (1) medial geniculate body (3rd neuron) primary auditory cortex (temporal lobe) and (2) caudal collicullus of tectum ( reflex pathways, eg. reflexive turning of head towards a sudden sound). 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. Signs of unilateral peripheral vestibular disease include: - Nystagmus (rhythmic flicking movement of the eyes) horizontal or rotatory (never vertical), fast phase away from side of lesion - head tilt (towards lesion) - circling or rolling (towards side of lesion) - ataxia without weakness; compensatory broad-based stance - may be contralateral hypertonus from involvement of vestibulospinal tract. 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

GUSTATORY (TASTE) PATHWAYS NB Remember most of the sensation experienced as taste is actually smell.

Sensory afferent pathways from the taste buds involve several cranial nerves: rostral 2/3rds of the tongue travels with the lingual nerve (branch of maxillary V) but then splits off as the chorda tympani of the facial (VII) geniculate ganglion medulla caudal 2/3rds of the tongue via glossopharyngeal n. (IX) epiglottis and area of larynx via vagus n (X). Taste sensation is projected to the primary sensory cortex (via thalamic relay). Local brainstem relays cause autonomic reflexes eg. salivation.

EYE & EAR ANATOMY - CHECKLIST

KEY WORDS orbit, orbital ligament eyelids / palpebrae canthus / commissures cilia tarsus tarsal / Meibomian glands conjunctiva 3rd eyelid / nictitating membrane nictitans gland, lacrimal gland lacrimal puncta / ducts / sac nasolacrimal duct sclera, lamina cribrosa cornea, limbus vascular tunic / uveal tract iris ciliary body / processes / zonules choroid, tapetum lucidum fundus, macula, fovea retina rods, cones lens KEY CONCEPTS
Form of the bony orbit and its foramina. Position of the eye and relationship to surrounding peri-orbital & retro-orbital structures. The function and integumentary specializations of the eyelids. The extent of the conjunctival sac. Lacrimal apparatus function, production & drainage of tears. The nested, laminar structure of the fibrous, vascular, and nervous tunics of the bulbus oculi. Production & flow of aqueous / vitreous humour. Movement of eyeball, extra-ocular muscles & their innervation. Intraocular examination: fundus, macula, optic disc, retinal vessels. The upside-down arrangement of retinal layers. Refraction of light by cornea and lens; accommodation by ciliary muscle. Neural pathways for vision; corneal & palpebral reflexes; menace response.

ear pinna tragus external auditory canal, acoustic meatus auricular / annular / scutular cartilage cerumen tympanic membrane auditory (Eustachian) tube auditory ossicles: incus, malleus, stapes tympanic bulla vestibule; oval window, round window utricle, saccule, macula cochlea; cochlear duct scala vestibuli / tympani semicircular canals; ampulla, cupola spiral organ of Corti; cilia endolymph, perilymph

External vs middle vs inner ear. Vestibular vs cochlear components of inner ear & vestibulocochlear nerve. Cartilage vs bony portions of ear canal. Semi-sealed, air-filled, mucousmembrane-lined nature of middle ear. Anatomical principles of sound (vibration) amplification. Passage of adjacent nerves through the middle ear, eg facial n., chorda tympani Bony vs membranous labyrinth. Perilymph vs endolymph. Fluid dynamics in cochlea, ie. why the round window? Kinetic (ampullae) vs static (maculae) detection of position & movement. Vestibular & auditory neural pathways, conscious vs unconscious. Typical signs of vestibular disease.