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    Sensation

    &

    Perception

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    17 Somatosensation

    Touch

    Heat/Cold

    Pain

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    Administrative stuff

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    The final

    Friday, December 10th, 10.30 am -12.30 pm

    In HGS 101

    Comprehensive, with an emphasis on things

    that havent been covered in previous exams.

    Like the midterms. Interpreting graphs,

    drawing diagrams, multiple choice and short

    written answers.

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    The early final

    For graduating seniors

    Their grades have to be in by Dec the 3rd.

    Format is similar as in regular final

    This leaves only one day: Dec the 2nd.

    But when on the 2nd? Someone has a mandatory

    class from 11-12, someone else has another finalfrom 2.30.

    What about really early (911 am) or really late

    (5-7 pm)?

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    The Review session

    When?

    Where?

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    Auditory system

    PostScript

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    Young adults are most sensitive for

    frequencies corresponding to human speech

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    Range of audible frequencies of several species at 60 dB

    SPL. No species covers the whole range.

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    SomatosensationTouch

    Temperature

    Pain

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    Organ: The skin

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    SomatosensationTouch

    Temperature

    Pain

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    The skin receptors (transducers)

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    The skin receptors (transducers)

    a) Merkel b) Meissner

    c) Ruffini d) Pacinian

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    The skin receptors (transducers)

    Meissner corpuscle, shallow, stack of cells

    Merkel disk, shallow, disk-shaped

    Ruffini endings, deep, branched cylinder

    Pacinian corpuscle, very deep, onion-like

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    Spatiotemporal fiber properties

    Time constants

    short long

    RF

    structure

    Small, Sharp

    boundaries

    Large, fuzzy

    boundaries

    Rapidly adapting,

    punctate

    (RA-P)

    Slowly adapting,

    punctate

    (SA-P)

    Rapidly adapting,

    diffuse

    (RA-D)

    Slowly adapting,

    diffuse

    (SA-D)

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    The skin receptors (transducers)

    Meissner corpuscle: RA-P

    Merkel disk: SA-P Ruffini endings: SA-D

    Pacinian corpuscle: RA-D

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    Spatiotemporal fiber properties

    Time constants

    short long

    RF

    structure

    Small, Sharp

    boundaries

    Large, fuzzy

    boundaries

    (RA-P) (SA-P)

    (RA-D) (SA-D)

    Meissner

    Corpuscle

    Merkel

    Disk

    Pacinian

    Corpuscle

    Ruffini

    Ending

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    The skin receptors (transducers)

    Meissner corpuscle. Hooked up to RA-P fibers.

    Optimal stimulus: Pressure. Perception: Flutter.

    Merkel disk. Hooked up to SA-P fibers.

    Optimal stimulus: Light tapping. Perception: Pressure.

    Ruffini endings. Hooked up to SA-D fibers.

    Optimal stimulus: Lateral Stretching. Perception: Stretch.

    Pacinian corpuscle. Hooked up to RA-D fibers.

    Optimal stimulus: Vibration. Perception: Vibration.

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    One last word on skin receptors

    Merkel disks = Merkel discs = Merkel receptors

    Ruffini endings = Ruffini cylinders

    SA-P = SA-1, SA-D = SA2

    RA-P = RA-1, RA-D = RA2

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    On receptive fields on the skin

    The vision analogy holds

    Somatosensory receptive fields have a

    antagonistic center-surround structure

    For the following experiment, have two

    sharp (but relatively blunt) objects like

    toothpicks or pen-points ready.

    If you hurt yourself, dont sue anyone.

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    On receptive fields on the skin

    Excitatory

    region

    Inhibitory

    region

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    On receptive fields on the skin

    1 stimulus in

    excitatory

    region:

    Feels like 1

    strong point

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    On receptive fields on the skin

    2 stimuli inexcitatory

    regions, far

    apart:

    Feels like 2separate strong

    points

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    On receptive fields on the skin

    2 stimuli inexcitatory

    regions, close

    together (both

    inhibiting andexciting each

    other)

    Feels like 1 point

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    On receptive fields on the skin

    2 stimuli inexcitatory

    regions,

    intermediate

    distance (eachinhibits the

    other)

    Feels like 2 weak

    points

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    Relative sensitivity of body parts

    This rationale (measuring the two-point-threshold) was used to elicit the absolute

    sensitivity of different regions of skin. It allows to infer the spacing of receptive

    fields andin turn the spatial resolution ofa given skin region for a given body part.

    This was done very early (as early as 1880),using a compass.

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    2-point Thresholds on skin

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    Pathways for touch

    Afferent touch fibers enter the spinal cord in the dorsalroot.

    Some of the fibers synapse onto local inter-neurons whichin turn synapse onto motor neurons, implementing a spinalreflex arc (like a withdrawal reflex)

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    Ascending pathways for touch

    Other fibers travel upwardto the brain, carryingtouch-information to thebrain-stem.

    They are in the lemniscalpathway

    After synapsing in thebrainstem, fibers cross tothe opposite side of thebrain.

    After synapsing in thethalamus, these neuronsproject to cortical areas.

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    Cortical processing of touch

    The projection area is

    the somatosensory

    cortex (S1, S2) It is situated

    immediately posterior

    of the central sulcus.

    Both S1 and S2

    contain a full sensory

    MAP of the body.

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    The sensory homunculus

    Literally

    http://www.cs.uta.fi/~jh/homunculus.htmlhttp://www.cs.uta.fi/~jh/homunculus.html
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    Threshold/Homunculus

    Correspondence The lower the 2-point

    threshold for a given skin

    region (= higher spatialresolution), the more

    relativespace is devoted to

    cortical represenation/

    processing of that region insomatosensory cortex.

    Roughly an inverse

    correlation.

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    Sidenote

    Recently, the analogy between retinotopic

    mapping in vision and somatotopic mapping in

    somatosensation has come under fire. Critics of this model argue that there are many

    more body maps in the somatosensory cortices

    They also argue that the mapping is much more

    distributed and not somatotopic.

    Homunculus model too nice to be true.

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    Another analogy to vision

    Just like in the case of eye movements, where the

    brain needs to know if the eyes moved or the

    world moved, an efference copy is sent from themotor system to the sensory system.

    In the case of the somatosensory system, this is the

    somatosensory cortex.

    This is the reason why one cant tickle oneself.The body knows that its not an external stimulus

    and takes it into account (dismissing it).

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    Somatosensory efference copy and

    tickling If one is instructed to push oneself and then

    push another person with the same force,

    the other person experiences this as a muchstronger push (escalation)

    Blakemore et al. (1998):

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    By the way

    Brodmann Areas 1,2,3

    = Somatosensory cortex

    = S1, S2

    =Postcentral gyrus

    =Postcentral cortex

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    SomatosensationTouch

    Temperature

    Pain

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    Thermoreceptors

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    Thermoreceptors

    Free nerve endings act as thermoreceptors

    There are separate classes of thermoreceptors:

    Cold fibers and Warm fibers Cold fibers respond to a decrease in

    temperature, while warm fibers respond to anincrease in temperature.

    An individual fiber has a preferred temperature(exhibits a temperature tuning curve just likeV1 neurons are tuned for orientation)

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    Thermoreceptors

    Both cold and warm fibers dont respond to

    mechanical pressure.

    Preferred temperatures of cold fibers range from20 C to 45 C, while preferred temperatures of

    warm fibers range from 30 C to 48 C.

    This response to a preferred temperature is asustained response, it doesnt adapt like the

    additional increase/decrease response discussed

    previously.

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    Thermoreceptor tuning curves

    Explains paradoxical cold

    Jumping into a very hot bath transiently activatesboth hot and cold fibers.

    For a short time, the bath feels both hot and cold

    Cold

    Hot

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    Higher temperature processing

    Temperature information reaches the brain via the

    spinothalamic tract in the spinal cord.

    Its target in the brain is both Somatosensorycortex, but also Insular cortex.

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    SomatosensationTouch

    TemperaturePain

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    Pain as a special perceptual quality

    A quality that has complex phenomenologicalfacets (behavioral, sensory, emotional)

    Pain perception can be modulated by all kindsof factors, including behavioral states (stress,sex), cognitive states (hypnosis), mental states(trance), social norms and drugs.

    Its significance for society is enormous. E.g.:Should fishing be legal?

    Evolutionary significance to organism obvious.

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    Pain is not a purely sensory

    experience It shares components of an emotion (distress

    that is associated with pain).

    But it also shares sensory characteristics. Itsignals the presence of stimuli that are harmful

    to the organism.

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    Pain receptors

    Free nerve endings in the skin

    They are connected to various fibers:

    A

    small, myelinated. High conductance speed Csmall, unmyelinated. Slow conductance speed

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    Pain receptors

    Afast, sensitive to mechanical noxious stimuli.

    Cslow, sensitive to many noxious stimuli

    (chemical, etc.) This distinction has been used to explain the

    phenomenon ofdouble-pain:

    One noxious stimulus causes first a quick, sharp

    pain (mediated by A fibers) and is followed by a

    dull and burning pain (mediated by C fibers)

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    Pain pathways

    Pain information also

    reaches the brain via

    the spino-thalamic

    tract.

    There are projections

    to the Somatosensory

    cortex and theAnterior cingulate

    cortex

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    Cortical pain processing

    Sensory aspects of pain seem to be processed in the

    Somatosensory cortex.

    Emotional distress associated with pain seems to be processed

    in the Anterior Cingulate Cortex (ACC). Subjects with lesions in ACC could still accurately judge the

    intensity of pain. But they were not in the least bothered by it.

    On the other hand,

    subjects empathy forthe pain of others only

    elicits activity in

    ACC, not

    Somatosensory cortex.

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    The Gate control theory

    Devised by Melzack & Wall (1965)

    Tries to explain the fact that pain intensity can becognitively (top-down) controlled.

    Components of the model: Large fibers (involved in touch)

    Small fibers (involved in Pain perception)

    Substantia gelatinosa (the gate)

    T-cells (Transmission cells, sending pain signals tothe brain)

    If T-cells are sufficiently active, we experience pain.

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    The Gate control theory

    The large fibers are under cognitive control. If they are active, they activate

    the substantia gelatonisa (sg) in the spinal cord.

    If the sg is active, the activity ofT-cells is diminished.

    If the T-cells dont reach their activity threshold, no pain is experienced.

    If small fibers are active first, sg is inhibited, both fibers increase T-activity

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    Explanatory power of the theory

    Cognitive factors can diminish pain experience.

    Large fibers have descending tracts.

    Chemical Analgesics act to inhibit small fibers,disinhibiting the sg reducing pain.

    Rubbing the skin around a hurting area reduces pain

    by activating large (touch-sensitive) fibers, activating

    sg reducing pain.

    This effectively implements a peripheral pain control

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    Central control of pain

    There is a system of endogenous opiatesand corresponding receptors in the brain.

    Endogenous opiates are particularlyenkephalins and endorphins.

    In situations of stress, release of endorphinsreduces the expected pain centrally.

    Morphine also acts on these endogenousopiate receptors.