ANP 1106 A

7. Neurophysiology
ANP 1106 A – Anatomy & Physiology II

TABLE OF CONTENTS

Organisation of Nervous System ...................................................................................................................................5

Neuron ...........................................................................................................................................................................5
Ionotropic Receptor ..............................................................................................................................................5
Neurotransmitter Structure ..................................................................................................................................6
Neurotransmitter Action .......................................................................................................................................7
Temporal and Spatial Summation .........................................................................................................................8
Circuits...................................................................................................................................................................8
Sensory Receptors .........................................................................................................................................................9
Classification .........................................................................................................................................................9
Example ................................................................................................................... 9
Transduction .......................................................................................................................................................10
Intensity Coding ..................................................................................................................................................10
Adaptation Examples ..........................................................................................................................................11
Adaptation Summary ..........................................................................................................................................11
Sensory Systems ..........................................................................................................................................................12
1. Taste ........................................................................................................................................................................12
Organ ...................................................................................................................................................................12
Receptors ............................................................................................................................................................13
Sensory Pathway .................................................................................................................................................13
2. Smell ........................................................................................................................................................................14
Organ ...................................................................................................................................................................14
Epithelium ...........................................................................................................................................................14
Receptors ............................................................................................................................................................14
Sensory Pathway .................................................................................................................................................15
3. Hearing & Balance ...................................................................................................................................................15
Structures ............................................................................................................................................................15
Labyrinth .............................................................................................................................................................17
Fluids ...................................................................................................................................................................17
Hearing ................................................................................................................................................................18
Sound Waves ..............................................................................................................................................18
Wave Amplitude and Frequency ................................................................................................................18
Mechanical Stimulus ...................................................................................................................................19
Sensory Organ.............................................................................................................................................19
Transduction ...............................................................................................................................................20
Sensory Pathway.........................................................................................................................................21
Cortical Tonotopy .......................................................................................................................................21
Sound Localisation ......................................................................................................................................22
Balance ................................................................................................................................................................23
Static Balance..............................................................................................................................................23

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Organ .................................................................................................................................. 23
Transduction ....................................................................................................................... 23
Receptor Orientation.......................................................................................................... 24
Dynamic Balance.........................................................................................................................................24
Semicircular Channels ........................................................................................................ 24
Organ .................................................................................................................................. 24
Transduction ....................................................................................................................... 25
VOR (Vestibulo-Occular Reflex) .......................................................................................... 25
Sensory Pathway.........................................................................................................................................25
Hearing & Balance: Similar Transduction ............................................................................................................26
4. Vision .......................................................................................................................................................................27
Eye Lubrification ..................................................................................................................................................27
Eye Muscle ..........................................................................................................................................................27
Saccades and Smooth Pursuit .............................................................................................................................28
Convergence .......................................................................................................................................................28
Light.....................................................................................................................................................................29
Reflection ............................................................................................................................................................29
Absorbed and Reflected Light .............................................................................................................................29
Light Path ............................................................................................................................................................29
Pupil Size .............................................................................................................................................................32
Pupillary Light Reflex ...........................................................................................................................................32
Focus and Inversion.............................................................................................................................................33
Accommodation ..................................................................................................................................................33
Myopic and Hypermetropic ................................................................................................................................34
Presbyopia ...........................................................................................................................................................35
Cataracts .............................................................................................................................................................35
Photoreceptors ...................................................................................................................................................35
Cone Sensitivities ................................................................................................................................................35
Red-Green Perception Test .................................................................................................................................36
Rhodopsin Cycle ..................................................................................................................................................36
Transduction .......................................................................................................................................................37
Photoreceptor Activation ....................................................................................................................................37
Visual Fields .........................................................................................................................................................38
Synaptic Relays ....................................................................................................................................................38
Connections to Cortex .........................................................................................................................................39
Secondary Connections .......................................................................................................................................40
5. Somesthesia .............................................................................................................................................................41
Simple Receptors ................................................................................................................................................41
Pain Stimulation ..................................................................................................................................................41
Pain Transduction................................................................................................................................................42
Temperature Stimulation ....................................................................................................................................42
Temperature Transduction .................................................................................................................................42
Mechanical Stimulation.......................................................................................................................................43
Mechanical Stimulus Transduction .....................................................................................................................43

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Proprioceptors ....................................................................................................................................................43
Proprioceptor Function .......................................................................................................................................44
Sensory Pathways ...............................................................................................................................................44
Somatosensory Cortex ........................................................................................................................................46
Sensory Homunculus ...........................................................................................................................................46
Internal Organ Pain .............................................................................................................................................46
Referred Pain ......................................................................................................................................................46
Perception ...................................................................................................................................................................47
Three (3) Levels ...................................................................................................................................................47
Cortical Connections ...........................................................................................................................................47
Detection and Magnitude ...................................................................................................................................48
Spatial Discrimination .........................................................................................................................................48
Feature Abstraction ............................................................................................................................................48
Pattern Recognition ............................................................................................................................................49
Retinal Slip ...........................................................................................................................................................49
Center-Surround Cortical Responses ..................................................................................................................49
Simple Cortical Responses ..................................................................................................................................51
Complex Cortical Responses ...............................................................................................................................51
Sensory Filtering ..................................................................................................................................................52
Motor Systems.............................................................................................................................................................52
Sensorimotor Arc ................................................................................................................................................52
Motor Homunculus .............................................................................................................................................53
Primary Descending Tract ...................................................................................................................................53
Secondary Descending Tracts .............................................................................................................................53
Cerebellar And Striatal Loops ..............................................................................................................................54
Cerebellar Disease ...............................................................................................................................................55
Basal Ganglia Disease ..........................................................................................................................................55
Spinal Pattern Generator ....................................................................................................................................56
Muscles: Neuromuscular Junction ......................................................................................................................56
Neural Integration: sensorimotor arc .................................................................................................................57
Reflexes .......................................................................................................................................................................57
Somatic Reflexes .................................................................................................................................................57
Basic Reflex Arc ...........................................................................................................................................57
Withdrawal Reflex ......................................................................................................................................58
Tendon Reflex .............................................................................................................................................58
Stretch Reflex ..............................................................................................................................................58
Plantar Reflex ..............................................................................................................................................59
Autonomic Reflexes ............................................................................................................................................59
Organisation ...............................................................................................................................................59
Homeostasis ...............................................................................................................................................60
Example: Temperature Control .......................................................................................... 60
Sympathetic And Parasympathetic .............................................................................................................60
Effects .........................................................................................................................................................60
Neurotransmitters ......................................................................................................................................61

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Receptors ....................................................................................................................................................63
Volition ........................................................................................................................................................................64
Cortex Roles ........................................................................................................................................................64
Spoken Language ................................................................................................................................................65
Written Language ................................................................................................................................................66
Braille Language ..................................................................................................................................................66
Supporting Systems .............................................................................................................................................67
1. Memory ..................................................................................................................................................67
Working Memory Circuit ....................................................................................... 67
Memory Formation ............................................................................................... 68
Memory Consolidation .......................................................................................... 68
2. State of Mind ..........................................................................................................................................69
1. Reticular Activating System (RAS) & Activation ............................................................. 69
Reticular Activating System (RAS) ......................................................................... 69
Seizures.................................................................................................................. 69
2. Sleep & Wakefulness ...................................................................................................... 70
Sleep Stages ........................................................................................................... 70
Sleep Stages Overnight .......................................................................................... 71
Sleep Changes With Age ........................................................................................ 72
3. Aminergic Projections & Mood ...................................................................................... 72
Biogenic Amines .................................................................................................... 72
............................................................................................................................... 73
Mood Disorders ..................................................................................................... 73
4. Limbic System & Emotions ............................................................................................. 73
Emotions ................................................................................................................ 73
Limbic System Links Other Systems ...................................................................... 74
Our World is in the Cerebral Cortex ............................................................................................................................75

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ORGANISATION OF NERVOUS SYSTEM

• The nervous system is subdivided
1. Central Nervous System (CNS)
▪ This is where all the integration of
information occurs
▪ Composed of Brain & Spinal cord
2. Peripheral Nervous System (PNS)
▪ Nerves, formed of Afferent axons that
transmit sensory information towards the
CNS.
▪ Nerves, formed of axons that transmit
Efferent information to the:
1. Somatic Nervous System
o Control of skeletal
muscles and voluntary
movement
2. Autonomic Nervous System
o Involuntary; control of organs, smooth, & cardiac muscle
1. Sympathetic System
2. Parasympathetic System

NEURON

IONOTROPIC RECEPTOR
• Neurons
o Functional units of the nervous system
• Chemical Synaptic Transmission between neurons:
a. Once in the Presynaptic Terminal, the action potential triggers the opening of Ca++ Voltage-
Gated Channels
b. This allows entry of calcium to serve as a Second Messenger for the release of neurotransmitters
into the Synaptic Cleft
▪ There are many types of neurotransmitters:
• Peptides
• Amino acids
• Biogenic amines
• Acetylcholine
o The neurotransmitter at the neuromuscular junction

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▪ The most common neurotransmitter in the CNS are:

• Glutamate (excitatory)
• GABA (inhibitory)
c. The neurotransmitter in the synaptic cleft binds to a postsynaptic Ionotropic Receptor of a
ligand-gated channel (these channels are only found in the soma and dendrites)
▪ Ionotropic Receptor
• Receptor that opens an ion channel in response to a signal
▪ Ligand-Gated Ion Channels (LGICS)
• One type of ionotropic receptor or channel-linked receptor
• Group of transmembrane ion channels that are opened or closed in response
to the binding of a chemical messenger (i.e., a ligand) such as a
neurotransmitter.
d. This produces a synaptic potential (either Excitatory [as in the figure] or Inhibitory) in the
postsynaptic neuron
e. Termination of the neurotransmitter effects are produced by:
▪ Enzymes in the synaptic cleft that metabolize (cleave) the neurotransmitter
▪ Endocytosis of neurotransmitter by the presynaptic terminal so that it can be reused
again later

NEUROTRANSMITTER STRUCTURE
• Neurotransmitters are classified according to their chemical structure.

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NEUROTRANSMITTER ACTION
• The action of a neurotransmitter depends on its receptor
• Focus on GABA, Glycine, Glutamate, Aspartate, Acetylcholine
• There are different types of receptors for each neurotransmitter, and depending on which receptor it
binds to the action of the neurotransmitter will be different
o Some neurotransmitters act on receptors that produce opposite effects
▪ Ex: acetylcholine acts on nicotinic receptors to depolarize skeletal muscle cells
▪ Ex: acetylcholine acts on muscarinic receptors to hyperpolarize cardiac muscle cells
o Some neurotransmitters act on receptors that produce similar but still different effects
▪ Ex. GABA – Ionotropic and Metabotropic receptors, they don’t have the same results
• We don’t need to know the details of this chart; like he said, focus on the notes (not the diagrams)

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TEMPORAL AND SPATIAL SUMMATION
• Graded postsynaptic potentials result from the
summation of successive synaptic potentials which
can exhibit:
o Temporal Summation
▪ e.g. postsynaptic effects of one
terminal activated in rapid
succession
o Spatial Summation
▪ e.g. postsynaptic effects of several
terminals activated simultaneously
• Excitatory Postsynaptic Potential (EPSP)
o Produced if the ligand-gated channel of the
postsynaptic receptor is selective to the flow
of sodium, then sodium will enter and
depolarize or excite the postsynaptic neuron
• Inhibitory Postsynaptic Potential (IPSP)
o Produced if the ligand-gated channel is
selective to the flow of potassium then
potassium will leave and hyperpolarize or
inhibit the postsynaptic neuron producing…
• …an Action Potential, which will occur if the
membrane potential crosses its threshold (-70mV)
o Otherwise, the leak channels will restore the
membrane potential of -70 mV
▪ N.B. Think of -70mV as the lowest energy state of the membrane → membrane is
always trying to return to this potential

CIRCUITS
• The electrical properties of neurons allow communication through the connections forming circuits.
o We know that the phenomenal power of the nervous system derives from the cortical circuits
formed by neurons
o However, we don't know the complete connections of these circuits
• Only simple repeating principles are known. These include connections that:
a. Diverge to spread signals to several neurons
b. Converge to integrate signals from several neurons
c. Reverberate to hold information in memory
d. Produce parallel after-discharges to produce a longer-lasting effect.
• Our study will mostly examine simple serial connections from sensory receptors to the cerebral cortex
(area which contains complex circuits) and from the cerebral cortex to muscles for movement.

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SENSORY RECEPTORS

CLASSIFICATION
• Sensory Receptors → used to detect the state of the internal and external milieu
• 3 methods to classify the sensory receptors
1. According to the anatomical location of the stimulus. This includes:
I. Exteroceptors:
• Located close to the surface of the body
• Receive information of the external environment for: Hearing, Vision, Smell,
Taste, Touch, Pressure, Vibration, Temperature, Pain
II. Visceroceptors
• Located in the blood vessels and the internal organs
• Provide information for automatic control of the internal milieu without
requiring involvement of the conscious areas of the CNS
o *except for nociception which produces referred pain to another part
of the body
III. Proprioceptors
• Located in the muscles and the articulations
• Provide information on the position of the body
2. According to the type of stimulus. This includes
IV. Mechanoreceptors
• Detects mechanical stimuli
V. Thermoreceptors
• Detects thermal stimuli because temperature changes the energy of the
channel molecule which then forms a new stable conformation where channel
opening is altered
VI. Photoreceptors
• Detects visual stimuli or photons
VII. Chemoreceptors
• Detects molecules
VIII. Nociceptors
• Detects dangerous stimuli for the organism
3. According to the complexity of the structure.
These include:
IX. Complex receptors
• Sensory organs of the special
senses (namely, vision, taste,
smell, hearing and balance)
X. Simple receptors
• All the other receptors which
are from the somatic and
vegetative systems
Example
• There are several receptors and each has 3 classifications.
• For example, a Pacinian corpuscle in the skin is at the
same time a:
o Exteroceptor
o Mechanoreceptor
o Simple receptor

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TRANSDUCTION
• Transduction: where the energy of the stimulus is transformed into electric activity
• e.g. transduction of mechanical pressure into depolarization of Pacinian Corpuscles
• 2 types of potentials produced by stimulation of receptors
1. Generator Potential or Receptor Potential – A depolarization produced when the stimulus
produces the opening of ionic channels
2. Action Potential – Produced by depolarization beyond the threshold of excitation

INTENSITY CODING
• The frequency of action potential discharge corresponds to the intensity of the stimulus.

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ADAPTATION EXAMPLES
• Adaptation – The fact that the
initial response of a receptor
decreases after a period of time
• If you give a receptor a
constant stimulus, the
receptor won’t react
the same throughout
the entire duration of
the stimulus
a. Phasic
Receptor
adapts quickly
b. Tonic
Receptor
adapt slowly
• Ex. Meissner’s corpuscle adapts
quickly and on a very large scale,
whereas Merkel cells show very
little adaption. Overall,
however, all receptors show at
least SOME adaptation.

ADAPTATION SUMMARY
• The slowest adaptation is by
nociceptors and proprioceptors
• Reason: They must constantly inform the (CNS) of the stimuli for pain and body position,
respectively

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SENSORY SYSTEMS
• First stage for perception of the environment: transmit sensory information to the cerebral cortex
• This is carried out by the special and somatic senses
• There are complex mechanisms of the special senses which enable us to taste, smell, feel, hear,
maintain balance, and see

1. TASTE

ORGAN
• Surface of the tongue has structures called Papillae which gives it a rough texture
• 2 kinds of papillae
1. Fungiform Papillae
▪ On the soft palate, internal cheeks, pharynx, epiglottis, and predominantly on the end
and sides of the tongue where there are 10000 of them and
2. Circumvallate Papillae
▪ Forms a series of 7-12 towards the back of the tongue
• Papillae contain taste buds each consisting of 40-60 Epithelial Cells forming a taste pore
• 3 types of epithelial cells
1. Supporting Cells
2. Gustatory Cells
▪ Chemoreceptors containing microvilli which adapts to stimuli in 1-5 minutes (they do
not produce an action potential)
3. Basal Cells
▪ Renew the supporting and gustatory cells every 7-10 days
• The receptors for the different fundamental flavors are partially mixed on the tongue, but in general as
we move from the tip → back of the tongue there is a change in the chemical sensitivity from:
• Salty → sweet → sour (acid) → bitter flavors
▪ Umami is found all over the tongue – monosodium glutamate taste
• Ex: By drinking wine we perceive some flavors from the receptors on the tip of the tongue but
then there is a bitter after taste because the wine flows slowly towards the back of the tongue
and stimulates the receptors for bitter flavors.

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RECEPTORS
• The right side of the figure shows 5 types of gustatory cells
• Each cell type has receptors specific for the stimulus to which it responds
• One sensory hair cell has only 1 type of sensory receptor found on all three cilia of the cell
• The mechanisms of transduction
are as follows
A. Salty Flavors
• Sodium (Na+) in food
crosses a Na+ channel to produce
depolarization and release of
neurotransmitter
B. Sour Flavors
• Hydrogen ions (H+) in
food can enter by a type of Na+ channel
(the degree of permeability to Na+ is too
low in the absence of H+ to generate a
depolarization) producing a
depolarization and the release of
neurotransmitter
C. Sweet, Bitter and Umami
Flavors
• Molecules bind to the
metabotropic receptor which activates a
G protein that stimulates the increase in
IP3
• This then causes
release of intracellular Ca++ stores to
trigger neurotransmitter release
• Even though they use the
same mechanism, we don’t mix them on
the same cells or else we wouldn’t be able
to distinguish between them

SENSORY PATHWAY
• Substances in solution stimulate
the chemoreceptors of gustatory cells
which then release neurotransmitters to
activate neurons of one of the following 3
cranial nerves:
• Facial (VII)
• Glossopharyngial (IX)
• Vagus (X)
• These cranial nerves project to the
solitary nucleus in the medulla
• The neurons of the solitary nucleus
project to the thalamus
• This then projects to the insular
cortex for perception of taste

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2. SMELL

ORGAN
• The olfactory organ is in the roof of the nasal cavity.

EPITHELIUM
• The receptor area of the olfactory system is made of an epithelium which contains 3 kinds of cells:
1. Olfactory Cells
▪ These have with olfactory cilia that detect odors
• N.B. The olfactory cells are bipolar neurons forming the first cranial nerve with
axons crossing the cribiform plate of the ethmoid bone to reach the olfactory
bulb
2. Supporting Cells
3. Basal Cells
▪ These regenerate the other cells
▪ Olfactory cells are among the few types of neurons that can be regenerated and it
occurs about every 60 days

RECEPTORS
• Many olfactory receptors; their mechanisms of transduction are incompletely known
• General mechanism:
1. An odor molecule activates a metabotropic receptor
2. This causes opening of a sodium channel (or closing
of a potassium channel) to depolarize the cell

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SENSORY PATHWAY
• Volatile and water-soluble substances stimulate the olfactory cells which project to the Mitral Cells of the
Olfactory Bulb
• These then send projections via the Olfactory Tract to several areas (2 main pathways listed):
1. The projections via the Thalamus to the Orbito-Frontal Cortex (shown in red) provide the ability
to perceive the
quality of odors
2. The other pathways
(shown in blue) give
rise to emotional
responses to odors
▪ This
pathway
doesn’t go
through the
hypothalam
us and thus
we don’t
perceive the
odor; we
just have an
emotional
response to
it.
• Other receptors contribute to
the appreciation of flavors (in addition to taste receptors)
• Sense of smell is responsible for 80% of taste sensations
▪ N.B. it is for this reason that one loses taste with a cold
▪ Only 20% of taste is derived from tongue receptors
• Thermoreceptors give the perception of temperature
• Ex: a hot coffee is preferred
• Mechanoreceptors provide perception of the texture of foods
• Ex: many don't like the fuzz on peaches)
• Nociceptors activate to provide spicy tastes

3. HEARING & BALANCE

STRUCTURES
• The structures for hearing and balance are together. The ear includes 3 parts:
1. Outer Ear
▪ Starts at the Pinna
• The helix forms the thicker cartilaginous edge and the lobule forms the soft
part without cartilage
▪ Passes along the External Auditory Canal
▪ Ends at the Tympanic Membrane
• It is thin and translucent formed of fibrous conjunctive tissue
▪ The external auditory canal contains glands that secrete Cerumen
• Cerumen = a brownish-yellow wax that repels insects
▪ Pressure waves enter the air (ex. Woofer moves in and out creating tranverse waves)

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2. Middle Ear
▪ Forms the Tympanic Cavity
• Contains Ossicles…
o Malleus
o Incus
o Stapes
• …for transmitting movements of the Tympanic Membrane to the Oval Window
• Tympanic cavity open into the Pharyngotympanic Tube that leads to the
Nasopharynx
▪ Pressure in the middle ear
• Ex. When you got on the plane, as you elevate the pressure from outside
pushing on the eardrum decreases, so the air inside the ear pushes outwards
on the eardrum, causing ears to “pop”
• As you land, the pressure starts building up on the outside, pushing inwards on
the tympanic membrane
• Both situations can be fixed by yawning or swallowing
o These evacuate the air in the middle ear via the auditory tube
3. Inner Ear (Labyrinth)
▪ The inner ear contains the sensory receptors.

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LABYRINTH
• The inner ear is in the temporal bone and is formed of a Bony Labyrinth covered with endostium. It
contains three different parts, each of which has its own Membranous Labyrinth:
1. Cochlea (the bony wall)
▪ Cochlear Duct – the membranous labyrinth within the cochlea which transduces sound
2. Vestibule (bone)
▪ Utricule and Saccule – the membranous labyrinth in the vestibule which transduce head
posture
3. Semicircular Canals (bone)
▪ Semicircular Ducts – the membranous labyrinth in the semicircular canals which
transduce head movements.
▪ Oriented on the 3 planes of plane (xyz) detecting movement of the head in any of
direction of motion

FLUIDS
• Endolymph
o Liquid inside the membranous labyrinth
o Similar to the intracellular liquid
o Formed by the Stria Vascularis of the membrane (high K+ concentration-> important for
transduction)
• Perilymph
o Liquid between the walls of the bony and membranous labyrinth
o Similar to extracellular or cerebrospinal fluid
o Derived from a Perilymphatic Duct leading to the subarachnoid space

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HEARING

SOUND WAVES
• Sounds are caused by changes in air pressure.
• Branches of tuning fork vibrate left and right (frequency dependent on rigidity of fork)
• Compresses adjacent area creating high and low pressure as it moves back and forth

WAVE AMPLITUDE AND FREQUENCY

• Sound waves have:
• Amplitude

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▪ Large amplitude wave = loud sound of high intensity
▪ Small amplitude wave = soft sound of low intensity
▪ The intensity is measured in decibels
• 0 dB = threshold of hearing
• 50 dB = normal conversation
• 90 dB = rock concert
• 130 dB = pain
• Frequency
▪ A short duration wave corresponds to a high frequency
sound (high pitch)
▪ A long duration wave corresponds to a low frequency
sound (low pitch)

MECHANICAL STIMULUS
• How the mechanical stimulus of waves is transmitted to a nerve impulse:
a. Sound Waves (variations of air pressure) enter the external auditory canal and apply pressure on the
Tympanic Membrane
b. Movements of this membrane are transmitted to the Malleus, Incus, and Stapes which strikes the
Oval Window
c. Movement of this elastic window enduces waves in the Perilymph propagating through the Scala
Vestibuli, around the Helicotrema, and thence the Scala Tympani ending on the Round Window
where the energy of the waves are absorbed by bulging of its membrane.
d. These waves induce oscillations of the Cochlear Duct and its sensory structure called the Spiral Organ
of Corti which is specialized to detect sound waves.
e. The Basilar Membrane extends from the base of the Cochlear Membrane (near the tympanic cavity)
to the Apex or Helicotrema.
• The highest frequency sounds vibrate the base of the basilar membrane
• Lowest frequency sounds vibrate its apex
• See www.blackwellpublishing.com/matthews/ear.html
• Therefore, dendritic branches of the Cochlear Nerve…
• …at the base of the basilar membrane will transmit information from high frequency waves
• …at the apex will transmit information from low frequency waves.
• The stronger the sound the greater
will be the amplitude of the basilar
membrane vibrations and
consequently the greater the number
of sensory hair cells and cochlear
nerve fibers that will be stimulated.
• This leads to the perception
of a higher intensity sound.
• There are muscles on the first and
last Ossicle (Malleous and Stapes)
which are recruited 40 ms after the
beginning of a strong sound to
reduce the amplitude of the waves.

SENSORY ORGAN

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• The spiral organ rests on the basilar membrane and it is composed of:
• Supporting Cells
• Sensory Hair Cells containing Stereocilia that are attached to the Tectoral Membrane.
▪ The Outer Hair Cells are stimulated by efferent fibers to contract in order to either
boost or dampen sounds.
▪ The Inner Hair cells detect sounds.
• No basal cells
▪ Once either cell is damaged, you lose hearing because there is no regeneration

TRANSDUCTION
• Movement of the Stereocilia cause depolarization of the inner hair cells. Bending of the stereocilia…
i. …in one direction leads to opening of potassium channels and depolarization (bending
toward the Chinocilium - the longest hair in the set)
▪ Bending to the right pulls on cytoskeletal wire, opening the gate on each ear which
is selective for potassium, endolymph containing K+ causes ions to rush in and
depolarize the cell→ transduction)
ii. …in the opposite direction leads to closing of these channels and repolarization
▪ N.B. sensory hair cells are bathed in endolymph which is higher in potassium and
lower in sodium ions than the intracellular fluid of these cell; thus opening of
potassium channels lead to depolarization
• Depolarization of hair cells causes neurotransmitter release which excites dendrites of the Cochlear
Nerve
o N.B. part of the VIII Cranial Nerve also called Vestibulocochlear Nerve

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SENSORY PATHWAY
• Fibers of the Cochlear Nerve join the Vestibulocochlear nerve and then branch off to reach neurons
of the cochlear nucleus.
• These relay information to the Contralateral Superior Olive which relays in the Inferior Colliculus and
the Thalamus which then relays to the Auditory Cortex in the Temporal Lobe.
• There are 2 kinds of deafness:
• Conduction Deafness
▪ Problem with the mechanical transmission of sound waves produces
• N.B. If a patient can only hear the sound from a tuning fork placed on the
temporal bone (vibrations of the tuning fork are transmitted to the hair
cells) then one can conclude that there is conduction deafness
• Sensorineural Deafness
▪ Problem with the transmission of electrical activity produces
• N.B. loud sounds destroy sensory hair cells

CORTICAL TONOTOPY
• The temporal cortex has a Tonotopic Organization
o There is an orderly numerical organization of sound frequencies reaching neighboring areas
▪ Low frequencies are anterior in primary auditory cortex of the temporal lobe
▪ High frequencies are posterior in primary auditory cortex of the temporal lobe

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SOUND LOCALISATION
• Sound is localized by:
• The temporal difference between the arrival of sound in the two ears
• The difference in intensity of sound which enters each ear. Sound coming from the left will
reach the left ear first and be louder in the left ear

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BALANCE

STATIC BALANCE

ORGAN
• The Saccule and Utricule each contain a Macula.
o Macula contains an epithelial membrane,
containing supporting and sensory hair cells,
onto which rests the Otolithic Membrane.
o This Otolithic Membrane contains crystals
called Otoliths whose weight on the
otolithic membrane causes displacements of
the Stereocilia and Kinocilium according to
the orientation of the head

TRANSDUCTION
• The Force of Gravity is the stimulus which acts on the
sensory structure named Macula which transduces
the stimulus into neuronal impulses.
• Bending of the Stereocilia towards the
Kinocilium produces depolarization.
• The position information provided by the
macula is of the head since this structure is
fixed to the head.
• The depolarization of the sensory hair cells
causes the release of neurotransmitter on the Vestibular Fibers of the Vestibulocochlear
Nerve which branches off to the Vestibular Nuclei.

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RECEPTOR ORIENTATION
• Since the orientation of hair
cells differs, only a fraction
of the cells will be
stimulated by a particular
orientation of the head.
o Knowledge of head orientation will derive from the
fraction of hair cells stimulated.

DYNAMIC BALANCE

SEMICIRCULAR CHANNELS
• There are 3 semicircular canals, each directed to detect
movement of the head in one of the 3 spatial dimensions.

ORGAN
• Each canal contains an Ampula which contains the sensory organ named Crista Ampullaris that
transduces acceleration of the head into neuronal impulses.
• Each Crista Ampullaris contains an Epithelial Membrane, formed of supporting and sensory hair
cells, on which lies the Cupula (a gelatinous mass).
o Cupula: hairy membranous part attachd to the sensoyr organ , when it bends, you will bend
the hair cells→ leading to bendign toward kinocilium and channels opening up to allow K+
diffusion into cell→ depolarization.
• Bathes in endolymph (high K+ concentration)
• No basal cells, meaning that you cannot regenerate the supporting cells or hair cells when damaged

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TRANSDUCTION
• Movement 1fof the head
causes movement of the Crista
Ampullaris through the
endolymph in the semicircular
ducts.
• The Cupula bends under force
against the endolymph in the
semicircular ducts which causes
bending of the stereocilia and
kinocilia and evokes
depolarization and transmitter
release onto vestibular fibers of
the vestibulocochlear nerve.

If you spin you force the water

in the cup to bend the hair,
similar to the endolymph bending the crista

VOR (VESTIBULO-OCCULAR REFLEX)

• The fact that one can read even if the bus
strikes a bump is due to The Vestibulo-
Occular Reflex triggered by the crista
ampullaris.
o A brief movement of the head will
produce a brief movement of
endolymph
o But after continuous motion of
the head, as on a merry-go-round,
the endolymph is turning with the
head and is essentially not flowing
around the canals
▪ When the merry-go-
round stops, the head
stops but the endolymph
continues its circular
motion and stimulates
the hair cells which
evoke Nystagmus
(involuntary eye
movements) and
Dizziness

SENSORY PATHWAY
• Fibers of the Vestibular Nerve join the
Vestibulocochlear Nerve which then branch
off to synapse onto cells of the
1. Lateral Vestibular Nuclei
2. Superior Vestibular Nuclei

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▪ This has some projections (bilateral) to the brainstem to participate in reflexes
• e.g. eye movements
▪ And other bilateral projections to the ventral posterior thalamus which thence
projects to the vestibular cortex in the parietal lobe
▪ Raw information about head movement and position

HEARING & BALANCE: SIMILAR TRANSDUCTION

• In summary, mechanical transduction of stimuli into electrical activity by the spiral organ of Corti, the
maculae, and the crista ampullaris are similar:
o There is a membrane with specialized supporting cells and sensory hair cells which are
depolarized by displacement of the cilia and thence release neurotransmitter onto the
neurofibers.
• The differences relate to the structure of the specialized membrane:
o For the spiral organ of Corti there is a tectorial membrane
o For the maculae there is an otolithic membrane
o For the crista ampullaris there is a cupula.

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4. VISION

EYE LUBRIFICATION
• The eye is lubricated by the lacrimal apparatus which includes:
o Lacrimal Gland producing a lacrimal secretion…
▪ Mucus
▪ Antibodies
▪ Lysozyme
• we blink to spread these fluids around
o …which leaves by Excretory Ducts of the lacrimal gland.
• The fluid normally
reach the Lacrimal
Punctum in the
nasal corner of
the eye leading to
the Lacrimal
Canal and
Nasolacrimal Duct
which pours in the
Inferior Meatus of
the nose.
• An excessive
secretion
overflows and run
on the face as
tears.
• Cold stimulates
lots of fluid
secretion from
lacrimal gland

EYE MUSCLE
• The eyes move in order that the object of interest falls onto the Fovea of the Retina where sensitivity
is greatest.
• Eye movements are produced by 3 pairs of extrinsic muscles:
1. Superior and Inferior Rectus Muscles
2. Medial and Lateral Rectus Muscles
3. Superior and Inferior Oblique Muscles (rotation of eye— obviously not 360° though)

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SACCADES AND SMOOTH PURSUIT
• The nervous system produces 2 types of eye movements:
1. Saccades
▪ Fast eye movements to change the point of visual attention
2. Smooth Pursuit
▪ Automatic eye movements to follow the movement of an object in visual attention
• The only volunatary decision we make is to follow an object, our nervous
system then automatically moves the eyes to stay locked on to it. We don’t
consciously adjust the speed and movement of the eyes
• We can’t slowly move our eyes to scan a scene (they just jump from one
point to another)

CONVERGENCE
• Convergence of the eyes
occurs for close objects so
that it falls on the Fovea
Centralis of both eyes to
permit stereoscopic vision.
• The eyes would then diverge
to look far away
• For other eye movements,
both eyes move in parallel
fashion.

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LIGHT
• The visible spectrum includes electromagnetic
wavelengths from:
400 Nm (blue) ---------> 750 Nm (red).
• A prism shows that white light is a mixture of all
colors.

REFLECTION
• Reflection
o The ability of light to be reflected on a surface
o The color of an object indicates the color reflected by the object

ABSORBED AND REFLECTED LIGHT

• Reflected Light
o White light is different from the perception of colors when mixed for painting because the
color viewed is that which is reflected.
o White paint reflects light of all color wavelengths so it is perceived as white.
• Absorbed Light
o A mixture of all paint colors produces black paint which does not reflect light and is thus is
perceived as black.

LIGHT PATH
• Light passes through the following structures to reach the photoreceptors:
1. The Cornea
2. Anterior Chamber
3. Pupil
4. Posterior Chamber
5. Lens
6. Posterior Segment
7. Ganglion Cells
8. Bipolar Cells
9. Photoreceptors
• The surface of the eyeball contains a delicate mucous membrane called Conjunctiva.

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• The eye itself is formed of 3 Layers or Tunics: fibrous, vascular and sensory (retina).
1. Fibrous Tunic
▪ Cornea
• A transparent part of the fibrous tunic
• Can be transplanted without rejection because it does not have any blood
vessels to carry immune response cells
▪ Sclera
• Part which forms the white of the eyeball
• Used as anchoring for the Ocular Muscles and joins the Dura Mater at the
back which forms the Envelope of the Optic Nerve
2. Vascular Tunic (Uvea)
▪ Iris
• Found anteriorly
• Composed of smooth muscles in radial and circular layout for dilating and
constricting the Pupil
o Pupil = the opening of the iris which lets light pass into the eye
▪ Ciliary Body
• Found anteriorly
• Made of smooth muscles for changing the shape of the lens to which it is
attached via suspensory ligaments.
• Ora Serrata
o Junction between the Ciliary Body and the Choroid
▪ Choroid
• Found posteriorly
• Its vessels irrigate the anterior 1/3 of the retina
3. Sensory Tunic (Retina)
▪ Includes a:
• Pigmented Layer
o Covers the choroid, ciliary body, and the posterior surface of the
iris
o Function: absorb light so that it does not reflect on the interior
walls of the eyeball
• Neural Layer
o Covers only the choroid
• Anterior Segment
o is the area in front of the lens and contains aqueous humour
▪ The ciliary body blood vessels constantly keeps this fluid present in the anterior
segment
o 2 divisions within the anterior segment
▪ Anterior Chamber: area in front of the iris
▪ Posterior Chamber: area behind the iris
• Posterior Segment
o Area behind the lens that contains vitreous humour
• Macula Lutea
o By looking into the eye one can see a yellow spot or which contains at its center the Fovea
Centralis which is the size of a pin-head.
• The Optic Disc or Blind Spot
o Part of the neural layer
o Deprived of photoreceptors since it is the site for the passage of nerves and blood vessels
which irrigate the posterior 2/3 of the retina

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• Photoreceptors:
o 2 Types:
1. Cones
• Sensitive to color
• Presence is numerous in the Fovea Centralis but decreases towards the
periphery
2. Rods
• Insensitive to color
• More sensitive to weak light
• Distributed mostly in the periphery
o Photoreceptors synapse onto Bipolar Cells which in turn synapse with Ganglion Cells whose
axons form the Optic Nerves
• Lens
o Divides the eyeball into:
1. Posterior Segment
• Containing the Vitreous Humor
o Vitreous Humor = which is a gelatinous substance
2. Anterior Segment
• Containing the Aqueous Humor
o Formed by filtration through the capillaries at the Ciliary Process
o Evacuation of the aqueous humor is done by the scleral Venous
Sinus (Canal Of Schlemm)
▪ Glaucoma = blockage of these sinuses
• The anterior segment is divided by the Iris into
o Anterior Chamber
o Posterior Chamber

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PUPIL SIZE
• Pupil
o Function: controls the amount of light
entering the eye.
• Muscles fibers organized in a circular function in
the center → reduce size of pupil
• Muscle fibers organized Radially on the outer
circle→ pull to enlarge the size of pupil

PUPILLARY LIGHT REFLEX

• Pupillary Reflex
o A fast contraction of the circular smooth muscles of the iris in response to light
o Mechanism/Pathway:
▪ Light
activates….
▪ Retinal Ganglion Cells
that relay to…
▪ Pretectal Nucleus Neurons
which relay to….
▪ Edinger-Westphal Nucleus Neurons on both the left and right sides
which pass via…
▪ Third Cranial Nerve
to relay to….
▪ Ciliary Ganglion Neurons
that innervate…
▪ Constrictor Muscle of the Iris

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FOCUS AND INVERSION
• Focusing
o Process that involves the convergence of
point sources through refraction of light by a
lens
o The lens of the eye:
▪ Focuses light onto a single point on the retina
▪ Causes the image to be inverted on the retina
▪ Ex. Focus on a camera
▪ Albino, lacks melanin, so has difficulty focusing because the light gets reflected
rather than absorbed and hits several spots in the retina
• Inversion
o Visual image on the retina is kept inverted along the path to the primary visual cortex

ACCOMMODATION
• Accommodation Reflex
o Bends the light rays towards the focal point of
the retina
o Function: bring the visual field is in focus
▪ Relaxation of the ciliary body
increases tension on the Suspensory
Ligaments and causes stretching of
the lens
• Result: permits far vision
▪ Contraction of the ciliary body
decreases tension of the Suspensory
Ligaments and allows the lens to
bulge from its elastic properties
• Result: permits near vision
o Accommodation of the lens is used to clearly
see objects that are at a distance between the
Punctum Proximum and Remotum
▪ An object that is closer than the
Punctum Proximum (typically to 25
cm) cannot be seen clearly because
accommodation of the lens (bulging
of the lens produced by contraction
of the ciliary muscles) is at a
maximum
▪ An object at the Puntum Remotum
(typically 2-6 meters) or further does
not require accommodation of the
lens to be seen clearly because the
light entering the eye is parallel

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MYOPIC AND HYPERMETROPIC
• Emmetropia
o Normal vision is 20/20
▪ 20/20 = distance you must be in order to
clearly see an object that a normal subject
sees clearly at 20 feet
▪ Focal point is right on the level of the retina
▪ If fraction is greater than 1, than better than
normal vision, less than 1 is worse
• Myopia
o Subject has poor far vision
o The image forms in front of the retina
▪ It's not a defect of the lens but rather that
the eye is too deep
o Correction requires concave lenses to push back focal point onto the retina
• Hyperopia
o Subject has poor near vision
o The image forms behind the retina
▪ It's not a defect of the lens but rather that the eye is not deep enough
o Correction requires convex lenses
o People with hyperopic or myopic still have regular accommodation.
• Astigmatism
o Due to unequal curvatures of the lens or cornea
o Correction requires a combination of both concave and convex lenses

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PRESBYOPIA
• Presbyopia
o Accommodation is decreased
o Due to loss of the elasticity of the lens
o Developed with age
▪ Level of accommodation decreases between the ages of 10-50

CATARACTS
• Cataract
o Opacity of the lens
o Clouds and blocks vision
o First treatment of cataracts in arab countries (pushed needle through
eye to move lens out of the way)

PHOTORECEPTORS
• Photoreceptors
o Found on the retina
o Function: Detect light
o Made of:
▪ Outer Segment (Light Receiving Area)
• Contains visual pigment (molecules)
for photoreception
▪ Inner Segment (Metabolic Area)
• Connecting Stalk – connects the two
segments

CONE SENSITIVITIES
o Blue, Green and Red Cones
▪ Detect the visible spectrum
▪ Maximally sensitive for light at a particular
wavelength….
• Blue cones = 420 nm
• Green cones = 530 nm
• Red cones = 560 nm
▪ …but also respond to lesser extents for light
at wavelengths near this maximum sensitivity

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RED-GREEN PERCEPTION TEST
• Color Blindness
o Most common one is a difficulty in distinguishing the colors red and green
o Cause: a defective X chromosome for the coding of red or green cones

RHODOPSIN CYCLE
• Visual pigment is made of 2 molecules
1. Retinal
2. Opsin
▪ Rods: Scotopsin
▪ Cones: 3 kinds of Photopsins (to detect the colors blue, green, and red)
• It is the retinal that is photosensitive:
o in the dark the retinal binds to the opsins to form the visual pigment
o in the light the retinal dissociates with the opsin (the color absorbed by the retinal depends
on the opsin to which it binds)
• Below is the visual cycle for rhodopsin which is the visual pigment of the rods:
o Light + Rhodopsin =
Scotopsin + All-Trans Retinal
o Dark + All-Trans Retinal +
Isomerase + ATP = 11-Cis
Retinal
o Dark + 11-Cis Retinal +
Scotopsin = Rhodopsin
• Vitamin A loses 2 hydrogen’s to form
an external source of 11-Cis Retinal.
• Effects of light and dark on
pigmentation
o Light reduces the quantity of
visual pigment → reduced
sensitivity to light
o Darkness allows the
regeneration of the visual
pigment →increased light
sensitivity
▪ Adaptation to dark
is slower because
ATP and 2
metabolic steps are
required to
regenerate
sufficient quantity
of visual pigment
• Photoreceptors are not always active
in daylight because:
o As we glance at different
objects in our environment,
some photoreceptors would
receive the black parts of
the object which do not
reflect light

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TRANSDUCTION
• Photons (light waves) transform Rhodopsine of the rods into:
o All-Trans Retinal, and
o Scotopsin
▪ Scotopsin triggers a metabolic cascade which results in: CIC
i. Conversion of cyclic-GMP into GMP
ii. Inactivation of the photoreceptor
iii. Closure of sodium channels

PHOTORECEPTOR ACTIVATION
• In the dark, there is plenty of cyclic-GMP, which stimulates:
1. Opening of the sodium channels
2. Depolarization of the cells
• In the light there is a reduction in cyclic-GMP, which leads to:
1. Closing of the sodium channels
2. Repolarisation to inactivate the photoreceptor.

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VISUAL FIELDS
• Visual Field
o Width of space which one can see
without moving
• Divided by the nose into the left and right visual
fields
• There is overlap of the parts of the visual field
seen by each eye.
o Overlap allows depth perception

SYNAPTIC RELAYS
• Photoreceptors
…synapse onto…
• Bipolar Cells
…which in turn synapse with…
• Ganglion Cells whose axons form the optic
nerves.
o Ganglion cell receives information only
from a single cone or from several rods.

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CONNECTIONS TO CORTEX
• Information coming from the rods and the blue, green and red cones follow separate pathways to the
primary visual cortex without mixing so that one can perceive the different colors
• Contralateral Visual Field
1. Visual Stimuli
…from the contralateral visual field are sensed by…
2. Ganglion Cells
…which project to the…
3. Lateral Geniculate Cells of the thalamus
…whose axons form the…
4. Optic Radiation
…terminating on cells of the…
5. Primary Visual Cortex in the Occipital Lobe → only receives information from the
contralateral visual field
• Ipsilateral Visual Field
1. Visual Stimuli
…from the Ipsilateral Visual Field strike the…
2. Nasal Retina
…which have…
3. Ganglion Cells
…that project through the…
4. Optic Chiasm
…to reach the….
5. Contralateral Lateral Geniculate Nucleus of the Thalamus
• Optic Nerve vs. Optic Tract
o Optic Nerve = The ganglion cell axons between the eye and the Optic Chiasm
o Optic Tract = The part of ganglion cell axons between the Optic Chiasm and the Thalamus

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• Deficits: (Diagram above)
A. Blind in right eye
B. Cut optic chiasm: no crossing over, so we can see only information coming through ipsalateral
signals, info from left visual field of left eye
C. Damage at level of right side optic radiation (after geniculate nucleus)
D. Lateral geniculate nucleus projection that fans out (optic radiation) and sometimes ppl who have a
stroke will lose part of the radiation, losing one quadrant of vision
E. See (c); with macular sparing = Retention of macular function in spite of losses in the adjacent
visual field (ex. as with homonymous hemianopia) due to the fact that most cortical lesions are
not large enough to affect the whole extensive cortical area representing the macula, thus leaving
some of the area unaffected

SECONDARY CONNECTIONS
• Apart from projections towards the lateral geniculate nucleus of the thalamus for visual perception,
ganglion cells have other projections towards:
1. Suprachiasmatic Nucleus (SCN) of the hypothalamus
▪ Which takes part in the biorhythms from the light and dark cycles
2. Superior Colliculi
▪ Which participates in fast movements (saccades) of the eyes towards a visual
stimulus
▪ Even people with blind sight (cortical damage), eyes will automatically still move
towards a strong visual stimulus (reflex of eyes).
3. Pre-Tectal Nuclei
▪ Which participates in the pupillary light reflex where light in one eye produces
pupillary constriction in both eyes.

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5. SOMESTHESIA

SIMPLE RECEPTORS
• Simple receptors in the somatic system detect the stimuli to the body and those in the vegetative
system detect stimuli to the internal organs
• There are 2 kinds of simple receptors:
1. Free Nerve Endings. These include:
a. Non-Specialized Naked or Free Nerve Endings
• Detect pain, temperature, or strong pressure throughout the body
b. Merkel Discs
• Detect light touch at the bottom of the epidermis
c. Hair Follicle Receptors
• Detect movement of hair
2. Encapsulated Dendritic Endings. These include:
a. Meissner's Corpuscle
• Of greater density in the hands, lips,
and nipples
• Have a discriminative tactile function
b. Krause Corpuscles
• Similar to Meissner's corpusclebut,
but are located in mucous
membranes like the mouth
c. Pacinian Corpuscle
• In the dermis and the subcutaneous
tissue
• Respond to changes in pressure since
the capsule partly adapts to
maintained pressure
d. Ruffini Corpuscle
• In the dermis, the subcutaneous
tissue, and the articular capsules
• Respond to stretching
e. Golgi Tendon Organs
• In the tendon
• Respond to applied tension
f. Muscle Spindles
• In the muscle
• Respond to changes in muscle length.

PAIN STIMULATION
• Painful stimulus will cause the release of substances… o Serotonin
…that will stimulate Free Nerve Endings to produce action potentials that will then o Prostaglandins
1. Cause the nerve terminals to release Substance P o Leukotrines
▪ Substance P stimulates…
• Blood Vessels
• Mastocytes
▪ …which in turn produce inflammation
2. Propagate the action potentials towards the CNS

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PAIN TRANSDUCTION
• The substances released by painful stimuli will act like
Ligands binding to Metabotropic Receptors
o This produces postsynaptic depolarization

TEMPERATURE STIMULATION
• Thermoreceptors
o Sensitive to either heat or cold

TEMPERATURE TRANSDUCTION
▪ Changes in temperature cause
→ change in the conformation of the receptor such
that
→it opens or closes the ionic channel

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MECHANICAL STIMULATION
• Touch is detected by Mechanoreceptors in the skin:
o Merkel's discs
o Hair follicle receptors
o Meissner's corpuscles
o Krause corpuscles
o Ruffini's corpuscles
o Pacinian corpuscles
▪ Figure below: Capsule of the
Pacinian corpuscle allows
receptor to detect increases or
decreases in pressure as during
vibrating stimuli
▪ Receptor responds only to
increases in pressure when
capsule is removed

MECHANICAL STIMULUS TRANSDUCTION

• All mechanoreceptors…
o e.g. Ruffini's corpuscles, Golgi tendon
organs, Muscle spindles
• …have channels with gates that open when stretch
or pressure is transmitted to the cell membrane with
its Cytoskeletal Cables that are attached to the
channel gate

PROPRIOCEPTORS
• Proprioception
o Sense associated with posture or movement
o Detected by mechanoreceptors like:
▪ Ruffini's corpuscles in the articulations
▪ Golgi tendon organs
▪ Muscle Spindles (these especially)

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• Have a capsule containing Intrafusal Muscle Fibers surrounded with
sensory endings (primary and secondary)
• Sensory endings measure the length of the central part of the intrafusal
fibers (this corresponds to the length of the extrafusal muscle)
• Sensitivity of the muscle spindles can be modified by activation of gamma
motoneurons that cause contraction of the extremities of the intrafusal
fibers

PROPRIOCEPTOR FUNCTION
• Golgi Tendon Organ
o Detects tension in the tendon produced by Extrafusal Muscle Contraction
• Muscle Spindles
o Detects differences between Extrafusal and Intrafusal Muscle Fiber length produced either
by passive muscle stretch or by contraction of intrafusal muscle fibers
o Besides allowing perception of muscle length, muscle spindles can trigger the stretch reflex
which corrects perturbations in muscle length.

SENSORY PATHWAYS
• It is by a relay of 3 neurons that the information of a stimulus on the body is conveyed to the cerebral
cortex for perception
o First Order Neuron
…sensory neuron which forwards information to the CNS where it synapses on the…
Ascending pathway o Second Order Neuron
which conveys …which in turn projects to the…
information to the o Third Order Neuron in the Thalamus
sensory cortex …which projects to the…
o Cortex
▪ Perception is achieved by the networks of neurons in the cerebral cortex
• There are 2 main pathways to transmit sensory information to the cortex

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1. Anterolateral Pathway
▪ Conveys sensations of:
• Pain
• Temperature
• Strong pressure
▪ Stimulus transmitted by first order neuron (sensory) synapses in the spinal cord on the
second order neuron which sends projections crossing towards the contralateral side
and ascending to the thalamus in order to make synapse on the third order neuron
▪ This thalamic neuron (third order) then projects to the primary somatosensory cortex
2. Medial Lemniscus Pathway
▪ Conveys sensations of:
• Discriminative touch
• Proprioception
▪ Stimulus is transmitted by first order neuron (sensory) which projects up the dorsal
columns of the spinal cord to synapse on a second order neuron in the Dorsal Column
Nuclei (Gracilis or Cuneatus in the medulla) which sends projections crossing to the
contralateral side and up to the thalamus where it synapses on a third order neuron
▪ This thalamic neuron (third order) then projects to the primary somatosensory cortex

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SOMATOSENSORY CORTEX
• The primary somatosensory cortex lies in the Postcentral Gyrus

SENSORY HOMUNCULUS
• The various parts of the body are represented by a Homunculus in the primary somatosensory cortex

INTERNAL ORGAN PAIN

• Sensory fibers also come from internal organs → providing pain information when there is organ damage

REFERRED PAIN
• Visceral Organs (like the kidneys, stomach, and heart)
o Homunculus does not have sensory representations for these organs
o They have Nociceptors
▪ These join the same pathways of other somatic sensations which are represented in the
homunculus of the primary somatosensory cortex
• Ex: Nociceptors of the heart join the sensory pathways of the arm and shoulder
leading to the homunculus in the primary somatosensory cortex
• This explains the referred pain of the heart to the arm and shoulder

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PERCEPTION

THREE (3) LEVELS

• 3 levels of transmission to achieve perception. Here we will use the somatic senses as example, but this
equally applies to the special senses.
1. Receptor Level
▪ Includes the sensory neuron (first order) which forwards information to the cns to
synapse on the second order neuron.
2. Ascending Pathway Level
▪ Includes second and third order neurons
that convey information to the primary
sensory cortex.
3. Perceptual Level
▪ Contains the cortical neurons that will
analyze sensory information to provide
perception.
• It is necessary to distinguish between sensation and
perception:
o Sensation occurs at the unconscious level
o Perception occurs at the conscious level
o Ex. If someone were in a very hot room and noticed
the heat only 10 minutes later then one would
conclude that only the sensation of heat existed
during the first 10 minutes and this was followed by
sensation plus perception.
• Somatic sensory information arrives first in the primary
somatosensory cortex (raw information)
• It is analyzed by the surrounding association sensory cortex
(processed information)
• Lesions:
o A lesion of the Primary Somatosensory Cortex
▪ Will eliminate sensation and perception
resulting in the symptom of anesthesia
o A lesion of the Tactile Association Cortex
▪ can produce tactile agnosia (where the
patient can identify an object by looking at it
but not by manipulating it)
o A lesion of the Visual Association Cortex
▪ Can produce a visual agnosia where the
patient cannot identify an object by looking
at it but can easily identify it by manipulating it.

CORTICAL CONNECTIONS
• This figure shows 300 interconnected neurons. By contrast, there are 10-50
thousand interconnected neurons in 1 mm cube of cortex.
• This complex connectivity has slowed progress in understanding
perception. Consequently, we will just describe some general principles of
perception.

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DETECTION AND MAGNITUDE
• Perceptual Detection
o The simplest perception
o It allows us to know if the stimulus
exists!
• Magnitude Estimation
o Allows us to know if the stimulus is weak
or strong!
o To perceive the intensity of a stimulus,
the cortex uses the discharge frequency
of a single neuron…
▪ low frequency corresponds to a
stimulus of weak intensity
▪ high frequency corresponds to a
stimulus of strong intensity
o …and the number of cells active
▪ More cells means stronger
stimulus

SPATIAL DISCRIMINATION
• Spatial Discrimination
o Allows us to know if the stimulus is large!
o The Limit Of Space Discrimination:
▪ Measured as the minimum distance
that 2 points can be perceived
▪ Depends on the density of receptors
▪ If the receptors are 20 mm apart then
you will not be able to detect 2 stimuli
that are 5 mm apart
• If you move the two points near only one receptor, both
stimuli go to that receptor, then only one signal is sent to
brain and we perceive only 1 pin prick
o Ex. On tongue we can detect features of objects
best, worst with our back
• Ability to detect precise size information depends on the
density of receptors on the skin.

FEATURE ABSTRACTION
• Feature Abstraction
o Allows us to know characteristics of stimulus!
o The skin does not have receptors specialized to
detect the rough, hard, or smooth characters of
a stimulus which are required for us to
distinguish among silk, velvet, and leather
o The nervous system analyzes information
transmitted by all its touch receptors (e.g.
Merkel discs, Meissner's corpuscles, Pacinian
corpuscles) in order to extract the characters
that make it possible to distinguish objects

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PATTERN RECOGNITION
• Pattern Recognition
o Allows to us know the pattern of a stimulus!
o Types of patterns:
▪ Tactile patterns
▪ Visual patterns (e.g. faces)
▪ Auditory patterns (e.g. melodies)
• Figure below: If you center your focus on the pencil then the red dot will be in the middle of your fovea.
o Each square represents a neuron in the cortex,
o Neurons only perceive ONE square, so neurons must communicate with each other to figure out
the complete perceived image
o If head is bumped, the center of focus changes, and image must be reanalyzed
▪ Solution: vestibular ocular reflex, easier for visual system to move eyes back where they
were, instead of reanalyzing the whole image.

RETINAL SLIP
• Moving eyes → center of focus has now changed from the previous image
o Different cortical cells are receiving the image of the pencil
o Therefore, the visual analyses to perceive the pencil has to occur again
• If this change in focus was produced by a perturbation to the head, then the Vestibulo-Occular Reflex
(VOR) would reflexively move the eyes back on the original center of focus
o So that the same cortical cells receive the image
o Thus we don't have to analyse the image again

CENTER-SURROUND CORTICAL RESPONSES

• Visual Perception
o Very complex
o Important for several functions of the nervous system.
▪ This is reflected by the fact that 50% of the cortex of the monkey is used for visual
processing as compared to 11% of the cortex for processing somatosensory information
and 3% of the cortex for processing auditory information.
• To identify an image, the visual cortex processes the raw visual data that it receives.
o We only know the first parts of this information processing.

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• There is a Center-Surround Antagonism
o Purpose of the processing is to clearly extract the point sources of visual stimuli
i. Some neurons exhibit an On-Center Activity
• Where light in the center of the receptive field increases neuronal activity while
light surrounding the center decreases neuronal activity
ii. Other neurons show the opposite response or Off-Center Activity
• The activity of the cells depends on the difference in the intensity of light on
the center and on the periphery of the receptive field
▪ In this way provides visual contrast information

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SIMPLE CORTICAL RESPONSES
• Cells with Center-Surround Responses →project to simple cells which detect edges of objects
• A vertical bar of light would cause strong firing of these cells
• A horizontal bar of light would not cause firing

COMPLEX CORTICAL RESPONSES

• These simple cells in turn →project to complex cells whose role is not clearly known

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SENSORY FILTERING
• There are numerous receptors on the body which are constantly
stimulated by our clothes, our posture, or other objects like a chair.
o Our brain cannot constantly supervise all this information.
• Therefore, the nervous system has mechanisms to block
information.
o Ex) The inhibition of the Ascending Lemniscal Pathway at
the level of the dorsal column nuclei
o This inhibition can block the stimuli from the weight of
our clothes on our body
• This mechanism also exists for other stimuli such as for the visual
stimuli.
o Ex) Several drivers have at one time noted that they drove
a long stretch of a familiar highway without really noticing
the road.
o That is due to the fact that the nervous system blocks
information that it can predict.

MOTOR SYSTEMS

SENSORIMOTOR ARC
• Motor systems
o Those circuits of the nervous system which enable us to make movements.
o It is the Frontal Cortex contains the neurons of the motor cortex that take part in voluntary
movements.
▪ Most Anterior Parts of the frontal cortex (e.g. premotor or association motor cortex)
• Takes part in the most abstract planning of movements
• A lesion here causes Apraxia where there is loss of movement planning but not
of muscular force
▪ Most Posterior Parts of the frontal lobe (e.g. primary motor cortex)
• Takes part in the execution of fine movements
• A lesion here produces Paresis where there is a loss of fine voluntary
movements but feeble movements remain because they are carried out by the
secondary motor pathways
o The motor cortex…
▪ …asks the Cerebellum how to coordinate the movement
▪ …asks the Basal Ganglia how to initiate the movement
o Once it receives the answers, the motor cortex sends a signal via the Corticospinal Tract to the
motoneurons of the spinal cord to carry out the movement.

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MOTOR HOMUNCULUS
• The voluntary movement
commands derive from the
primary motor cortex
• Primary motor cortex has a
Somatotopic Organization →
called the Motor Homunculus

PRIMARY DESCENDING TRACT

• The Corticospinal Tract consists of a neuron in the primary
motor cortex that sends a descending projection to the
spinal cord. It is divided into 2 tracts.
1. Lateral Corticospinal Tract
▪ Crosses to the contralateral side at the
level of the Pyramidal Decussation in the
Medulla
▪ Continues down in the lateral white
matter until it reaches its Target Spinal
Cord Motoneuron for the control of distal
limb muscles
• Thus distal muscles often receive
controlateral cortical control.
2. Ventral Corticospinal Tract
▪ Travels ventrally in the spinal cord white
matter
▪ Reaches its target Controlateral or
Ipsilateral Spinal Motoneurons for the
control of proximal muscles.
• Thus proximal muscles often
receive bilateral cortical control
• The primary motor cortex also sends Corticobulbar Fibers
to the Facial Motoneurons
o Control of facial movements
o The forehead receives bilateral cortical control
o The lower part of the face receives only
controlateral cortical contro
o The projections from the primary motor cortex are
for fine control of movements.

SECONDARY DESCENDING TRACTS

• There is also Synergistic Motor Control
o Used to assist fine movements
▪ e.g. stabilize the shoulder and arm during
manual tasks
o These are secondary descending tracts such as:
▪ Rubrospinal Tract
• To control limbs to assist hand
movements

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▪ Reticulospinal Tracts
• To control
posture
▪ Vestibulospinal Tract
• To maintain
balance
▪ Tectospinal Tract
• To adjust the
head for visual
stimuli.
• Lesion of the ventral root:
o Eliminates movements without
affecting sensations
• Lesion of the dorsal roots:
o Eliminates sensory inputs
• Lesion of the spinal nerve:
o All movements, sensations and reflexes are lost
• Complete transection of the spinal cord
o Only voluntary movements and conscious sensations are lost (reflexes remain).
o Ex) Complete transection at the level of L1 will produce a complete loss of voluntary movements
(paraplegia) and sensations (anesthesia) of the lower limbs but reflexes can persist because the
sensory inputs to the spinal cord and the motoneuron output from the spinal cord are intact.

CEREBELLAR AND STRIATAL LOOPS

• Figure right:
o Summarizes the
connections involved
in the control of
movements
o Note the feedback
loops from the
motor cortex:
1. To the
cerebellum
2. To the basal Corticospinal
ganglia tract
o These 2 feedback
loops are to assist
the cortex in the
control of
movements
• Lesions of the cerebellum and
basal ganglia
o Produce clear deficits
in the control of
movements

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CEREBELLAR DISEASE
• Cerebellar lesion
o Ataxia (lack of coordination)
▪ Which is described as a drunken sailor's gait
▪ Errors of arm movement amplitude
• I.e. Dysmetria (length of movement is wrong)
o Hypometria
▪ Where the movement stops
before reaching the target
o Hypermetria
▪ Where the movement
exceeds the target
▪ Intentional Tremor
• Where the hand trembles as the patient tries
to be precise as when inserting a key in a lock.
o Loss of coordination
▪ Also shown by Dysdiadokinesia
• An inability to make fast rhythmic movements
(ex. rapid pronation and supination of hand)
o Scanning speech
▪ Bad coordination of the voice
• Fundamental cause of cerebellar symptoms is a loss of control of the muscle spindles
o This causes the nervous system to not have exact knowledge:
▪ Muscle length
▪ Posture (consequence of lack of knowledge of muscle length)
o Loss of muscle spindle control also produces a reduction of Muscular Tone
▪ Muscular Tone = resistance to passive movement of a limb

BASAL GANGLIA DISEASE

• A patient with basal ganglia disease loses control of the initiation of movements. There are two extremes:
1. Bradykinesia - The movement is difficult to initiate
▪ Cause: a loss of Dopamine release from neurons of the Substancia Nigra Pars Compacta
• Ultimately leads to excessive inhibition of Thalamic Projections to the cortex
▪ The patient with Parkinson's Disease shows Bradykinesia
• Increase in the muscular tone
• Tremor at rest
• Loss of the expression of the face
• Forward inclined posture
• Shuffling gait (short, uncertain steps,
with minimal flexion and toes
dragging)
2. Hyperkinesia - The movement is initiated involuntarily
▪ Cause: Lesions of the Subthalamic Nucleus (i.e.
degeneration of the Caudate and Putamen)
lead to excessive activation of Thalamic
Projections to the cortex
▪ A patient with Huntington's Chorea shows
Hyperkinesia
• Rapid involuntary movements

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SPINAL PATTERN GENERATOR
• The spinal cord contains…
o Motoneurons
o Networks of neurons; can produce
rhythmic movements of walking
▪ The basic program for walking
or locomotion is in spinal cord
▪ However, these spinal neural
networks are normally
activated by descending
voluntary commands from the
Cerebral Cortex
▪ These commands alter the
basic locomotor pattern to
adapt to the terrain

MUSCLES: NEUROMUSCULAR JUNCTION

• The voluntary action chosen by the nervous
system is carried out by the skeletal muscles
1. The Axonal Endings of motoneurons
…release Acetylcholine onto…
• There's 3-4 times more neurotransmitter released than necessary
• This means so opening of these channels produces a depolarization which is
always above the threshold for excitation of the action potential
• This is then propagated throughout the muscle to produce muscular
contraction
• Therefore, an action potential in the motoneuron will always produce an action
potential in the muscle and contraction
• This is why the motoneuron and its target muscle fibers are called a motor unit
2. Neuromuscular Junction
…which then binds to…
3. Ligand-Dependent Channels on the skeletal muscle

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NEURAL INTEGRATION: SENSORIMOTOR ARC
• Neural Integration
o Between the input of sensory information and the
output of motor commands
o Occurs for the actions of Reflexes and for the voluntary
actions
▪ Reflexes: We will start with them since their
mechanisms are simple, very well-known
▪ Neural Integration: For voluntary actions; we
will then speak about them because their
mechanisms are far more complex and much less understood.

REFLEXES

SOMATIC REFLEXES

BASIC REFLEX ARC

• Reflex: is an automatic Action triggered by a Stimulus
o Action - can be a movement, salivation, an increase in heart rate or others
o Stimulus - can be mechanical, visual, auditory or others
• The reflexes are classified 2 ways
1. According to the control of the reflex:
a) Involuntary reflex
▪ e.g. Stretch reflex
b) Learned or conditioned reflex
▪ e.g. Automatic movements by a goal keeper
2. According to the reflex target:
a) Autonomic reflexes
▪ Effectors are the glands or the smooth muscles or cardiac muscles
b) Somatic reflexes
▪ Effectors are the skeletal muscles
• Here is a list of the elements which form part of a reflex:
1. Receptor
To convert the stimuli into an electric signal
2. Sensory Neuron
To forward information to the CNS
N.B. Simple receptors belong to the sensory neuron
3. Integration Center
Which includes the neurons of the CNS that will transform the information of the
sensory neurons into commands for the motoneurons
Polysynaptic Reflexes - integration is done by interneurons
Monosynaptic Reflexes -
integration is done by the
motoneuron itself
4. Motoneuron
To activate the effector
5. Effector
E.g. Muscle or gland

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WITHDRAWAL REFLEX
• Arms
o Placing the hand in fire → flexion of the arm
• Legs
o Walking on glass → flexion to remove the painful
stimulus (flexor reflex), extension of the opposite leg
to maintain an upright position (crossed extensor
reflex)
• The expression of reflexes can be partially controlled by the
voluntary actions of the cerebral cortex
o Normally : Protection & Balance reflexes have priority
over > the control of voluntary movements
o Exceptional Cases: It is possible to voluntarily modify
a reflex
▪ Ex: If we burn ourselves by taking a pot of
boiling water, we are able to inhibit the
reflex withdrawal in order to avoid dropping
the boiling water on a child.

TENDON REFLEX
• Tension in the agonist muscle (i.e. prime mover) causes:
o Inhibition of the agonist
o Excitation of the antagonist
• GOAL: To prevent the development of too much tension in the
agonist
o Too much tension = ripping the tendon out of its
insertion into the bone
o Thus, this is a protective reflex
o Common in athletes who generate a lot of force
• Can be inhibited in certain conditions (i.e. blocked this reflex
to generate force in times of extreme stress)

STRETCH REFLEX
• Stretching of the agonist causes…
o …excitation of the agonist
o …inhibition of the antagonist
• GOAL: To return agonist muscle length to its original length
• The gamma-motoneurons define the desired muscle length for
posture and the reflex corrects and perturbations in posture
o Ex. Tapping with hammer onto patellar tendon, the
tap forces a rapid microscopic stretch (FAST) of the
quadriceps, which is detected by muscle spindles who
send signals to spinal cord, causes muscles that was
stretch to contract back to original length, causing
extension of leg
o This reflex works for almost every muscle
• Spinal Cord Transection
o Causes this reflex to become exaggerated → may lead
to muscle spasms

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PLANTAR REFLEX
• Plantar Reflex
o Expressed as an extension of the big toe
o Abduction of the other toes
o Also called the Sign Of Babinski
• How it happens:
o Stimulation of the sole of the foot activates sensory neurons projecting to spinal cord
interneurons that will produce the sign of Babinski…
o …UNLESS the Corticospinal Projections are available to modify the behavior of the spinal cord
interneurons forming this reflex.
▪ Lesion of the cortciospinal pathway
• Scratching the sole of the foot in patients with this lesion causes plantar reflex
▪ Not fully developed corticospinal tract
• Seen in children less than one year; they too will show plantar reflex when the
sole of their foot is scratched

AUTONOMIC REFLEXES

ORGANISATION
• Hypothalamus
o Controls the internal milieu
o Receives information of the state of all the internal
organs and systems
o To maintain homeostasis, the hypothalamus acts on two
systems:
1. Endocrine System
• Secretes hormones in the blood supply
that travel to target tissues and
stimulate homeostatic responses
• We will not be covering the endocrine
system
2. Autonomic Nervous System
• Uses neurons to control the target
organs

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HOMEOSTASIS
• Homeostasis
o Involves mechanisms that
take corrective actions to
prevent changes in the
value of a variable

EXAMPLE: TEMPERATURE CONTROL

• Figure below: An example of
homeostatic mechanisms for the
control of body temperature.
• Not on exam! (no figures on exam,
remember?)

SYMPATHETIC AND
PARASYMPATHETIC
• All (almost all) targets of the
autonomic nervous system receive
inputs from both divisions:
o Sympathetic → T1-L2
o Parasympathetic → Cranial nerves 3, 7, 9 and 10 and S2-S4

EFFECTS
• Sympathetic and Parasympathetic divisions usually act on the same target organs, but their actions are
opposite → i.e. If one causes contraction, the other causes relaxation
o Sympathetic division
▪ Highly mobilized in extreme situations
• e.g. The “E” system: Exercise, Excitement, Emergency, Embarrassment
▪ When there is an extreme situation, the body must be prepared to respond quickly. The
sympathetic division causes:
• Pupils to dilate → improved vision
• Bronchioles to dilate → increased ventilation
• Heart to beat stronger, faster → increased blood flow
• Circulation to be re-directed
From……Digestive, urinary and genital organs
To………..Skeletal muscles
o N.B. there is not enough blood to irrigate all organs
o Parasympathetic division
▪ Involved in routine situations
• e.g. The “D” system: Digestion, Defecation, Diuresis
▪ In routine situations, the parasympathetic division reacts opposite to the sympathetic
division and causes:
• Pupils to constrict → regular, lowered vision
• Bronchi to constrict → regular, lowered ventilation
• Heart to beat weaker, slower → reduced blood flow blood
• Circulation to be re-directed
From……Skeletal muscles
To………..Digestive, urinary and genital organs

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o There is always a balance between the activities of the 2 divisions: Sympathetic Tone and
Parasympathetic Tone
▪ Thus both are active but one is more active than the other
o The autonomic system can independently select targets that must be controlled.
▪ Sympathetic or parasympathetic neurons are not always active on all their connections
with target organs
• e.g. Bladder does not contract every time we relax

NEUROTRANSMITTERS
• The outputs of the autonomic system include a chain of 2 neurons:
1. Preganglionic Neuron – 1st neuron
▪ It’s body is in the CNS
▪ Preganglionic Axon
• It synapses with the cell body of the 2nd neuron, the postganglionic neuron
▪ Neurotransmitter:
• Acetylcholine
2. Postganglionic Neuron – 2nd neuron
▪ It’s Autonomic Ganglion is outside the CNS
▪ Postganglionic Axon
• Projects to the target effector
▪ Neurotransmitter

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• Released directly into the extracellular space
o This then drains into the blood and acts on several effectors irrigated
by blood vessels:
▪ Smooth muscle
▪ Heart (cardiac) muscle
• Varies with the division which may be:
o Sympathetic → Norepinephrine
o Parasympathetic → Acetylcholine
• Adrenal Medulla: It’s postganglionic neurons are an exception because these
cells release:
o Epinephrine (80%)
o Norepinephrine (20%)
• Other features that distinguish sympathetic and parasympathetic divisions of autonomic nervous system
1. Origins of the preganglionic cell bodies
▪ Parasympathetic division
• Brain (cranial nerves 3,7,9,10)
• Sacrum (S2-S4)
▪ Sympathetic division
• Thoraco-lumbar region of the spinal cord (T1-L2)
2. Length of pre- and post-ganglionic axons:
▪ Parasympathetic division
• Pre = long
• Post = short
▪ Sympathetic division
• Pre = short
• Post = long
3. Location of ganglia:
▪ Parasympathetic ganglia
• In the target organ (effectors)
▪ Sympathetic ganglia
• In proximity to the spinal cord
• Around the spine, sympathetic ganglia form:
o Sympathetic Trunk - on both sides
o Prevertebral Ganglia - at the front

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RECEPTORS
1. Cholinergic (name = linked to Acetylcholine)
Receptors – 2 types
1. Nicotinic Receptors
▪ Named because of their
activation by Nicotine
(source = tobacco)
▪ Found on:
• Skeletal muscle
• Postganglionic
neurons
2. Muscarinic Receptors
▪ Named because of their
activation by Muscarine (source = wild mushroom)
▪ Found on:
• Target organs of postganglionic neurons
o We can identify these 2 receptor types by the application of Nicotine or Muscarine…
BUT… it is noted that it is Acetylcholine that stimulates both nicotinic and muscarinic
receptors in the human body
2. Adrenergic (name = linked to Epinephrine & Norepinephrine) Receptors – 2 types
1. Alpha
▪ Divided into subclasses alpha 1, alpha 2
2. Beta
▪ Isoproterenol (drug) acts only on beta-adrenergic receptors
▪ Divided into subclasses beta 1, beta 2, beta 3,
o We cannot distinguish between these 2 classes of adrenergic receptors with Norepinephrine and
Epinephrine…
BECAUSE… these hormones stimulate all adrenergic receptors
o The 2 classes and 5 subclasses were identified thanks to the specificity of certain drugs
▪ Advantage of the classification of receptors
• One can easily treat receptors that are connected to a medical problem rather
than treating all receptors
• Ex. Asthma Attack (bronchioles constrict), corrected by using:
o Norepinephrine
▪ This can correct the problem since the body normally releases
norepinephrine to dilate the bronchioles through their beta
receptor
▪ However, this injection will stimulate all the alpha and beta
receptors in the body (we cannot apply only to specific
receptors as do the postganglionic neurons that innervate the
lungs)
o Isoproterenol
▪ We will only stimulate beta receptors, correcting the problem
▪ However, this will cause other problems because the heart
also has beta receptors (Heart has beta 1 receptors and the
lungs have beta 2 receptors)
o Salbutamol
▪ This will correct the problem by activating only the beta 2
receptor
▪ This won’t cause many side effects

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VOLITION
• Volition
o Ability to decide on a particular action
o Requires higher motor centres

CORTEX ROLES
• GOAL of Nervous System = To adapt to changes in the internal and external environments for survival
o Autonomic Nervous System
▪ Has automatic responses to stimuli in order to adapt to changes in the internal milieu
▪ Genetic memory
• Automatic responses passed down from generation to gen. through genes
• Genes form the neuronal circuits generating these responses
• This genetic memory also exists for the involuntary reflexes such as the flexion
reflex that automatically protects us from painful stimuli in the external milieu
▪ But there do not exist automatic responses for all stimuli in the external milieu
• Ex. Response to a flat tire on the way to a final exam is not automatic
o Cerebral Cortex
▪ One must use the higher mental functions of the cerebral cortex in order to find an
appropriate response or solution
• Ex. Repair it or take a taxi
▪ Circuits of the cerebral cortex
• Essential for individual to be able to analyze situation and choose a solution
• Association cortices
o So far, we learned that the following two types of information exist:
▪ Sensory information
• Reaches the cortex in areas named Primary Sensory Cortex
▪ Motor information
• Leaves the cortex from the Primary Motor Cortex
o Surrounding these sensory and motor areas are association cortices to:
▪ Perceive sensory information
• Sensory Association Cortex
▪ Plan movements
• Motor Association Areas
• Reasoning & Higher Thought
o Most complex analyses occurs in:
▪ Areas of the cortex between sensory perception and movement planning
▪ This includes several areas of the cortex, but…
• Prefrontal Cortex seems to be more (most) specialized for reasoning
o Lesion to the prefrontal (orbitofrontal) cortex:
▪ Patient would show behaviors which indicate lack of consid.
of consequences of actions (no higher planning)
▪ They instead act reflexively (i.e. the simplest function)
o Cerebral hemispheres are specialized for 2 other important functions
▪ Right Parietal Cortex
• Reconstructs spatial coordinates of the body and surroundings so that we
always know our posture relative to our surroundings
• i.e. an internal GPS
▪ Left Hemisphere
• Specialized for language

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SPOKEN LANGUAGE
• Reasoning
o It is difficult to identify the mechanisms of reasoning which lead us to choosing a plan of action
o Therefore, we won’t talk about it much
• Language
o The most advanced function of the nervous system
o Serves as a good example of neural integration
▪ Importance of the language for thinking or reasoning
• Show by the fact that we say, “I can't hear myself think,” when there is too
much noise
o Language includes the ability to express oneself to someone else and to understand someone
else. Hearing a spoken word is a function of the primary auditory cortex in the temporal lobe
o But to understand the word is a function of Wernicke's area in the medial temporal lobe
o Language includes the ability to:
▪ Express oneself to someone else
▪ Understand someone else
• Hearing a spoken word
o Function of the Primary Auditory Cortex in the Temporal Lobe
• Understand the word
o Function of Wernicke's Area in the medial Temporal Lobe
o To answer a question orally:
▪ Neurons in Wernicke's area
…send projections to…
▪ Broca's area in the lateral frontal lobe
…which plans the…
▪ Oral movements
…and then sends the commands to the…
▪ Primary motor cortex
…which will execute the…
▪ Oral movements

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WRITTEN LANGUAGE
• Seeing the written word
o Function of the Primary Visual Cortex in the occipital lobe
• Understand the word
o Function of Wernicke's Area in the medial temporal lobe
• To answer a question orally:
1. Neurons in Wernicke's area
…send projections to…
2. Broca's area in the lateral frontal lobe
…which plans the…
3. Oral movements
…and then sends the commands to the…
4. Primary motor cortex
…which will execute the…
5. Oral movements

BRAILLE LANGUAGE
• Somesthetic Information
o Wernicke's area can integrate this information ON TOP OF integrating auditory and visual
information
o This is used by the blind when reading Braille
• Therefore, Wernicke's area is critical for the integration of the information connected to language.
• Lesions of the following language areas will cause specific deficits:
o Primary Auditory Cortex
▪ Deafness
o Primary Visual Cortex
▪ Blindness
o Wernicke's Area
▪ Cannot understand language whether spoken or written

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▪ Know when to talk based on vision but can’t understand, can’t they form real sentences
• Just babbling nonsense
o Projections From Wernicke's Area To Broca's Area
▪ Cannot repeat words
o Broca's Area
▪ Cannot plan oral movements for speech
▪ Broken, speech that is trying to convey a real meaning
o Primary Motor Cortex
▪ Paralysis of oral movements

SUPPORTING SYSTEMS
• Underlying volitional control are:
1. Memory
2. State of mind, determined by….
i. Reticular Activating System (RAS)
ii. Sleep
iii. Aminergic Projections
iv. The Limbic System
Functions of these 4 parts are very abstract, but, contribute greatly to behavior
• Optimal volitional control depends on:
1. Memory of past events
2. State of mind
i. Activation provided by the RAS
ii. Wakefulness
iii. Mood provided by the aminergic system
iv. Emotions provided by the limbic system

1. MEMORY

Working Memory Circuit

• All neural functions result from…
1. Genetic memory
▪ e.g. Baby can suckle because of the genetically determined connections
2. Acquired memory
▪ Essential to analyze immediate situations using information acquired in the past
▪ Divided into:
1. Short-Term Memory
o Holds information temporarily
▪ e.g. Memory required to follow a conversation
o Information retained here possibly due to reverberating circuits
2. Long-Term Memory
o Holds information for longer than a few week

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Memory Formation
• Information retained in short-term memory is also possibly due to synaptic potentiation
• After tetanus to pathway 1 (strong stimulation representing information to remember)….
o …a stimulus to pathway 1 produces a stronger response
o …but not a stimulus to pathway 2
• Synaptic potentiation can also underlie long term memory.

Memory Consolidation
• Information held in long-term memory → due to anatomical changes in the connections between neurons
o Hippocampus (in the temporal lobe)
▪ Necessary to consolidate declarative (e.g. phone number) short-term memory into long-
term memory
▪ Bilateral lesion of the Hippocampal Area
• Patient still has Declarative Short-Term Memory
…which allows him to carry a conversation with his doctor…
• And he still has long-term memory
…which allows him to speak about things that occurred before the
lesion… (proving that hypothalamus is not the site of memory storage)
• But he cannot generate new long-term memories
…thus can never recognize his new doctor even after having seen him
regularly for more than 20 years after the surgery
• On the other hand, he shows improved performance on an agility test (Non-
Declarative Memory)
...even if he doesn’t remember doing the agility test before
Ex. Test where patient had to trace a star while looking thru mirror,
first day it took him an hour, 2nd day he did it on first shot, but he
never remember having done the test before
o Thus, Motor Systems (like the cerebellum) are responsible for the consolidation of Non-
Declarative Memory (e.g. riding a bicycle) into long-term memory

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2. STATE OF MIND

1. RETICULAR ACTIVATING SYSTEM (RAS) & ACTIVATION

Reticular Activating System (RAS)

• Reticular Activating System
o Sensory Afferents
…send off branches to the…
o Brainstem
…which relay cholinergic projections to…
o Thalamic Nuclei
…projecting to widespread areas of the…
o Cortex
…to provide a…
o Background Level of Activity that lights up the cortex
• Brainstem lesions of the RAS
o Would induce coma because Ascending Projections of the RAS underlies the State Of Arousal
o Occurs with:
1. Sudden onset from trauma, hemorrhage or exogenous toxins (sedatives and alcohol)
and slow onset from progressive tumor or swelling from infection
2. Metabolic abnormalities such as renal failure and diabetic ketoacidosis
o Does not occur following:
1. Cortical infarction
• Ex. Patient falls during infarction of the cerebral cortex but remains awake

Seizures
• Electroencephalogram
o Instrument for measuring the neuronal activity of the brain
o Amplitude of the electroencephalogram
▪ Corresponds to the number of neurons that discharge synchronously
o Frequency
▪ Increases with a more active state of the cortex
• Status Epilepticus
o Prolonged, 30 minutes, or repeated generalized seizure activity without regain of consciousness
in between seizures which may lead to death
o RAS can be recruited by epileptic activity
o Seizures can be partial or generalized to both hemispheres
o 2 generalized seizures that always lead to loss of consciousness
1. Petit Mal (Absence Seizures)
• Patient can maintain a normal posture
• But is unaware of his surroundings because of the disruption of information
processing by the seizure activity
2. Grand Mal (Tonic-Clonic Seizures)
• Electrical activity recruits the RAS which excites the entire cortex
• But the movements are due to invasion of the motor cortex
o To control epilepsy, Anticonvulsants Drugs are used to either:
a) Block sodium and calcium fluxes across neurons,
b) Enhance inhibitory GABA effects, or
c) Inhibit excitatory neurotransmitter release (glutamate and aspartate)

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2. SLEEP & WAKEFULNESS

Sleep Stages
• Sleep
o Circadian Rhythm of 24 hours
▪ Suprachiasmatic Nucleus of the hypothalamus receives inputs from the retina to
function as a biological clock for the sleep-wake cycle
o Vigilance of the cerebral cortex while awake
▪ Relies on impulses from the RAS of the brainstem
▪ Lesion of RAS produces coma
o The ability to sleep
▪ Activated by another part of the brainstem
▪ Lesion of this part of brainstem prevents sleep
o Sleep vs. Coma
▪ Sleep = Partial unconsciousness which can be ended by sensory stimulation

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▪ Coma = Cannot be ended by sensory stimulation even when strong and prolonged
• Coma is not similar to Syncope
o Syncope = Temporary loss of conscience which ends when blood flow
returns to the brain
o When analyzing neurons, after such long bouts of sleeplessness, they don’t have any lack of NTs
or ATP etc.
▪ Not completely sure what causes need for sleep (might be something to do with having
to consolidate memories)
• 2 types of sleep
1. Slow Wave Sleep
▪ Sleep becomes progressively deeper along 4 stages:
1. Relaxation starts
2. EEG becomes irregular
3. Theta and Delta waves appear, vital signs (temperature, breathing, pulse and
pressure) start to drop, skeletal muscles relax
4. EEG is dominated by delta waves
▪ Note that sleepwalking and movements are during slow wave sleep
2. Paradoxical Sleep:
▪ Named “paradoxical” → because cortical activity is almost as in the awake state but the
person is asleep
▪ Paradoxical sleep is the period of REM (Rapid Eye Movements) which is associated with
the period of dreams but the person is easily awakened
• Results from strong cortical activity
• Other muscles are flaccid as if they were paralyzed

Sleep Stages Overnight

• During the night
o Depth of slow wave sleep decreases
o Duration of paradoxical sleep increases
▪ Therefore, the longest dreams are close to the morning

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Sleep Changes With Age

• Stage 4 of slow wave sleep
o Shortens and disappears with age
• Hours and percentage of paradoxical sleep
o Decreases with age
o Forms nearly 50% of sleep in a child
o But forms about 25% of sleep at age 10

3. AMINERGIC PROJECTIONS & MOOD

Biogenic Amines
• Biogenic Amines
o Neurons in the brainstem
o They provide Ascending Projections that set the mood of the cortex
▪ These release Biogenic Amine Neurotransmitters such as Serotonin, Dopamine, And
Norepinephrine
• On widespread regions of cortex
• Rather than the specific point-to-point projections we've been talking about so
far for sensory systems

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Mood Disorders
• Normal behaviour
o There is level of biogenic neurotransmitter release associated with normal behavior
• Depression
o Associated with insufficient release of biogenic amines
o Treatment:
▪ Antidepressants
• Which either block norepinephrine or serotonin uptake at the synapse
▪ Mitochondrial Monoamine Oxidase Inhibitors
• Prevent degradation of the biogenic amines
• Schizophrenia
o Associated with excess release of biogenic amines
o Treatment:
▪ Neuroleptic Drugs
• Block serotonin receptors
• Block dopamine receptors →Will cause signs of Parkinson's disease

4. LIMBIC SYSTEM & EMOTIONS

Emotions
• A comparison of nervous systems across species reveals 3 parts of the nervous system
1. Reptilian Part
▪ Formed of:
• Spinal cord
• Brainstem
• Diencephalon
• Basal ganglia
▪ Functions:
• Automatic control of the internal milieu by the autonomic nervous system
• Automatic control of the external milieu by the involuntary somatic reflexes

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2.
Neomammalian Part
▪ Formed of:
• Cortex
• Subcotical areas (to which the cortex is connected)
▪ Functions:
• Higher mental functions
3. Paleomammalian Part
▪ Formed of:
• Limbic system
o Found between the neomammalian part and the reptilian part
▪ Function:
• Generate emotions
• Emotions
o Limbic System
▪ Emotional behaviors depend largely on this system
▪ This system consists of an area of the Forebrain
• Diencephalon and Surrounding Cortical Structures
o Normal and necessary functions
▪ Ex. Interest, joy, surprise, distress, anger, disgust, contempt, fear, shame, and guilt
▪ Role of emotions
• To destabilize the individual so that he takes an action
o Emotions dictate most of what we choose to do
o Without them we would have no drive
o Ex. Hunger causes the individual to find food
▪ Therefore, we are driven to avoid certain unpleasant
emotions
o Ex. Pleasure causes the individual to repeat the actions which
produced it
▪ Therefore, we are driven to repeat certain pleasant emotions

Limbic System Links Other Systems

• Limbic system can be activated either by the Autonomic or Somatic Nervous System

oCASE: Let’s consider a drop in blood volume due to evaporation of fluids from the body
▪ Partial correction of loss of blood by the hypothalamus (which reduces excretion of
urine and redirects blood to the key organs such as the heart and brain)
Autonomic NS ▪ Stimulation of limbic system to generate the thirst emotion (hypothalamus cannot
correct extreme blood volume loss)
• Thirst emotion is so strong that the cerebral cortex cannot ignore it and must
find water to relieve this emotion

o CASE: Let’s imagine you got a flat tire on the way towards your final exam
▪ Cortex would perceive the situation and activate the limbic system which would
generate a strong emotion of anxiety which would motivate you to use your higher
Somatic NS
mental functions of the cortex cerebral to find a way to attend the exam
• This is activation of the limbic system by the voluntary somatic system
▪ Furthermore, anxiety will act through the autonomic system to produce physiological
Autonomic NS changes such as increased heart rate to give you the energy to work towards a solution
• In extreme cases this can produce signs of stress

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• Therefore, when there is no automatic solution for a stimulus coming from the internal or external milieu,
o There is automatic activation of the limbic system to generate an emotion
o The intensity of the emotion will depend on the stimulus
o Such emotions will motivate us to find a solution by using our higher mental functions in the
cortex cerebral

OUR WORLD IS IN THE CEREBRAL CORTEX

• We live in the world recreated by our cerebral cortex
• All the sensory information (including sensory information derived from our own movement) is
reproduced as best as possible in the cortex
• Hallucinations are errors either in faithfully reproducing the true sensory information or in correctly
interpreting it
• Problems would occur if there were mixing between actual sensory stimuli and memory of previous
sensory stimuli
• If we could put someone to sleep and stimulate the cortical areas in the same way that they are
normally stimulated by touch, light, sound, smell and other stimuli then the person would perceive
the artificial environment that we generate
• Furthermore, if we paralyse their muscles but stimulate the senses as
though the muscles worked then the subject would think that he is
moving within his environment
o e.g. if subject tries to generate elbow flexion, there would be
no movement but you could stimulate the senses the same
way that they would normally be stimulate during elbow
flexion
• This is exactly the plot of the movie Matrix.

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