5) Exteroceptive Sensory Systems
- Auditory (hearing)
- Somatosensory (touch)
- Olfactory (smell)
- Gustatory (taste)
- Vision
Primary Sensory Cortex
Receives most of its input directly from the thalamic relay
nuclei of that system
Secondary Sensory Cortex
Receives most input from primary sensory cortex of that
system or from other areas of the secondary cortex of that
same system
Association Cortex
Any area of the cortex that receives input from more than 1
sensory system (most from the secondary)
Hierarchial Organization of System
Based on specificity and complexity of system structure's
function - the higher level of damage, the greater the effect
Simple to Complex:
Receptors - Thalamic Relay Nuclei - Primary Sensory Cortex -
Secondary Sensory Cortex - Association Cortex
Sensation
Process of detecting the presence of stimuli
Perception
Higher-order process of integrating, recognizing, and
interpreting complete patterns of sensations
Functional Segregation
Each cerebral cortex level contains functionally distinct areas
that specialize in different kinds of analyses
Parallel Processing
Simultaneous analyses of a signal in different ways by the
multiple parallel pathways of a neural network = "2
streams" :
- w/ conscious awareness
- w/o conscious awareness
Sensory Systems' History (evolution)
hierarchal, functionally homogenous, & serial
TO
hierarchal, functionally segregated, parallel
(systems are characterized by a division of labor = multiple
specialized areas at multiple levels are interconnected by
multiple parallel pathways...perceptions are a product of the
combined activity of different interconnected cortical areas =
"binding problem")
Sounds
Vibrations of air molecules - always complex patterns of
vibrations, not "pure tones"/a sine wave
Amplitude corresponds with...
Loudness
Frequency corresponds with...
Pitch
Complexity corresponds with...
Timbre
Fourier Analysis
mathematical procedure to break down complex waves into
their component sine waves
Fundamental Frequency
The pitch of a complex sound may not be directly related to
the frequency of any of the sound's components = because
what is perceived is the HIGHEST frequency of which the
various component frequencies of a sound are multiples
Path of Ear
Sound waves - outer ear - auditory canal - vibrates the
eardum (tympanic membrane) - 3 ossicles - oval window -
cochlea - organ of corti
Tympanic Membrane
Eardum
3 Ossicles
Small bones of the middle ear:
- malleus (hammer)
- incus (anvil)
- stapes (stirrup)
Oval Window
membrane
Cochlea
Snail-like tube, containing fluid, with an internal membrane
called the Organ of Corti
Organ of Corti
Auditory receptor organ, has 2 membranes:
- basilar membrane
- tectorial membrane
Basilar Membrane
HAIRS are mounted here
Tectorial Membrane
Rests on the hair cells
Each pressure change...
travels along as a wave and stimulates hair cells
Auditory Nerve
Branch of cranial nerves
Auditory System Organization is: TONOTOPIC (vs.
retinotopic in the visual system)
Nerves are specialized according to frequency...higher
frequencies produce greater activation closer to the windows
Semicircular Canals
Receptive organs of the VESTIBULAR SYSTEM: carries info
about the direction and intensity of head movements, which
helps us maintain our balance
Visual System Pathway vs. Auditory PathwayS
retina - geniculate - striate
VS.
a NETWORK of auditory pathways
Superior Olives
On both sides of brain stems - for the localization of sounds
in space
- medial = timing
- lateral = amplitude
Inferior Colliculi
Superior Olives connect to it via the "lateral lemniscus"
Medial Geniculate Nuclei (of thalamus)
Projects to the primary auditory cortex
- organized in columns, tonotopically (function = frequency)
- temporal lobe
Areas of Association Cortex
Where sensory systems interact
- prefrontal cortex (anterior auditory pathway, more involved
in identifying sounds = "WHAT")
- posterior parietal cortex (posterior auditory pathway, more
involved in locating sounds = "WHERE")
Auditory Pathways
Superior Olives - Lateral Lemniscus - Inferior Colliculi -
Medical Geniculate Nuclei (thalamus) - Primary Auditory
Cortex (temporal lobe) - Secondary auditory cortex (more
complex neural responses) - Areas of Association cortex
(prefrontal cortex, posterior parietal cortex)
Damage to Ears
- Very few permanent hearing deficits
- Can lose ability to localize sound and discriminate
frequencies
- Severe ear problems = from damage to the inner/middle
ear rather than central damage
'conductive deafness' vs. 'nerve deafness' vs. tinnitus
conductive = damage to ossicles (sm. ear bones)
nerve = damage to cochlea/auditory nerve (= loss of hair cell
receptors...need a cochlear implant)
tinnitus = ringing of the ears
Somatosensations
Sensations from your body
3 Interacting systems mediate the somatosensory system:
1) Exteroceptive System
2) Proprioceptive System
3) Interoceptive System
Exteroceptive System
Senses external stimuli applied to the skin:
- mechanical (touch)
- thermal (temperature)
- nociceptive (pain)
Proprioceptive System
Monitors info about the position of the body that comes from
receptors in the muscles, joints, and organs of balance
Interoceptive System
Provides general info about conditions within the body (blood
pressure, temperature, etc.)
Cutaneous (in the skin) Receptors
specialized:
- free nerve endings
- pacinian corpuscles
- Merkel's disks & Ruffini endings
- Stereognosis
Free Nerve Endings
Neuron endings w/ no specialized structures on them
(sensitive to temperature and pain)
Pacinian Corpuscles (vs. Merkel's disks & Ruffini endings)
Adapt rapidly to sudden skin displacements
(vs. slower adapters/responders = R & M's)
Stereognosis
Identification of objects by touch
By which roots do the neural fibers that carry info from the
cutaneous receptors into the spinal cord use?
Dorsal Roots
Dermatome
Area of body innervated by the left and right dorsal roots of a
given segment of the spinal cord
2 Major Pathways of the Somatosensory System
work hand in hand:
1. Dorsal-Column Medial-Lemniscus System
2. Anterolateral System
Dorsal-Column Medial-Lemniscus System
Carries info about touch and proprioception:
1. sensory neurons enter spinal cord via dorsal root
2. ascend ipsilaterally in the dorsal columns
3. synapse in the dorsal column nuclei of the medulla, where
axons decussate
4. ascend in the medial lemniscus
5. to the contralateral ventral posterior nucleus
6. -- some neurons project to the primary somatosensory
cortex OR others to the secondary somatosensory
cortex/posterior parietal cortex
Anterolateral System
Carries info about pain and temperature:
1. dorsal root neurons synapse as soon as they enter the
spinal cord
2. axons decussate - "spinothalamic tract" = project to the
ventral posterior nucleus of thalamus
OR
ascend ipsilaterally - "spinoreticular tract" = reticular
formation - parafasicular nuclei - intralaminar nuclei of
thalamus
Somatotopic
Human Primary Somatosensory Cortex (S1) is organized
according to a map ("somatosensory homunculus") of the
body surface
- only small areas of S1 receive input from large areas of the
body
- can divide receptive fields into excitatory and inhibitory
areas, columnar organization (4 'strips')
- neurons: anterior to posterior = simple to complex
S1 vs. S2 (primary vs. secondary somatosensory cortexes)
S2 lies ventral to S1, receives info from S1 from BOTH sides
of the body (vs. S1's contralateral-ness)
Association Cortex of Somatosensory
Posterior Parietal Lobe
Damage to S1
reduced ability to detect light touch and identify objects by
touch
Damage to Association Cortex (posterior parietal cortex)
'bimodal neurons' = respond to activation of 2 different
sensory systems
Astereognosia
Inability to recognize objects by touch
Asomatognosia
Failure to recognize parts of one's own body (often unilateral
- left side b/c of right brain damage)
Anosognosia
Failure of neuropsychological patients to recognize their own
symptoms
Contralateral Neglect
Tendency not to respond to stimuli that are contralateral to a
right-hemisphere injury
Why is the Perception of Pain "PARADOXICAL" (a logical
contradiction)
1. warns us to stop something dangerous (evolutionarily
beneficial)
2. no obvious cortical representation except for "anterior
cingulate cortex" being linked to the emotional reaction to
pain (not perception of it)
3. it can be so effectively suppressed by cognitive and
emotional factors
Gate-Control Theory
Ability of cognitive and emotional factors to block pain
(descending from brain pain-control)
Periaqueductal Gray (PAG)
Stimulation has pain-blocking effects
Endorphins
Internally produced opiate analgesics
Neuropathic Pain
Severe chronic pain in the absence of a recognizable pain
stimulus (ex: after a healed injury, or an amputation)
Olfaction & Gustation = what?
Smell and Taste monitor chemical content of the
environment, produce FLAVOR
Pheromones
released by many species, chemicals that influence the
physiology and behavior of others of the same species
Olfactory Mucosa
Where receptor cells lie = in a layer of mucus-covered tissue
Olfactory Bulbs
Where axons synapse on neurons that project via the
olfactory tracts to the brain
Receptor Organization
RANDOMLY scattered - but proteins project to the same
location of the bulb
Olfactory Glomeruli
Discrete clusters of neurons near the surface of the olfactory
bulbs (where olfactory receptor axons terminate)
**each receives input from 1000's of receptor cells
Glomeruli Layout
- mirror symmetry between left and right bulbs
- glomeri are arranged the same among bulbs of many
species and is similar in related species
Are olfactory recepter cells replacable?
YES!
Piriform Cortex
PRIMARY OLFACTORY CORTEX = An area of the medial
temporal cortex (lobe)
Olfactory path (2 paths)
Primary Olfactory Cortex (piriform/temporal lobe)
- does NOT pass thru the thalamus - lymbic system
(emotional response to odors)
OR
- projects via MEDIAL DORSAL NUCLEI (thalamus) -
orbitofrontal cortex (frontal lobes) (conscious perception of
odors)
Taste Buds
Taste receptors found in clusters in the oral cavity -
"papillae"
Receptors for 5 Primary Tastes
1. Sweet
2. Sour
3. Bitter
4. Salty
5. Unami (meaty)
Solitary Nucleus
Where fibers terminate in the medulla, synapsing on neurons
that project to the VENTRAL POSTERIOR NUCLEUS of the
thalamus
Gustatory Pathway
front of tongue - back of tongue - back of oral cavity -
solitary nucleus - ventral posterior nucleus (thalamus) -
primary gustatory cortex (lateral fissure) - 2ary gustatory
cortex
Anosmia
Inability to smell
Ageusia
Inability to taste - rare, sometimes due to same side ear
damage
Simultanagnosia
Attention disorder - specifically visual (trouble focusing on
more than 1 object at a time)
= bilateral damage to dorsal stream (responsible for visually
localizing objects in space)
Selective Attention
We consciously perceive only a small subset of the many
stimuli that excite our sensory organs at any 1 time and
largely ignore the rest
= improves perception of the stimuli that are its focus
= interferes with perception of the stimuli that are not its
focus
Attention can be focused in 2 ways:
- Internal Cognitive Processes (endogenous attention - ex:
looking for keys) "top-down"
- External Events (exogenous attention - ex: something fell
over nearby) "bottom-up"
Cocktail-Party Phenomenon
Brain can block from conscious awareness all stimuli except
those of a particular kind while still monitoring the blocked
out stimuli in case something comes up that requires
attention
Change Blindness
Noting NO change between 2 similar/same photos = we have
no memory for parts of the scene that are not the focus of
our attention
Ventral vs. Dorsal Stream Activities
Ventral = attention to shape/color, identify face
Dorsal = attention to movement, note position difference of
face in a picture
Sensation & Perception - When we smell a fragrant flower, are we experiencing a
sensation or a perception? In everyday language, the terms "sensation" and
"perception' are often used interchangeably.
However, as you will soon see, they are very distinct, yet complementary processes. In
this section, we will discuss some concepts central to the study of sensation and
perception and then move on to discuss vision and the perception of pain (it is not
possible in the scope of these notes to discuss all the senses).
I. Sensations and Perceptions
Sensations can be defined as the passive process of bringing information from the
outside world into the body and to the brain. The process is passive in the sense
that we do not have to be consciously engaging in a "sensing" [Link] can
be defined as the active process of selecting, organizing, and interpreting the
information brought to the brain by the senses.
A) HOW THEY WORK TOGETHER:
1) Sensation occurs:
a) sensory organs absorb energy from a physical stimulus in the environment.
b) sensory receptors convert this energy into neural impulses and send them to the
brain.
2) Perception follows:
a) the brain organizes the information and translates it into something meaningful.
B) But what does "meaningful" mean? How do we know what information is important
and should be focused on?
1) Selective Attention - process of discriminating between what is important & is
irrelevant (Seems redundant: selective-attention?), and is influenced by motivation.
For example - students in class should focus on what the teachers are saying and the
overheads being presented. Students walking by the classroom may focus on people in
the room, who is the teacher, etc., and not the same thing the students in the class.
2) Perceptual Expectancy - how we perceive the world is a function of our past
experiences, culture, and biological [Link] example, as an American, when I look
at a highway, I expect to see cars, trucks, etc, NOT airplanes. But someone from a
different country with different experiences and history may not have any idea what to
expect and thus be surprised when they see cars go driving by.
Another example - you may look at a painting and not really understand the message
the artist is trying to convey. But, if someone tells you about it, you might begin to see
things in the painting that you were unable to see before.
ALL OF THIS IS CALLED Psychophysics
C) Psychophysics can be defined as, the study of how physical stimuli are
translated into psychological experience.
In order to measure these events, psychologists use THRESHOLDS.
1) Threshold - a dividing line between what has detectable energy and what does not.
For example - many classrooms have automatic light sensors. When people have not
been in a room for a while, the lights go out. However, once someone walks into the
room, the lights go back on. For this to happen, the sensor has a threshold for motion
that must be crossed before it turns the lights back on. So, dust floating in the room
should not make the lights go on, but a person walking in should.
2) Difference Threshold - the minimum amount of stimulus intensity change needed to
produce a noticeable change.
the greater the intensity (ex., weight) of a stimulus, the greater the change needed to
produce a noticeable change.
For example, when you pick up a 5 lb weight, and then a 10 pound weight, you can feel
a big difference between the two. However, when you pick up 100 lbs, and then 105 lbs,
it is much more difficult to feel the difference.
3) Signal-Detection Theory - detection of a stimulus involves some decision making
process as well as a sensory process. Additionally, both sensory and decision making
processes are influenced by many more factors than just intensity.
a) Noise - how much outside interference exists.
b) Criterion - the level of assurance that you decide must be met before you take action.
Involves higher mental processes. You set criterion based on expectations and
consequences of inaccuracy.
For example - at a party, you order a pizza...you need to pay attention so that you will
be able to detect the appropriate signal (doorbell), especially since there is a lot of noise
at the party. But when you first order the pizza, you know it won't be there in 2 minutes,
so you don't really pay attention for the doorbell. As the time for the pizza to arrive
approaches, however, your criterion changes...you become more focused on the
doorbell and less on extraneous noise.
II. SIGHT/VISION
A) The visual system works on sensing and perceiving light waves. Light waves vary in
their length and amplitude:
a) wave length (also referred to as frequency, since the longer a wave, the less
often/quickly it occurs) - affects color perception (ex., red=approx 700, yellow approx
600)
b) wave amplitude (this is the size/height of the wave) - affects brightness perception.
B) Structure of The EYE:
1) Cornea - the round, transparent area that allows light to pass into the eye.
2) Lens - the transparent structure that focuses light onto the retina.
3) Retina - inner membrane of the eye that receives information about light using rods
and cones. The functioning of the retina is similar to the spinal cord - both act as a
highway for information to travel on.
4) Pupil - opening at the center of the iris which controls the amount of light entering the
eye. Dilates and Constricts.
5) Rods & Cones - many more rods (approximately 120 million) than cones (approx 6.4
million).
a) cones - visual receptor cells that are important in daylight vision and color vision.
the cones work well in daylight, but not in dim lighting. This is why it is more difficult to
see colors in low light.
most are located in the center of the retina...called the FOVEA, which is a tiny spot in
the center of the retina that contains ONLY cones...visual acuity is best here.
SO...when you need to focus on something you attempt to bring the image into the
fovea.
b) rods - visual receptor cells that are important for night vision and peripheral vision.
the rods are better for night vision because they are much more sensitive than cones.
in addition, the rods are better for peripheral vision because there are many more on the
periphery of the retina. The cones are mostly in and around the fovea but decrease as
you go out.
to see best at night, look just above or below the object...this keeps the image on the
rods.
C) Seeing In Color - we can see many colors, but only have 3 types of cones that
receive information about color. We have cones that pick up light waves for red, green,
and blue.
Color Vision Theories:
1) Trichromatic Theory - this theory indicates that we can receive 3 types of colors (red,
green, and blue) and that the cones vary the ratio of neural activity (Like a projection
T.V.). The ratio of each color to the other then determines the exact color that we see.
2) Opponent-Process Theory - color perception depends on the reception of pairs of
antagonist colors. Each receptor can only work with one color at a time so the opponent
color in the pair is blocked out. Pairs = red-green, blue-yellow, black- white (light-dark).
Note: Most every Introductory Psychology book has a demonstration on the Opponent-
Process theory. Please look for the one in your book and give it a try.
DOES COLOR EXIST? People just assume that because we see colors, that they
actually exist in the world. In other words, that when they see the color red, that red is a
real, physical, tangible, "thing". But is it, or is color just a matter of our perception? If we
had different types of nervous systems, we would see things differently (literally) and so
wouldn't we think those other things we saw were the real "things"? Let's examine this
question of perception a bit further.
II) PERCEPTION
Much of our understanding of how and why we perceive things comes from Gestalt
Psychology
For example - one of the most well known Gestalt principles is the Phi Phenomenon,
which is the illusion of movement from presenting stimuli in rapid succession. When you
see a cartoon or running Christmas lights, you see movement (although none actually
exists) because of this principle.
A) Gestalt Principles of Perceptual Organization
1) figure-ground - this is the fundamental way we organize visual perceptions. When we
look at an object, we see that object (figure) and the background (ground) on which it
sits. For example, when I see a picture of a friend, I see my friends face (figure) and the
beautiful Sears brand backdrop behind my friend (ground).
2) simplicity/pragnanz (good form) - we group elements that make a good form.
However, the idea of "good form" is a little vague and subjective. Most psychologists
think good form is what ever is easiest or most simple. For example, what do you see
here: :
do you see a smiling face? There are simply 3 elements from my keyboard next to each
other, but it is "easy" to organize the elements into a shape that we are familiar with.
3) proximity - nearness=belongingness. Objects that are close to each other in physical
space are often perceived as belonging together.
4) similarity - do I really need to explain this one? As you probably guessed, this one
states that objects that are similar are perceived as going together. For example, if I ask
you to group the following objects: (* * # * # # #) into groups, you would probably place
the asterisks and the pound signs into distinct groups.
5) continuity - we follow whatever direction we are led. Dots in a smooth curve appear to
go together more than jagged angles. This principle really gets at just how lazy humans
are when it comes to perception.
6) common fate - elements that move together tend to be grouped together. For
example, when you see geese flying south for the winter, they often appear to be in a
"V" shape.
7) closure - we tend to complete a form when it has gaps.
B) Illusions - an incorrect perception caused by a distortion of visual sensations.
1) Muller-Luyer Illusion
2) Reversible Figures - ambiguous sensory information that creates more than 1 good
form. For example, the picture of two faces looking toward each other that is also a
vase. I am sure most every Introductory Psychology book has this example.
3) Impossible Figures - objects that can be represented in 2-dimensional pictures but
can not exist in 3-dimensional space despite our perceptions. You know the artist,
Escher who draws the pictures like...the hands drawing each other, the waterfall that
goes down and stays level at the same time, etc...
C) THE PERCEPTION OF PAIN
Pain is an unpleasant yet important function for survival: warning system (but not all
pain is needed for survival).
There are two different pathways to the brain on which pain can travel - information
brought from free nerve endings in the skin to the brain via two different systems:
1) fast pathways - registers localized pain (usually sharp pain) and sends the
information to the cortex in a fraction of a second. EX. - cut your finger with a knife.
2) slow pathways - sends information through the limbic system which takes about 1-2
seconds longer than directly to the cortex (longer lasting, aching/burning).
Factors in Pain Perception - not an automatic result of stimulation:
1) expectations - research shown that our expectations about how much something will
hurt can effect our perception.
Melzack - indicated that believing that something will be very painful helps us prepare
for it.
For example - child birth: Lamaze method falsely leads us to believe it won't be painful.
Maybe if we know it will be bad we can adequately prepare to handle it.
another example - placebo effect - if we believe pain has stopped, it may.
2) personality - people with negative types of personalities often have more pain.
For example - a very uptight person may experience muscle pains, back pains, etc.
3) mood - bad moods, angry, unhappy, etc, can lead to the experience of increased
pain.
For example - study manipulated moods of subjects then asked them to complete
questionnaires of pain perception. Those in negative mood group reported significantly
more pain than other subjects.
So, it seems that our brains can regulate, control, determine, and even produce pain.
THEORIES OF PAIN PERCEPTION
1) Gate Control Theory (Melzack & Walls, 1965) - incoming pain must pass through a
"gate" located in the spinal cord which determines what information about pain will be
sent to the brain. So, it can be opened to allow pain through or closed to prevent pain
from being perceived.
The Gate - actually a neural network controlled by the brain. Located in an area of the
spinal cord called the Substansia Gelatinosa. There are two types of nerve fibers in this
area:
a) large - sends fast signals and can prevent pain by closing the gate.
b) small - sends slower signals which open the gate. So - when pain occurs it is
because the large fibers are off and the small are on, opening the gate.
Since the gate is controlled by the brain, he factors discussed earlier (expectations,
mood, personality) influence the functioning of the gate.
Contradiction to Gate Control Theory:
1) endorphins - the body's own pain killers (morphine-like). May explain acupuncture,
acupressure, pain tolerance during last two weeks of pregnancy, etc.
BUT- endorphins may work with the gate control theory - maybe pain is perceived,
endorphins are released, so the brain no longer needs the signals and closes the gate.
PHANTOM LIMBS
Ability to feel pain, pressure, temperature, and many other types of sensations including
pain in a limb that does not exist (either amputated or born without).
The feelings and the pain are sometimes so life-like that person attempts to pick things
up with phantom hand, step with phantom foot or leg, etc. Often person feels phantom
moving in perfect coordination with the rest of the body - some report a missing arm
extending outward at a 90 degree angle so they turn sideways when going through a
doorway.
May occur right after amputation or not until years later.
Often felt as part of the body (belonging to the rest of the body). EX. - with a missing
leg, some report having a phantom foot but not the rest of the leg. Still, the foot feels as
though it is part of the body.
Explanations:
1) the neuroma explanation - remaining nerves in the stump grow into nodules
(neuromas) at the end of the stump continue to fire signals. Signals follow the same
pathways the brain as when the appendage existed.
2) the spinal cord explanation - neurons in the spinal cord that are no longer receiving
information from the lost appendage continue to send information to the brain.
Problem - studies have shown that when areas in the spinal cord are severed often
feelings still being perceived from areas that meet the spinal cord in lower areas (below
separation in spinal cord).
3) the brain explanation - signals in the somatosensory circuits of the brain change
when the limb is lost which produce the phantom...the brain compensates for the loss or
altered signals. This has been expanded - brain contains a network of fibers that not
only respond to stimulation but continually generates a pattern of impulses that indicate
that the body is intact and functioning. Thus, the brain creates the impression that the
limb exists and is al right. This system may be prewired.
4) the hardwired explanation - we may have a biological makeup to be born with all of
our appendages. So, when we are born w/o one or lose one, the nerves are still there
and are still going to send the information.