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Chapter 12
The Somatic Sensory System
.
Sensation and Perception
The function of each sensory system is to
provide the CNS with a representation of
the external world
Sensation -
detection of stimulus and
recognition that the event occurred.
Perception -
interpretation &
appreciation of that event.
Sensory Systems
Neurons in Sensory Systems signal
events by a combination of rate,
spatial and temporal codes.
– Intensity of stimulus - frequency (rate) of APs
– Spatial quality - location of activated neurons
(Braille)
– Temporal quality - music
Sensory Systems:
Chemical Senses (ch. 8):
• Smell (olfactory system)
• Taste (gustatory system)
Sensory Systems:
Auditory and Vestibular Systems (ch. 11)
Sensory Systems:
Visual System (ch. 9 & 10)
Sensory Systems:
Somatic Sensory System (ch. 12)
Somatic Sensation
Enables body to feel, ache, chill
Somatic Sensation
Sensitive to stimuli
Somatic Sensation
Responsible for feeling of touch and pain
Somatic Sensation
Somatic sensory system: Different from other systems in two ways
• Receptors: Distributed throughout the body instead of being concentrated in a small specialized location
• Responds to different kinds of stimuli
Somatic Sensation
We can think of it as a group of at least four senses which are?
touch, temperature, pain, and body position
Somatic Sensation
It is a collective category for all the sensations that are not seeing, hearing, tasting, smelling, and the vestibular sense of balance
Touch
Types and layers of skin
– The two types of skin: Hairy and glabrous (hairless)
– Epidermis (outer layer of skin) and dermis (inner layer of skin)
Touch
Functions of skin
– Protective function
– Prevents evaporation of body fluids
– Provides direct contact with world
Touch
Mechanoreceptors
– Most somatosensory receptors are mechanoreceptors
Touch
Look at figure on page 389
Pacinian Corpuscle
A mechanoreceptor that lies deep within the dermis, selective for high-frequency vibrations.
Rapidly adapting- Tend to respond quickly at first but then stop firing even thugh the stimulus continues
Meissner's Corpuscle
Meissner's corpuscles (or tactile corpuscles) are a type of mechanoreceptor. They are a type of nerve ending in the skin that is responsible for sensitivity to light touch.

-Lower frequencies than Pacinian corpuscle
Rapidly adapting- Tend to respond quickly at first but then stop firing even thugh the stimulus continues
Merkel's disk
the disklike expansion of the end of a nerve fiber together with a closely associated Merkel cell that has a presumed tactile function—called also Merkel's corpuscle
Slowly adapting- Generate a more sustained response during a long stimulus
Ruffini's ending
Found in both hairy and glabrous skin
Slowly adapting- Generate a more sustained response during a long stimulus
Mechanosensitive ion channels
Gating depends on stretching, or changes in tension, of the surrounding membrane
Touch
Look at figures on page 390
Mechanoreceptors
Receptive field size and adaptation rate
Look at figure on 390
Mechanoreceptors
Two-point discrimination- A simple measure of spatial resolution

-spatial resolution- is the minimum distance between two adjacent features

Ex. You can start with a paper clip bent into the shape of a U. Touch on finger, you should have no problem telling that there are two separate points touching your finger. Now bend the wire to bring the points closer together, and touch them to your finger again. When does it feel like one point?
the Importance between fingertips (highest resolution) and elbows

1. There is a much higher density of mechanoreceptors in the skin of the fingertip than on other parts of the body
2. The fingertips are enriched in receptor types that have small receptive fields
3. There is more brain tissure devoted to the sensory information of each square mm of fingertip than elsewhere
4. There may be special neural mechanisms devoted to high-resolution discriminations.

Look at figure on page 392
Primary Afferent Axons- Axons bringing information from the somatic sensory receptors to the spinal cord or brain stem
– Aα, Αβ, Αδ, C (These are in order of decresing sizes)- Axons from skin sensory receptors
– C fibers mediate pain and temperature and is unmyelinated and are the slowest because no myelin
– Aβ mediates touch sensations
Look at figure on page 393
The Spinal cord
Spinal segments (there are 30 of them)- spinal nerves divided into 4 groups in the spinal cord

-4 groups are called, Cervical, Thoracic, Lumbar, and Sacral
Look at figures on page 394, 395, and 396
The Spinal cord
Dermatomes- 1-to-1 correspondence with
segments and dermatomes

Dermatome- Area of the skin innervated by the right and left dorsal roots of a single spinal segment
Touch: The Spinal cord
Sensory Organization of the spinal cord
• Divisions
– Cervical (C)
– Thoracic (T)
– Lumbar (L)
– Sacral (S)
Sensory Organization of the spinal cord
Division of spinal gray matter: Dorsal horn; Intermediate zone; Ventral horn
– Myelinated Aβ axons (touch-sensitive)- enables us to form complex judgements about the stimuli touching the skin
Look at figure on page 397
Dorsal Column–Medial Lemniscal Pathway
– Touch information ascends through dorsal column, dorsal nuclei, medial lemniscus, and ventral posterior nucleus to primary somatosensory cortex

1.Dorsal column- Carry information about tactile sensation toward the brain
2.Dorsal column nuclei- Terminate the axons of the dorsal column
3.Medial lemniscus- Rises through the medulla, pons, and midbrain, and its axons synapse upon neurons of the Ventral Posterior nuclus of the thalamus.
4.Ventral Posterior nuclus- project to specific regions of primary somatosensory cortex
Look at figure on page 398
The Trigeminal Touch Pathway
– Trigeminal nerves
– Cranial nerves
Second order sensory neurons
The neurons that receive sensory input from primary afferents
ipsilaterally
Touch information from the left side of the body is represented in the activity of cells in the left dorsal column nuclei.
Quick fact
No sensory information goes directly into the neocortex without first synapsing in the thalamus
The Trigeminal Touch Pathway
– Trigeminal nerves
– Cranial nerves
Second order sensory neurons
The neurons that receive sensory input from primary afferents
ipsilaterally
Touch information from the left side of the body is represented in the activity of cells in the left dorsal column nuclei.
drug form:
fluid
inhalation, by nose
inhalation, by mouth
inhalation, by neblizer
Somatosensory Cortex
– Brodmann’s Area 3b (or S1): Primary somatosensory cortex

-Primary- area 3b is the PRIMARY somatic sensory cortex because
• Receives dense input from VP nucleus of the thalamus
• Neurons: Responsive to Somatosensory stimuli (not other sensory stimuli)
• Lesions impair somatic sensations
• Electrical stimulation evokes somatic sensory experiences
Look at figures on page 401
Somatosensory Cortex
Other areas
• Postcentral gyrus- Area 3a,1, and 2
Areas 1 and 2- receive dense inputs from area 3b.
-3b-->1 mainly texture information
-3b-->2 Mainly size and shape
• Posterior Parietal
Cortex- Areas 5 and 7
Somatosensory Cortex
Cortical Somatotopy

-Look at figure on page 402
Homunculus- sometimes called a somatotopic map which is the mapping of the body's surface sensations onto a sturcture in the brain
-Map is not always continuous but can be broken up
-Map is not scaled like the human body
Somatosensory Cortex
Cortical Somatotopy
Importance of mouth
– Tactile sensations: Important for speech
– Lips and tongue: Last line of defense when deciding if it is delicious or if it could break your tooth
Somatosensory Cortex
– Cortical Map Plasticity- What happens to the somatotopic map in cortex when an input, such as the finger, is removed?
Look at figure on page 403
Somatosensory Cortex
– Remove digits or overstimulate – examine
somatotopy before and after
• Conclusions of experiments
– Reorganization of cortical maps
» Dynamic
» Adjust depending on the amount of sensory
experience
Somatosensory Cortex
The Posterior Parietal Cortex
• Involved in perception, interpretation of
spatial relationship and movement planning
Somatosensory Cortex
Damage to The Posterior Parietal Cortex
• Agnosia- Inability to recognize objects even though simple sensory skills seem to be normal
Somatosensory Cortex
Damage to The Posterior Parietal Cortex
• Astereoagnosia- Cannot recognize common objects by feeling them
Somatosensory Cortex
Damage to The Posterior Parietal Cortex
• Neglect syndrome- Part of the body or a part of the world (the entire visual field left of the center of gaze) is ignored or suppressed, and its very existance is denied.
Pain- Dont study tooo much on this portion till slide 73
Nociceptors- Besides mechanoreceptors, somatic sensation depends strongly on nociceptors, the free, branching, unmyelinated nerve endings that signal that body tissue is being damaged or is at risk of being damaged.

-The membrans of nociceptors contain ion channels that are activated by these types of stimuli

-Normally respond only when stimuli are strong enough to damage tissue
Pain
Pain and nociception are not the same thing
– Pain - feeling of sore, aching, throbbing
– Nociception - sensory process that provides the
signals that trigger pain
Pain
Nociception and the Transduction of Painful Stimuli
Types of Nociceptors

-Polymodal nocireceptors- The majority of nociceptors respond to mechanical, thermal, and chemical stimuli and are therefore called polymodal nociceptors

-Mechanical nocireceptors- Showing selective responses to strong pressure

-Thermal nocireceptors- Showing selective responses to burning heat or extreme cold.
Pain
Nociception and the Transduction of Painful Stimuli
Hyperalgeia - Our body's ability to control its own pain
• Primary (occurs within the area of damaged tissue) and secondary hyperalgesia (Tissues surrounding a damaged area may become supersensitive by the process of secondary hyperalgesia)
• Bradykinin (binds to specific receptor molecules that activate ionic conductances in some nociceptors), prostaglandins, and substance P (these three makes it more sensitive to thermal or mechanical stimuli
Pain
Primary Afferents and Spinal mechanisms
– First pain (fast and sharp) and second pain (duller, longer-lasting)
– Referred pain: Angina

Look at figure on page 412
IMMUNOLOGY
study of the molecules, cells, organs,
and systems responsible for the
recognition of foreign material
Ascending Pain Pathways
Spinothalamic Pain Pathway
-Information about pain in the body is conveyed from the spinal cord to the brain by the Spinothalamic Pain Pathway
Brown-Séquard Syndrome- The constellation of sensory and motor signs following damage to one side of the spinal cord is called Brown-Séquard Syndrome
Ascending Pain Pathways
The Trigeminal Pain Pathway
Pain information from the face and head takes a path to the thalamus that is analogous to the spinal path
Ascending Pain Pathways
The Thalamus and the Cortex
• Touch and pain systems remain segregated
• Pain and temperature information sent to various
cortical areas
Ascending Pain Pathways
Look at figure on page 414
Ascending Pain Pathways
Look at figure on page 415
The Regulation of Pain
– Afferent Regulation- Pain evoked by activity in nociceptors can also be reduced , for ex. rubbing on shin when you bruise it.
– Descending Regulation- pain suppression (feel no pain)
– The endogenuos opiates
• Opioids and endorphins- act by binding tightly and specifically to several types of opioid receptors in the brain and that the brain itself manufactures endogenous morphin-like substances, collectivly called endorphines.
Look at figure on page 417
The Regulation of Pain
– Descending regulation
– PAG = pariaqueductal gray matter of the midbrain
– PAG can influnce the raphe nuclei of the medulla- depress the activity of nociceptive neurons.
Thermoreceptors
– “Hot” and “cold” receptors
– Varying sensitivities
Look at figure on page 419
Thermoreceptors
Hot and cold receptors
Look at figure on page 420
The Temperature Pathway
– Organization of temperature pathway
• Identical to pain pathway
– Cold receptors coupled to Aδ and C
– Hot receptors coupled to C
Concluding Remarks
• Sensory systems exhibit similar organization
and function
• Sensory types are segregated within the
spinal cord and cerebral cortex
• Repeated theme
– Parallel processing of information
• Perception of object involves the seamless
coordination of somatic sensory information
Dont study too much up to here, next ones studyy!!!
Lecture 9: Cortical and Descending Motor Systems.
Chapter 13 & 14
We will turn our attention to the system that actually gives rise to behavior
Look at slide 2!!
helpful
Motor Programs
Motor system: All of our Muscles and neurons that control muscles
Motor Programs
Role: Generation of coordinated movements
Motor Programs
Motor control can be divided into two parts which are:
• Spinal cord's command and control of coordinated muscle contraction
• Brain's control and command motor programs in spinal cord
The Somatic Motor System
Types of Muscles
Two types of Striated muscle:
• skeletal (bulk of
body muscle mass) and functions to move bones around joints
• Cardiac (heart)
The Somatic Motor System
Types of Muscles
Smooth muscle lines the digestive tract, arteries, and related structures and is innervated by nerve fibers from the autonomic nervous system
-Plays a role in peristalsis (movement of material through the intestines) and the control of blood pressure and blood flow.
The Somatic Motor System
Look at figures on page 425
Somatic Musculature
– Axial muscles are responsible for movement of the Trunk
– Proximal muscles move Shoulder, elbow, pelvis,
knee movement
– Distal muscles move Hands, feet, digits (fingers and
toes) movement
The Lower Motor Neuron
– Lower motor neuron: Innervated (stimulate through nerves.) by ventral horn of
spinal cord
Look at figure on page 427
The Lower Motor Neuron
– Upper motor neuron: Supplies input to the spinal cord
Somatic musculature is innervated (stimulate through nerves.) by somatic motor neurons
.
The Motor Unit- One alpha motor neuron and all the muscle fibers it innervates collectively make up the elementary component of motor control called the motor unit
Look figure on page 429
Two lower motor neurons
• Alpha neurons- directly trigger the generation of force by muscles
• Gamma neurons
-Muscle contraction results from the individual and combined actions of those motor units
.
Motor neuron pool
collection of alpha motor neurons the innervates a single muscle.
Graded Control of Muscle Contraction by Alpha Motor Neurons
– Varying firing rate of motor neurons
Graded Control of Muscle Contraction by Alpha Motor Neurons
– Recruit additional synergistic motor units
-depends on how many muscle fibers are in that unit
- motor units are recruited in the order of smallest first, largest last.
Inputs to Alpha Motor Neurons
-Has three sources of input
• ALS (Amyotrophic Lateral Sclerosis) = Lou Gehrig’s Desease
– Selective degeneration of alpha motor neurons

3 sources:
1. The dorsal root ganglion cells with axons that innervate a specialized sensory apparatus embedded within the muscle known as a muscle spindle.
2. Derives from upper motor neurons in the motor cortex and brain stem
3.Is the largest input to an alpha motor neuron derives from interneurons in the spinal cord
Look at figure on page 430
Neuromuscular Matchmaking
• Alternate nerve input
– Switch in muscle phenotype (physical
characteristics) as a consequence of increased or
decreased activity.
Neuromuscular Matchmaking
Increased in activity leads to Hypertrophy: Exaggerated growth of muscle fibers as seen in body builders
Neuromuscular Matchmaking
Prolonged inactivity leads to Atrophy: Degeneration of muscle fibers
ex. Joints in a cast is immobilized
Excitation-Contraction Coupling
Muscle contraction
Alpha motor neurons release Ach
• Innervate muscle fibers
Excitation-Contraction Coupling
Muscle contraction
ACh produces large EPSP in muscle fibers (via nicotinic Ach
receptors
Excitation-Contraction Coupling
Muscle contraction
EPSP evokes action potential
Excitation-Contraction Coupling
Muscle contraction
Action potential (excitation) triggers Ca2+ release, leads to
fiber contraction
Excitation-Contraction Coupling
Muscle contraction
Relaxation, Ca2+ levels lowered by organelle reuptake
Muscle contraction is initiated by the release of ACh (actylcoline) from the axon terminals of alpha motor neurons
.
Excitation-Contraction Coupling
• Myasthenia Gravis
– NMJ Disease
Excitation-Contraction Coupling
• Myasthenia Gravis
– autoimmune disease; nicotinic ACh Receptors dysfunction
Excitation-Contraction Coupling
• Myasthenia Gravis
– Drugs: AChE blockers
Muscle Fiber
Structure
Look at figure on page 434 &435
The Molecular Basis of Muscle Contraction
Z lines: Division of myofibril into segments by disks
The Molecular Basis of Muscle Contraction
Sarcomere: segment comprised of Two Z lines and myofibril in between
The Molecular Basis of Muscle Contraction
Thin filaments: Series of bristles- on each side of the Z lines
The Molecular Basis of Muscle Contraction
Thick filaments: Between and among thin filaments
The Molecular Basis of Muscle Contraction
Sliding-filament model:
• Binding of Ca2+ to troponin causes myosin to bind to
actin
• Myosin heads pivot, cause filaments to slide
Part of the muscle fiber- Sarcolemma
Muscle fibers are enclosed by an excitable cell membrane called the Sarcolemma
Part of the muscle fiber- Myofibrils
Within the muscle fiber are a number of cylindrical structures called Myofibrils
Part of the muscle fiber- Sarcoplamic
Surrounds the myofibrils and is an extensive intracellular sac that stores Ca
Part of the muscle fiber- T tubules
Action potentials sweeping along the sarcolemma gain access to the sarcoplamic reticulum deep inside the fiber by way of netweork of tunnels called T tubules
Summarization of EXCITATION of the excitation-contraction coupling
1. An action potential occurs in an alpha motor neuron axon
Summarization of EXCITATION of the excitation-contraction coupling
2. ACh is released by the axon terminal of the alphamotor neuron at the neuromuscular junction
Summarization of EXCITATION of the excitation-contraction coupling
3. Nicotinic receptor channels in the sarcolemma open, and the postsynaptic sarcolemma depolarizzes (EPSP)
Summarization of EXCITATION of the excitation-contraction coupling
4. Voltage-gated sodium channels open, and an action potential is generated in the muscle fiber, which sweeps down the sarcolemma and into the T tubules
Summarization of EXCITATION of the excitation-contraction coupling
5. Depolarization of the T tubules causes Ca release from the sarcoplasmic reticulum
Summarization of CONTRACTION of the excitation-contraction coupling
1. Ca binds to troponin
Look at figure on page 436
Summarization of CONTRACTION of the excitation-contraction coupling
2. Myosin binding sites on actin are exposed
Summarization of CONTRACTION of the excitation-contraction coupling
3. Myosin heads bind actin
Summarization of CONTRACTION of the excitation-contraction coupling
4. Myosin heads pivot
Summarization of CONTRACTION of the excitation-contraction coupling
5. Myosin heads disengage at the expense of ATP
Summarization of CONTRACTION of the excitation-contraction coupling
6. The cycle continues as long as CA and ATP are present
Summarization of RELAXATION of the excitation-contraction coupling
1. As EPSPs end, the sarcolemma and T tubules return to their resting potentials
Summarization of RELAXATION of the excitation-contraction coupling
2. Ca is sequestered by the sarcoplasmic reticulum by an ATP-driven pump
Summarization of RELAXATION of the excitation-contraction coupling
3. Myosin binding sites on actin are covered by troponin
• Steps in Excitation-
Contraction Coupling
– Excitation: Action potential, ACh release,
EPSP, action potential in muscle fiber,
depolarization
– Contraction: Ca2+, myosin binds actin, myosin
pivots and disengages, cycle continues until
Ca2+ and ATP present
– Relaxation: EPSP end, resting potential, Ca2+ by
ATP driven pump, myosin binding actin covered
Spinal Control of Motor Units
First source: Sensory feedback from muscle
Look at figure on page 438
The Myotatic Reflex
Stretch reflex: Muscle pulled--> tendency to pull back
The Myotatic Reflex
Feedback loop-
ex. When a weight is placed on a muscle and the muscle starts to lengthen, the muscle spindles are streched. The stretching of the spindle leads to depolarization of the Ia axon endings due to the opening of mechanosensitive ion channels. Therefore it synaptically depolarizaes the alpha motor neurons, which respond by incresing their action potential frequency. This causes the muscle to contract, thereby shortening it.
Principles of the feedback control systems are that a set point is determined (in this case, the desired muscle length), deviations from the set point are detected by a sensor (the Ia axon endings), and deviations are compensated for by an effector system (alpha motor neurons and extrafusal muscle fibers), returning the system to the set point of the myotatic feedback loop
The Myotatic Reflex
Discharge rate of sensory axons: Related to muscle length
-As the muscle is streched, the discharge rate goes up
-As the muscle is shortened and goes slack, the discharge rate goes down.
The Myotatic Reflex
Monosynaptic (monosynaptic myotatic reflex arc)- because only one synapse separates the primary sensory input from the motor neuron output.
Example: knee-jerk reflex
The Myotatic Reflex
Look at figure on page 441
Gamma Motor Neurons
Muscle spindle- contains modified skeletal muscle fibers within its fiberous capsule
Intrafusal fibers: gamma- modified skeletal muscle fibers within its fiberous capsule
Gamma Motor Neurons
Muscle spindle
Extrafusal fibers: alpha -
more numerous than intrafusal fibers and lie outside the spindle and form the bulk of the muscle
An important difference between extrafusal and intrafusal fibers is that..
-Extrafusal fibers are innervated by alpha motor neurons
-Intrafusal fibers receive their motor innervation by gamma motor neurons.
An important difference between extrafusal and intrafusal fibers is that..
-Alpha and Gamma motor neurons have the opposite effects on Ia axon output.
-Alpha activation alone decreases Ia activity, while gamma activation alone increases Ia activity
What happens when there is an Activation of alpha motor neurons?
It causes the extrafusal muscle fibers to shorten.
What happens when there is an Activation of Gamma motor neurons?
It causes the poles of the spindle to contract, keeping it "on-the-air"
Gamma Loop
Changing the activity of the gamma motor neurons changes the set point of the myotatic feedback loop
Gamma Loop
Provides additional control of alpha motor neurons and muscle contraction
Gamma Loop
Circuit
• Gamma motor neuron--> intrafusal muscle fiber
--> Ia afferent axon --> alpha motor neuron --> extrafusal muscle fibers
Proprioception from Golgi Tendon Organs
Reverse myotatic reflex function: Regulate muscle tension within optimal range

-Ib axons enter the spinal cord, branch repeatedly, and synapse on interneurons in the ventral horn. Some of these interneurons form inhibitory connections with alpha motor neurons innervating the same muscle.

-This reflex arc protects the muscle from being overloaded

-As muscle tension increases, the inhibition of the alpha motor neuron slows muscle contraction; as muscle tension falls, the inhibition of the alpha motor neuron is reduced, and muscle contraction increases.
Look at figure on page 443
Proprioception
Awareness of the position of one's body.
Golgi Tendon Organs
-Monitors force of contraction and muscle tension
Location: junction of the muscle and the tendon and are innervated by group Ib sensory axons, which are slightly smaller than the Ia axons innervating the muscle spindles
Important
-Spindle fibers are situated in parallel with muscle fibers
-golgi tendon organs are situated in series
--This is what distinguishes the types of information these two sensors provide the spinal cord: Ia activity from the spindle encodes muscle length information, while Ib activity from the golgi tendon organ encodes muscle tension information.
Proprioception from the joints
Proprioceptive axons in connective joint tissues
Proprioception from the joints
Respond to angle, direction and velocity of movement in a joint
Spinal Interneurons
Synaptic inputs
Actions of Ib inputs from golgi tendon organs on alpha motor neurons are entirely polysynaptic

Spinal interneurons receive synaptic input from
1.Primary sensory axons 2.Descending axons from brain
3.Collaterals of lower motor neuron axons
Inhibitory Input
Reciprocal inhibition: Contraction of one muscle set accompanied by relaxation of antagonist muscle, the extensors
Example: Myotatic reflex

Look at figure on page 445
The Generation of Spinal Motor
Programs for Walking
Central pattern generators
-Circuits that give rise to rhythmic motor activity
Look at figure on page 448
Spinal control of movement
Different levels of analysis
Spinal control of movement
Sensation and movement linked
Direct feedback
- The normal function of the alpha motor neuron depends on direct feedback from the muscles themselves and indirect information from the tendons, joints, and skin.
Spinal control of movement
Spinal cord contains an Intricate network of circuits for the control of movement
Brain Control of Movement
Chapter 14
.
The brain influences activity of the spinal cord for what?
Voluntary movements
Hierarchy of controls
– Highest level: Strategy
-Represented by the association areas of neocortex and basal ganglia of the forebrain: the goal of the movement and the movement strategy that best achieves the goal

– Middle level: Tactics
-Represented by the motor cortex and cerebellum: the sequences of muscle contractions, arranged in space and time, required to smoothly and accurately achieve the strategic goal.

– Lowest level: Execution
-Represented by the brain stem and spinal cord: activation of the motor neuron and interneuron pools that generate the goal-directed movement and make any necessary adjustments of posture
Sensorimotor system
Sensory information: Used by motor system
-The proper functioning of each level of the motor control hierarchy relies so heavily on sensory information that the motor system of the brain might properly be considered a sensorimotor system.

-At the highest level, sensory information generates a mental image of the body and its relationship to the environment

-At the middle level, tachtical decisions are based on the memory of sensory information from past movements.

-At the lowest level, sensory feedback is used to maintain posture, muscle length, and tension before and after each voluntary movement.
Principles of Motor Systems organization:
A. There is a hierarchy of motor subsystems.
Principles of Motor Systems organization:
B. Whenever possible a subsystems of the hierarchy
accomplishes movements through control of automatic and reflex behaviors organized at lower levels.
Principles of Motor Systems organization:
C. Almost all components of the motor system contain somatotopic maps.
Principles of Motor Systems organization:
D. Each subsystems receives somatosensory info.
Principles of Motor Systems organization:
E. Movements can be accomplished by means of the spinal cord (or brain stem) lower motor neurons, which innervate muscles.
Principles of Motor Systems organization:
F. The lower motor neurons may be reflexly activated at the
segmental level or may be voluntarily activated by a higher
subsystem in the hierarchy
Principles of Motor Systems organization:
G. The cerebellum and basal ganglia have some control over the subsystems of the motor hierarchy but not project to the
lower motor neurons.
Look at figure on page 455
Descending Spinal Tracts
Axons from brain descend along two major pathways:
Lateral Pathways
• Involved in voluntary movement of the distal musculature and are under direct cortical control
Descending Spinal Tracts
Axons from brain descend along two major pathways:
Ventromedial Pathways
• Control of posture and locomotion are under brain stem control
The Lateral Pathways
Voluntary movement --> under direct cortical control
Look at figure on page 453
The Lateral Pathways
Components
• Corticospinal tract- most important component
-Originating in the neocortex
-regulate the flow of somatosensory information to the brain

• Pyramidal tract
-At the junction of the medulla and spinal cord, the pyramidal tract crosses, this means that the right motor cortex directly commands the movement of the left side of the body and the left motor cortex controls the muscles on the right side.

• Rubrospinal tract
-originates in the "red nucleus" of the midbrain, named for its distinctive pinkish hue
-A major source of input to the red nucleus is the very region of frontal cortex that also contributes to the corticospinal tract.
The Lateral Pathways
The Effects of Lateral Pathway Lesions
Partial lesions in corticospinal and rubrospinal tracts
– Fractionated movement of arms and hands; they could not move their shoulders, elbows, wrists, and fingers independently
The Lateral Pathways
The Effects of Lateral Pathway Lesions
Damage of corticospinal tract
» Paralysis on contralateral side
The Ventromedial Pathways
Posture and locomotion --> under brain stem control
The Ventromedial Pathways use sensory information about balance, body position, and the visual environment to reflexively maintain balance and body posture
The Ventromedial Pathways
The Vestibulospinal tract
-Keeps head balanced on the shoulders as the body moves, and turn the head in response to new sensory stimuli.

-Originate in the vestibular nuclei of the medulla, which relay sensory information from the vestibular labyrinth in the inner ear. The vestibular labyrinth consists of fluid-filled canals and cavities in the temporal bone that are closely associated with the cochlea.

-Proojects bilaterally down the spinal cord and activates the cervial spinal circuits that control the neck and back muscles
The Ventromedial Pathways
The Tectospinal tract
-Originates in the superior colliculus of the midbrain, which receives direct input from the retina
The Ventromedial Pathways
The Pontine and Medullary Recticulospinal tract.

-The Pontine Recticulospinal tract enhances the antigravity reflexes of the spinal cord

-Helps maintain gravity by facilitating the extensors of the lower limbs

-The Medullary Recticulospinal tract has the opposite effect; it liberates the antigravity muscles from reflex control

-Activity in both reticulospinal tracts is controlled by descending signals from the cortex
The Ventromedial Pathways
Contains four descending tracts that originate in the brain stem and terminate among the spinal interneurons controlling proximal and axial muscles

The four tracts are?
1. Vestibulospinal tract,
2. Tectospinal tract,
3. The pontine reticulospinal tract,
4. The Medullary reticulospinal tract.
The Planning of Movement by
the Cerebral Cortex
Motor Cortex
– Area 4 and area 6 of the frontal lobe is called the motor cortex
-The control of voluntary movement engages almost all of the neocortex
Look at figure on page 460
What is Area 4 now referred to?
Primary motor cortex or M1
Somatotopic map of primary motor Cortex
Compare the Somatosensory Cortex and Motor Cortex
Figures on page 460 and page 402
Motor Cortex (Penfield)
Area 4 = “Primary motor cortex” or “M1”
Motor Cortex (Penfield)
Area 6 = “Higher motor area” (Penfield)
Lateral region --> Premotor area (PMA)
-The PMA connects primarily with reticulospinal neurons that innervate proximal motor units
Motor Cortex (Penfield)
Area 6 = “Higher motor area” (Penfield)
Medial region --> Supplementary motor area (SMA)
-The SMA sends axons that innervate distal motor units directly
Motor Cortex (Penfield)
Area 6 = “Higher motor area” (Penfield)
Motor maps in PMA and SMA
– Similar functions; different groups of muscles innervated
The Contributions of Posterior Parietal and Prefrontal Cortex
Represent highest levels of motor control
Decisions made about actions and their outcome
The Contributions of Posterior Parietal and Prefrontal Cortex
Area 5:
-Target of inputs from the primary somatosensory cortical areas 3,1,and 2
The Contributions of Posterior Parietal and Prefrontal Cortex
Area 7: Inputs from higher-order visual cortical areas such as MT
The Contributions of Posterior Parietal and Prefrontal Cortex
The parietal lobes are extensively interconnected with regions in the Anterior frontal lobes:
-Anterior frontal lobes (prefrontal): Abstract thought, decision making and anticipating consequences of action
These "prefrontal areas" along with the posterior parietal cortex, represent the highest levels of the motor control hierarchy, where decisions are made about what actions to take and their likely outcome.
The Contributions of Posterior Parietal and Prefrontal Cortex
-Prefrontal and parietal cortex both send axons that converge on cortical area 6

Area 6: Actions converted into signals specifying how actions will be performed
The Contributions of Posterior Parietal and Prefrontal Cortex
Per Roland --> Monitored cortical activation
accompanying voluntary movement by (PET)
• Results supported view of higher order motor planning
Neuronal Correlates of Motor Planning
Evarts: Recorded activity in motor areas of awake, behaving animals
Demonstrated importance of area 6 in planning movement (Where SMA and PMA are located)
Neuronal Correlates of Motor Planning
Evarts: Recorded activity in motor areas of awake, behaving animals
(ready,set, go!)
“ready”- readiness depends on activity in the Parietal and frontal lobes
Neuronal Correlates of Motor Planning
Evarts: Recorded activity in motor areas of awake, behaving animals
(ready,set, go!)
“set”- Supplementary and premotor areas (SMA and PMA)
-Where movement strategies are devised and held until they are executed
Neuronal Correlates of Motor Planning
Evarts: Recorded activity in motor areas of awake, behaving animals
(ready,set, go!)
“go”- Area 6
The Basal Ganglia
The major subcortical input to area 6 arises in a nucleus of the dorsal thalamus, called the ventral lateral (VLo) nucleus

-The input to this part of VLo, arises from the basal ganglia buried deep within the telencephalon.
Look at figure on page 465
-important both figures
The Basal Ganglia
Cortex
Projects back to basal ganglia
The Basal Ganglia
Cortex
Forms a “loop”
The Basal Ganglia
Function of the loop: Selection and initiation of willed movements
Anatomy of the Basal Ganglia
Caudate nucleus, putamen, globus pallidus, subthalamic nucleus

-Caudate and putamen together are called the striatum, which is the target of the cortical input to the basal ganglia.
-The globus pallidus is the source of the output to the thalamus.
-The other structures participate in various side loops that modulate the direct path:
Cortex-->striatum-->globus pallidus-->VLo--> cortex(SMA)
Anatomy of the Basal Ganglia
Substantia nigra: Connected to basal ganglia
The Motor Loop: Selection and initiation of willed movements
Origin of the most direct path in the motor loop through the basal ganglia originates with an Excitatory connection from the cortex to cells in putamen
Cortical activation of the putamen is excitation of the SMA by VL
The Motor Loop: Selection and initiation of willed movements
Cortical activation
1.Excites putamen neurons
The Motor Loop: Selection and initiation of willed movements
Cortical activation
2. Inhibits globus pallidus neurons
The Motor Loop: Selection and initiation of willed movements
Cortical activation
3. Release cells in VLo from inhibition
The Motor Loop: Selection and initiation of willed movements
Activity in VLo boosts the activity in SMA
The Motor Loop
Basal Ganglia Disorders:
Hypokinesia (increased inhibition of the thalamus by the basal ganglia, little movement) and hyperkinesia (a decreased in basal ganglia output leads to hyperkinesia, an excess of movement)
The Motor Loop
Basal Ganglia Disorders:
Parkinson’s disease
(characterized by hypokinesia)
Symptoms: Bradykinesia (slowness of movements), akinesia (difficulty in initiating willed movements), rigidity (increased muscle tone) and
tremors of hand and jaw
The Motor Loop
Basal Ganglia Disorders:
Parkinson’s disease
(characterized by hypokinesia)
Organic basis: Degeneration of substantia nigra inputs to striatum
The Motor Loop
Basal Ganglia Disorders:
Parkinson’s disease
(characterized by hypokinesia)
Dopamine treatment: Facilitates production of dopamine
to increase SMA activity
The Motor Loop
Basal Ganglia Disorders:
Huntington’s disease
(characterized by hyperkinesia)
Symptoms: Hyperkinesia, dyskinesia, dementia,
impaired cognitive disability, personality disorder
The Motor Loop
Basal Ganglia Disorders:
Hemiballismus (ballism
(characterized by hyperkinesia)
Violent, flinging movement on one side of the body
Initiation of Movement by the Primary Motor Cortex
Electrical stimulation of area 4
Contraction of small group of muscles
Initiation of Movement by the Primary Motor Cortex
The Input-Output Organization of M1
Betz cells: Pyramidal cells in cortical layer 5 (V)
Initiation of Movement by the Primary Motor Cortex
The Input-Output Organization of M1
Two sources of input to Betz cells
• Cortical areas
• Thalamus
Initiation of Movement by the Primary Motor Cortex
The Coding of Movement in M1
Activity from several neurons in M1 encodes force and direction of movement
Initiation of Movement by the Primary Motor Cortex
The Coding of Movement in M1
Look at figure on page 470
Initiation of Movement by the Primary Motor Cortex
The Coding of Movement in M1
-There are 3 important conclusions about how M1 commands voluntary movement which are,
1. Motor cortex: Active for every movement
Initiation of Movement by the Primary Motor Cortex
The Coding of Movement in M1
--There are 3 important conclusions about how M1 commands voluntary movement which are,
2. Activity of each cell: Represents a single “vote”
Initiation of Movement by the Primary Motor Cortex
The Coding of Movement in M1
--There are 3 important conclusions about how M1 commands voluntary movement which are,
3. Direction of movement: Determined by a tally (and averaging)
The Coding of Movement in M1(
The Malleable Motor Map
Experimental evidence from rats
– Microstimulation of M1 cortex normally elicits
whisker movement--> cut nerve that supplies
whisker muscles--> Microstimulation now causes
forelimb movement
The Coding of Movement in M1(
The Malleable Motor Map
Decoding M1 activity
– Helps patients with severe damage to their motor
pathways
The Coding of Movement in M1(
The Malleable Motor Map
The larger the population of neurons representing a type of movement, the finer the possible control
The Cerebellum
Function: Sequence of muscle contractions
Ataxia
• Uncoordinated and inaccurate movements
The Cerebellum
Function: Sequence of muscle contractions
Ataxia
• Caused by cerebellar lesions
The Cerebellum
Function: Sequence of muscle contractions
Ataxia
• Symptoms
– Dysynergia (decomposition of synergistic multijoint movement), dysmetric
The Cerebellum
The Motor Loop Through the Lateral Cerebellum
-- Pontine nuclei
• Axons from layer V pyramidal cells in the sensorimotor
cortex form massive projections to pons
The Cerebellum
The Motor Loop Through the Lateral Cerebellum
– Corticopontocerebellar projection
• 20 times larger than pyramidal tract
The Cerebellum
The Motor Loop Through the Lateral Cerebellum
– Function
• Execution of planned, voluntary, multijoint movements
The Cerebellum
The Motor Loop Through the Lateral Cerebellum
Look at figure on page 476
Concluding Remarks
Example of the baseball pitcher
Walking: Ventromedial pathways
Concluding Remarks
Example of the baseball pitcher
Ready to pitch
• Neocortex, ventromedial pathways
Concluding Remarks
Example of the baseball pitcher
Pitch signs and strategy
• Sensory information engages parietal and prefrontal cortex and area 6
Concluding Remarks
Example of the baseball pitcher
– Winds and throws
• Increased basal ganglia activity (initiation)
Concluding Remarks
Example of the baseball pitcher
– Winds and throws
• SMA activity --> M1 activation
Concluding Remarks
Example of the baseball pitcher
– Winds and throws
• Corticopontocerebellar pathways --> Cerebellum
Concluding Remarks
Example of the baseball pitcher
– Winds and throws
Cortical input to reticular formation --> Release of
antigravity muscles
Concluding Remarks
Example of the baseball pitcher
– Winds and throws
Lateral pathway --> engages motor neurons --> action
Look at concluding remarks in chapter 14
IMPORTANT SUMMARY
Chapter 9
The eye
.
Overview of the Visual System
Visual information is projected the following pathway:
• Retina
• Optic nerve
• Optic chiasm
• Optic tract
• Lateral geniculate nucleus (LGN)
• Visual radiations
• Primary visual cortex
• Higher visual cortex
Overview of the Visual System
The visual world is represented throughout the visual pathway in a map that basically is “upside down and backwards”
Retina
A thin layer of cells at the back of the eye that transduces light energy into neural activity by photoreceptors
-specialized to detect differences in the intensity of light falling on different parts of it.
• Photoreceptors: Converts light energy into
neural activity
• Detects differences in intensity of light
Optic nerve
The bundle of ganglion cell axons that passes from the eye to the optic chiasm.
-distributes visual information in the form of action potentials.
Optic chiasm
The structure in which the right and left optic nerves converge and partially decussate (cross) to form the optic tracts.
Optic tract
A collection of retinal ganglion cell axons stretching from the optic chiasm to the brain stem.
Important targets of the optic tract are the lateral geniculate nucleus and superior colliculus.
Lateral geniculate nucleus (LGN)
A thalamic nucleus that relays information from the retina to
the primary visual cortex.
• First synaptic relay in the primary visual pathway
• Visual information ascends to cortex --> interpreted and remembered
Visual radiations
a charted band of wavelengths of electromagnetic radiation obtained by refraction or diffraction.
Primary visual cortex
Brodmann’s area 17, located
at the pole of the occipital lobe; also called striate
cortex and V1.
Significance of vision
Relationship between human eye & camera

-Like the camera, the eye automatically adjusts to differences in illumination and automatically focuses itself on objects of interest.
Properties of Light
Look at figure on page 279
Light
Light and electromagnetic radiation (wave of energy) - it is all around us

-Light is the electromagnetic radiation that is visible to our eyes.
Light
Electromagnetic radiation
• Wavelength, frequency, amplitude
Hot colors: Orange, red;
Cool colors: blue, violet
Optics
Study of light rays and their
interactions
Reflection
– Bouncing of light rays off a
surface
Optics
Study of light rays and their
interactions
Absorption
– Transfer of light energy to a
particle or surface
Optics
Study of light rays and their
interactions
Refraction
– Bending of light rays from one
medium to another
-Images are formed in the eye by refraction
-reflection and absorption determine what light enters the eye
Optics
Study of light rays and their
interactions
Look at figure on page 280
The Structure of the Eye
Gross Anatomy of the Eye
Pupil: Opening where light enters the eye and reach the retina
The Structure of the Eye
Gross Anatomy of the Eye
Sclera: "White of the eye" which forms the tough wall of the eyeball
The Structure of the Eye
Gross Anatomy of the Eye
Iris: Gives color to eyes surrounds the pupil
The Structure of the Eye
Gross Anatomy of the Eye
Cornea: Glassy transparent external surface of the eye covers the pupil and iris
The Structure of the Eye
Gross Anatomy of the Eye
Optic nerve: Carries Bundles of axons from the retina and reaches the base of the brain near the pituitary gland
The Structure of the Eye
Gross Anatomy of the Eye
Look at figure on page 280
The Structure of the Eye
Ophthalmoscopic Appearance of the Eye
-Optic disk-retinal vessels originate from a pale circular region called the optic disk, where the optic nerve fibers exit the retina
Look at figure on page 281
Cross-Sectional Anatomy of the Eye
Aqueous Humor: Because the cornea lacks blood vessels it is nourished by the fluid behind it, the aqueous humor
Cross-Sectional Anatomy of the Eye
Ciliary muscles: Ligaments that suspend lens
Cross-Sectional Anatomy of the Eye
Lens: Change shape to adjust focus
Divides eyes into two compartments
– Aqueous humor in anterior chamber- watery fluid that lies between the cornea and the lens
– Jelly-like vitreous humor in posterior chamber- Lies between the lens and the retina; its pressure serves to keep the eyeball spherical
Image Formation by the Eye
Introduction
– Eye collects light reflected off objects in the environment, focuses onto retina to form images
-Bringing objects into focus involves the combined refractive powers of the cornea and lens
Image Formation by the Eye
Refraction of light by the cornea
-The cornea, rather than the lens, is the site of most of the refractive power of the eyes.
Look at figures on page 284
Image Formation by the Eye
Focal distance = refractive surface to the point where parallel light rays converge
-focal distance depends on the curvature of the cornea
-----The tighter the curve, the shorter the focal distance
Image Formation by the Eye
Accommodation by the Lens
Changing shape of lens allows for extra focusing power a process called accommodation
The Pupillary Light Reflex
The pupillary light reflex involves Connections between retina and brain stem neurons that control the muscles that constirct the pupils
The Pupillary Light Reflex
Continuously adjusting to different ambient light levels
The Pupillary Light Reflex
Consensual- Shining light into one eye causes the constriction of the pupils of both eyes
The Pupillary Light Reflex
Pupil similar to the aperture (hole) of a camera
-Constriction of the pupil has the effect of increasing the depth of focus, just like decreasing the aperture size on a camera lens
The Visual Field
Amount of space viewed by the
retina when the eye is fixated
straight ahead
-space in front of you without moving head just eyes.
Visual Acuity
– Ability to distinguish two
nearby points
look at figure on page 288, 9.10
Visual Acuity
– Visual Angle: Distances across
the retina described in degrees
Microscopic Anatomy of the Retina
Photoreceptors:
– Cells that convert light energy into neural activity

Look at figures on page 289
Microscopic Anatomy of the Retina
Direct (vertical) pathway:
-visual information to exit the eye
Photoreceptors--> bipolar cells--> ganglion cells
Microscopic Anatomy of the Retina
Direct (vertical) pathway
-visual information to exit the eye
– Horizontal cells
-Influences retinal processing
-receives input from the photoreceptors and project neurites laterally to influence surrounding bipolar cells and photoreceptors
Microscopic Anatomy of the Retina
Direct (vertical) pathway
-visual information to exit the eye
– Amacrine cells
-Influences retinal processing
-receive input from the bipolar cells and project laterally to influence surrounding ganglion cells, bipolar cells, and other amacrine cells.
Microscopic Anatomy of the Retina
Direct (vertical) pathway
-visual information to exit the eye
– Ganglion cells
-fire action potentials in response to light, and these impulses propagate down the optic nerve to the rest of the brain
2 important points
1. The light-sensitive cells in the retina are the photoreceptors. All other cells are influenced by light only by direct or indirect synaptic interactions with the photoreceptors
2 important points
2. The ganglion cells are the only source of output from the retina. No other retinal cell type projects an axon through the optic nerve.
The Laminar Organization
of the Retina
Cells organized in layers
The layers are Inside-out; light must pass from the vitreous humor through the ganglion cells and bipolar cells before it reaches the photoreceptors
Photoreceptor Structure
Conversion of Electromagnetic radiation into neural signals
Photoreceptor Structure
Every photoreceptor has Four main regions
1. Outer segment
Photoreceptor Structure
Every photoreceptor has Four main regions
2. Inner segment
Photoreceptor Structure
Every photoreceptor has Four main regions
3. Cell body
Photoreceptor Structure
Every photoreceptor has Four main regions
4. Synaptic terminal
Photoreceptor Structure
Types of photoreceptors
• Rods (longer) and cones (shorter)
Photoreceptor Structure
Types of photoreceptors
• Rods are 1000x more sensitive to light than cones
-Under nighttime lighting, only rods contribute to vision
-Conversely, under day-time lighting cones do the bulk of the work.
Photoreceptor Structure
Types of photoreceptors
• All rods contain the same
photopigment
-There are three types of cones, each containing a different photopigment, variations among pigments make the different cones sensitive to different wavelengths of light
-cones are responsible for our ability to see color
Photoreceptor Structure
Look at figure on page 290
Regional Differences in Retinal Structure
Varies from fovea to retinal periphery
Regional Differences in Retinal Structure
Peripheral retina
• Peripheral retina has a Higher ratio of rods to cones
Regional Differences in Retinal Structure
Peripheral retina
• Peripheral retina has a Higher ratio of photoreceptors to ganglion cells
Regional Differences in Retinal Structure
Peripheral retina
• The combined effect of this arrangement is that the peripheral retina is more sensitive to light

1.Rods are specialized in low light
2. There are more photoreceptors feeding information to each ganglion cell
Regional Differences in Retinal Structure
Look at figure on page 291
Regional Differences in Retinal Structure:
Cross-section of fovea: Pit in retina
Look at figure on page 292
Regional Differences in Retinal Structure:
Structure: Maximizes visual acuity
Regional Differences in Retinal Structure:
Central fovea: All cones (no rods)
• 1:1 ratio with ganglion cells
Regional Differences in Retinal Structure:
Central fovea: All cones (no rods)
• Area of high visual acuity
Phototransduction
Phototransduction in Rods
Depolarization (movement of positive charge across the membrane) occurs in the dark: “Dark current”
Phototransduction
Phototransduction in Rods
Hyperpolarization in the light
Phototransduction
Phototransduction in Rods
One opsin in rods: Rhodopsin
• Receptor protein that is activated by light
-rebound agonist is called retinal, absorption of light causes a change in the conformation of retinal so that it activates the opsin, this process is called bleaching because it changes the wavelengths absorbed by the rhodopsin.
Phototransduction
Phototransduction in Rods
Look at figure on page 295
Phototransduction in Cones
Similar to rod phototransduction
-Vision during the day depends entirely on the cones, whose photopigments require more energy to become bleached
Phototransduction in Cones
Different opsins
• Red, green, blue
Phototransduction in Cones
Look at figures on page 296
Color detection
Contributions of blue, green,
and red cones to retinal signal
Color detection
Spectral sensitivity
Color detection
Young-Helmholtz trichromacy
theory of color vision
-The brain assigns colors based on a comparison of the readout of the three cone types, when all cones are active then we perceive "white"
Dark and Light Adaptation
Dark adaptation: All-cone daytime vision ---20-25mins-->All-rod nighttime vision this is the process of Dark adaptation
Dark and Light Adaptation
Dark adaptation—factors
• Dilation of pupils
Dark and Light Adaptation
Dark adaptation—factors
• Regeneration of unbleached rhodopsin
Dark and Light Adaptation
Dark adaptation—factors
• Adjustment of functional circuitry
Dark and Light Adaptation
Calcium’s Role in Light Adaptation
The ability of the eye to adapt to changes in light level relies on Calcium concentration changes within the cones.
-When you step out into bright light, the cones are hyperpolarized as much as possible.
-Reason that this happens is because it stems from the fact that the cGMP-gated sodium channels we discussed previously also admit calcium

-When the channels close, a process is initiated that gradually reopens them even if the light level does not change (Ca channels)
Dark and Light Adaptation
Calcium’s Role in Light Adaptation
Indirectly regulates levels of cGMP--> Ca channels
Retinal Processing
Research by
Keffer Hartline, Stephen Kuffler, and Horace Barlow
Retinal Processing
Research study
Action potential discharges retinal
ganglion cells
• Retina: Stimulated with light
Retinal Processing
Transformations in the Outer Plexiform Layer
Photoreceptors
• Release neurons when depolarized
-The transmitter released by photoreceptors is the amino acid glutamate
-Look at figure on page 300
Retinal Processing
Bipolar Cell Receptive Fields
-Categorized into two classes, based on their responses to glutamate
1. In OFF bipolar cells, glutamate- gated cation channels mediate a classical depolarizing EPSP from the influx of Na
Retinal Processing
Bipolar Cell Receptive Fields
-Categorized into two classes, based on their responses to glutamate
2. ON biopolar cells have G-protein-coupled receptors and respond to glutamate by hyperpolarizing
NOTICE
notice that the names OFF and ON refer to whether these cells depolarize in response to light off (more glutamate) or to light on (less glutamate)
Retinal Output
Ganglion Cell Receptive Fields
On-Center (will be depolarized and respond with a barrage of action potentials when a small spot of light is projected onto the middle of its receptive field) and Off-Center cells (Respond to a small dark spot presented to the middle of its receptive field.
-However in both types of cells, the response to stimulation of the center is canceled by the response to stimulation of the surround
Retinal Output
Ganglion Cell Receptive Fields
Responsive to differences in illumination that occur within their receptive fields

-Look at figure on page 302 and 303
Retinal Output
OFF-center cell
Dark in center causes cell to depolarize, whereas dark in the surround causes the cell to hyperpolarize
-In uniform illumination, the center and surround cancel to yield some low level of response.
Types of Ganglion Cells
– Categories based on appearance, connectivity,
and electrophysiological properties
Two types of ganglion cells in monkey and human retina
M-type (Magno=large)(5% of ganglion cells) and P-type (90% of ganglion cells) (Parvo=small)
Color-Opponent Ganglion Cells
Color-opponent cells- Majority of these color-sensitive neurons
-reflecting the fact that the response to one wavelength in the receptive field center is canceled by showing another wavelength in the receptive field surround.
Look at figure on page 305
Parallel Processing
Simultaneous input from two eyes
Information from two streams is compared in
the central visual system
– Depth and the distance of object
Parallel Processing
Simultaneous input from two eyes
Information about light and dark: ON-center and OFF-center ganglion cells
Parallel Processing
Different receptive fields and response properties of retinal ganglion cells: M- and P- cells, and nonM-nonP cells
Concluding Remarks
Light emitted by or reflected off
objects in space --> imaged onto the retina
Concluding Remarks
Transduction
Light energy converted into membrane potentials
Concluding Remarks
Transduction
Phototransduction parallels olfactory transduction
• Electrical-to-chemical-electrical signal
Concluding Remarks
Mapping of visual space onto retinal ganglion cells not uniform
Chapter 10
Visual System II. The Central Visual System
.
Introduction
Neurons in the visual system
Neural processing resulting in perception
Parallel pathway serving conscious visual perception originate in the retina
The pathway serving conscious visual perception includes the lateral geniculate nucleus, primary visual cortex & higher order visual areas in temporal and parietal lobes (aka. area 17, V1, or striate cortex)
The Retinofugal Projection (neural pathway that leaves the eye)
The Optic Nerve- exit the left and right eyes at the optic disks, travel through the fatty tissue behind the eyes in their bony orbits, then pass through holes in the floor of the skull

Optic Chiasm- The optic nerves from both eyes combine to form the optic chiasm, which lies at the base of the brain. The axons originating in the nasal retinas cross from one side to the other. Because only the axons originating in the nasal retinas cross, we say that a partial decussation of the retinofugal projection occurs at the optic chiasm

Optic Tract- Following the partial decussation of at the optic chiasm, the axons of the retinofugal projections form the optic tracts
Decussation- The crossing of a fiber bundle from one side of the brain to the other is called decussation.
Look at figure on page 311
The Retinofugal Projection
Right and Left Visual Hemifields
Optic nerve fibers cross in the optic chiasm so that the left visual hemifield is "viewed" by the right hemisphere and the right visual hemifield is :viewed" by the left hemisphere
Look at figure on page 312
The Retinofugal Projection
Look at figure on page 313
The Retinofugal Projection
Targets of the Optic Tract
-Most of them innervate the lateral geniculate nucleus (LGN) of the dorsal thalamus
-The neurongs in the LGN give rise to axons that project to the primary visual cortex, this projection is called the optic radiation.
Look at figure on page 313
The Retinofugal Projection
While cutting the left optic nerve would render a person blind in the left eye only, a cut of the left optic tract would lead to blindness in the right visual field
Look at figures on page 314
The Retinofugal Projection
Nonthalamic Targets of the Optic Tract:
Hypothalamus: Biological rhythms, including sleep and wakefulness
The Retinofugal Projection
Nonthalamic Targets of the Optic Tract:
Pretectum: Size of the pupil; certain types of eye movement
The Retinofugal Projection
Nonthalamic Targets of the Optic Tract:
Superior colliculus (optic tectum): Orients the eyes in response to new stimuli (saccadic eye movements)
The Lateral Geniculate Nucleus (LGN)
-arranged in 6 layers for each eye and bent around the optic tract like a knee joint.
-segregation of the LGN neurons into layers suggests that different types of retinal information are being kept separate at this synaptic relay, and indeed this is the case: axons arising from M-type, P-type, and nonM-nonP ganglion cells in the two retinas synapse on cells in different LGN layers
Look at figure on page 317
The Lateral Geniculate Nucleus (LGN)
-LGN neurons receive synaptic input from the retinal ganglion cells, and most geniculate neurons project an axon to primary visual cortex by the optic radiation
The Segregation of Input by Eye and by Ganglion Cell Type
Look at figure on page 318
Magnocellular LGN layers
the ventral layers of LGN.
-layers 1 and 2 contain larger neurons
Parvocellular LGN layers
the dorsal layers of LGN
-layers 3-6 contain smaller neurons
Receptive Fields
Receptive fields of LGN neurons: Identical to the ganglion cells that feed them
Receptive Fields
Magnocellular LGN neurons: Large, monocular receptive fields with transient response
Receptive Fields
Parvocellular LGN cells: Small,monocular receptive fields with sustained response
Nonretinal Inputs to the LGN
Retinal ganglion cells axons: Not the main source of synaptic input to the LGN
Look at figure on page 319
Nonretinal Inputs to the LGN
Primary visual cortex: 80% of the synaptic inputs to the LGN
Nonretinal Inputs to the LGN
Neurons in the brain stem: regulate the influence on neuronal activity
Anatomy of the Striate Cortex
-LGN has a single major synaptic target: primary visual cortex
Look at figure on page 321
Anatomy of the Striate Cortex
Retinotopy- an organization whereby neighboring cells in the retina feed information to neighboring places in their target structures
Map of the visual field onto a target structure
(retina, LGN, superior colliculus, striate cortex) -
overrepresentation of central visual field
Anatomy of the Striate Cortex
Retinotopy
Discrete point of light: Activates many cells in the retina, and other target structures
Anatomy of the Striate Cortex
Retinotopy
Perception: Based on the brain’s interpretation of distributed patterns of activity
Anatomy of the Striate Cortex
Look at figure on page 320
Anatomy of the Striate Cortex
Lamination of the
Striate Cortex
Layers I - VI
Anatomy of the Striate Cortex
Lamination of the
Striate Cortex
Spiny stellate cells: Spinecovered
dendrites; seen in layer IVC
-Make local connections only within the cortex
Anatomy of the Striate Cortex
Lamination of the
Striate Cortex
Pyramidal cells: Spines; thick
apical dendrite; layers III, IVβ, V, VI
-only pyramidal cells send axons out of striate cortex to form connections with other parts of the brain.
Anatomy of the Striate Cortex
Lamination of the
Striate Cortex
Inhibitory neurons: Lack
spines; All cortical layers;
Forms local connections
Anatomy of the Striate Cortex
Lamination of the
Striate Cortex
Look at figure on page 321
Anatomy of the Striate Cortex
Inputs to the Striate Cortex
Magnocellular LGN neurons project to layer IVCα
Anatomy of the Striate Cortex
Inputs to the Striate Cortex
Parvocellular LGN neurons Project to layer IVCβ
Anatomy of the Striate Cortex
Inputs to the Striate Cortex
Koniocellular LGN axons: Bypasses layer IV to make synapses in layers II and III
Anatomy of the Striate Cortex
Ocular Dominance Columns
Look at figure on page 322
Layer IVC innervates superficial layers
Look at figure on page 323, 10.17
Outputs of the Striate Cortex:
Layers II, III, and IVB:
Projects to other cortical
areas
Look at figure on page 323, 10.18
Outputs of the Striate Cortex:
Layer V: Projects to the
superior colliculus and pons
Outputs of the Striate Cortex:
Layer VI: Projects back to
the LGN
Cytochrome Oxidase
Blobs
Cytochrome oxidase is a
mitochondrial enzyme
used for cell metabolism
Cytochrome Oxidase
Blobs
Blobs: Cytochrome oxidase
staining in cross sections
of the striate cortex
Receptive Fields
Layer IVC: Monocular; center-surround
Receptive Fields
Layer IVCα: Insensitive to the wavelength
Receptive Fields
Layer IVCβ: Center-surround color
opponency
Binocularity
Layers superficial to IVC: First binocular receptive fields in the visual pathway
Receptive Fields
Orientation Selectivity
Look at figure on page 325
Receptive Fields
Direction Selectivity
Neuron fires action potentials in response to moving bar of light

Look at figures on page 327
Receptive Fields
Simple cells: Binocular; Orientationselective;
Elongated on-off region with
antagonistic flanks responds to optimally oriented bar of light (location in the field)
Receptive Fields
Possibly composed of three LGN cell axons with center-surround receptive fields
Receptive Fields
Complex cells: Binocular; Orientationselective;
ON and OFF responses to the bar of light but unlike simple cells, no distinct on-off regions
Look at figure on page 329
Physiology of the Striate Cortex
Receptive Fields
Blob Receptive Fields
• Blob cells: Wavelength-sensitive; Monocular; No
orientation; direction selectivity
Physiology of the Striate Cortex
Receptive Fields
Parallel Pathways: Magnocellular;
Koniocellular; Parvocellular

-Look at figure on page 330
Physiology of the Striate Cortex
Receptive Fields
Cortical Module
-Look at figure on page 332
Beyond Striate Cortex
Dorsal stream
Analysis of visual
motion and the visual
control of action
Beyond Striate Cortex
Ventral stream
Perception of the
visual world and the
recognition of objects
-Look at figure on page 333
Beyond Striate Cortex
The Dorsal Stream (V1, V2, V3, MT, MST, Other dorsal areas)
Area MT (temporal lobe)
• Most cells: Direction-selective; Respond more
to the motion of objects than their shape
Beyond Striate Cortex
The Dorsal Stream (V1, V2, V3, MT, MST, Other dorsal areas)
Beyond area MT - Three roles of cells in area MST (parietal lobe)
– Navigation
Beyond Striate Cortex
The Dorsal Stream (V1, V2, V3, MT, MST, Other dorsal areas)
Beyond area MT - Three roles of cells in area MST (parietal lobe)
– Directing eye movements
Beyond Striate Cortex
The Dorsal Stream (V1, V2, V3, MT, MST, Other dorsal areas)
Beyond area MT - Three roles of cells in area MST (parietal lobe)
– Motion perception
The Ventral Stream (V1, V2, V3, V4, IT, Other ventral areas)
Area V4
Achromatopsia: Clinical syndrome in humans-caused by damage to area V4; Partial or complete loss of color
vision
The Ventral Stream (V1, V2, V3, V4, IT, Other ventral areas)
Area IT
Major output of V4
The Ventral Stream (V1, V2, V3, V4, IT, Other ventral areas)
Area IT
Receptive fields respond to a wide variety of colors and
abstract shapes
The Ventral Stream (V1, V2, V3, V4, IT, Other ventral areas)
Area IT
prosopagnosia
From Single Neurons to Perception
Visual perception
– Identifying & assigning meaning to objects
From Single Neurons to Perception
Hierarchy of complex receptive fields
Retinal ganglion cells: Center-surround structure, Sensitive to contrast, and wavelength of light
From Single Neurons to Perception
Hierarchy of complex receptive fields
Striate cortex: Orientation selectivity, direction selectivity, and binocularity
From Single Neurons to Perception
Hierarchy of complex receptive fields
Extrastriate cortical areas: Selective responsive to complex shapes; e.g., Faces
From Single Neurons to Perception
From Photoreceptors to Grandmother Cells
Grandmother cells: Face-selective neurons in area IT?
From Single Neurons to Perception
From Photoreceptors to Grandmother Cells
Probably not: Perception is not based on the activity of individual, higher order cells
From Single Neurons to Perception
Parallel Processing and Perception
Groups of cortical areas contribute to the perception of color,motion, and identifying
object meaning
Concluding Remarks
Vision
Perception combines individually
identified properties of visual objects
Concluding Remarks
Vision
Achieved by simultaneous, parallel
processing of several visual pathways
Concluding Remarks
Parallel processing
Like the sound produced by an orchestra of visual areas rather than the end product of an assembly line