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Transcript of Meta-Chemical Control of Brain and Behaviour .pdf
8/14/2019 Meta-Chemical Control of Brain and Behaviour .pdf
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eta-chemicaleta-chemicalControl of Brainontrol of BrainV
Su
no
a
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Contentsontents Chapt rshapt rs
I S nsit iv p riods in t hS nsitiv p r iods i n th d v lopm nt of brain and v lopm n t of b rain and
b h aviour haviour
II P rc p tual control I P rc ptual control
th oryh ory
III P rc ptual c ontrol II P rc ptual control practicractic
Exampl sxampl s
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Ch. Ih. I
Sensitive period –Sensitive period – point inpoint inone's existence wone's existence when effecthen effect
of experience on the brain isof experience on the brain isparticularly strong during aparticularly strong during a
limited period inlimited period indevelopmentdevelopment
Critical period point–Critical period point– in one'sin one'sexistence whenexistence when experienceexperience
provides informationprovides information
essential for normalessential for normaldevelopment, alteringdevelopment, altering
performance permanentlyperformance permanently ensitive periods in theensitive periods in thedevelopment of Brain andevelopment of Brain and
Behaviourehaviour
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Ch. Ih. I
!
Although reflected in behavior,Although reflected in behavior,actually property of neural circuitsactually property of neural circuits
HYPOTH!"! # experience during aHYPOTH!"! # experience during asensitive period modifiessensitive period modifiesarchitecture of a circuit inarchitecture of a circuit in
fundamental ways, causing certainfundamental ways, causing certainpatterns of connectivity to becomepatterns of connectivity to become
highly stable and energeticallyhighly stable and energeticallypreferredpreferred
$nderstanding at circuit level, as$nderstanding at circuit level, aswell as relationship betweenwell as relationship between
circuit properties and behavior,circuit properties and behavior,provides deeper insight into criticalprovides deeper insight into critical
role experience plays in shapingrole experience plays in shapingdevelopment of brain and behaviordevelopment of brain and behavior
Sensitivity" #hat$s it allensitivity" #hat$s it allabout%bout%
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Ch. Ih. I
&
%earning that occurs during sensitive%earning that occurs during sensitiveperiods lays the foundation forperiods lays the foundation for
future learning and exerts long#future learning and exerts long#lasting influence on development oflasting influence on development ofindividual&s social and emotionalindividual&s social and emotional
behaviorbehavior
ritical periods # special class ofritical periods # special class ofsensitive periods that result insensitive periods that result in
irreversible changes in brain functionirreversible changes in brain function
experience must be of a particularexperience must be of a particular(ind and it must occur within a(ind and it must occur within a
certain period if the behavior is tocertain period if the behavior is todevelopdevelop
Sensitivity and initialensitivity and initiallearnin'earnin'
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Ch. Ih. I
(
behavior results frombehavior results frominformation that has beeninformation that has been
processed through hierarchiesprocessed through hierarchies
of neural circuitsof neural circuits
to define sensitive periods andto define sensitive periods and
to explore why they occur andto explore why they occur andhow they might behow they might be
manipulated, must thin( aboutmanipulated, must thin( aboutthem at the level of circuitsthem at the level of circuits
)he circuitry ofhe circuitry ofBehaviourehaviour
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Ch. Ih. I
*
)ot all circuits are shaped during)ot all circuits are shaped duringsensitive periodssensitive periods
*aintain high degree of plasticity*aintain high degree of plasticity
throughout life, such as basolateralthroughout life, such as basolateralnucleus of the amygdala, molecularnucleus of the amygdala, molecular
layer of the cerebellar cortexlayer of the cerebellar cortex
+hen a circuit can select from large+hen a circuit can select from large
range of potential patterns ofrange of potential patterns ofconnectivity effect of experienceconnectivity effect of experiencecan have an enormous impact on itscan have an enormous impact on its
connectivityconnectivity
onversely, when range of potentialonversely, when range of potential
patterns of connectivity is highlypatterns of connectivity is highlyconstrained by genetic influences,constrained by genetic influences,
effect of experience iseffect of experience iscorrespondingly smallcorrespondingly small
+penin' of sensitivepenin' of sensitiveperiods initial conditionseriods initial conditions
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Ch. Ih. I
,
- information provided to circuit- information provided to circuitmust be sufficiently reliable andmust be sufficiently reliable and
precise to allow the circuit to carryprecise to allow the circuit to carryout its functionout its function
.-.- circuit must contain ade/uatecircuit must contain ade/uateconnectivity 0both excitatory andconnectivity 0both excitatory and
inhibitory connections- to processinhibitory connections- to processthe informationthe information
1- activated mechanisms that enable1- activated mechanisms that enableplasticity2plasticity2
a- capacity for altering axonal ora- capacity for altering axonal or
dendritic morphologiesdendritic morphologiesb- ma(ing or eliminating synapsesb- ma(ing or eliminating synapsesc- changing strengths of synapticc- changing strengths of synaptic
connectionsconnections
Sensitive periodensitive periodprereuisitesrereuisites
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Ch. Ih. I
.
may be enabled bymay be enabled by
- progress of development- progress of development.- individual&s experience.- individual&s experience
intense impulse activity 0canintense impulse activity 0canresult from experience- shownresult from experience- shown
to trigger2to trigger2- gene transcription and- gene transcription and
translationtranslation.- activate existing molecular.- activate existing molecular
cascades for processescascades for processesunderlying plasticityunderlying plasticity
Sensitive periods timin'ensitive periods timin'of initiationf initiation
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Ch. Ih. I
/
xperience→xperience→
- customi3es a developing neural- customi3es a developing neuralcircuit to the needs of the individualcircuit to the needs of the individual
.- provides precise information.- provides precise informationabout the individual or aboutabout the individual or about
environment that often cannot beenvironment that often cannot bepredicted and, therefore, geneticallypredicted and, therefore, genetically
encodedencoded
1- calibrates the circuits that process1- calibrates the circuits that processstereoscopic information andstereoscopic information and
customi3es circuits involved incustomi3es circuits involved in
processing speech sounds for theprocessing speech sounds for theparticular language0s- that will beparticular language0s- that will bespo(enspo(en0urin' a sensitive periodurin' a sensitive period
experience and sens-xperience and sens-
period plasticityeriod plasticity
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Ch. Ih. I
1
predisposition of a circuit topredisposition of a circuit tobe instructed by typicalbe instructed by typical
experience reflects2experience reflects2
- selectivity of the circuit&s- selectivity of the circuit&svarious inputs 0may bevarious inputs 0may beshaped during sensitiveshaped during sensitive
periods-periods-.- innate connectivity of the.- innate connectivity of the
circuit4circuit4 2eflectionseflections
of the circuitf the circuit
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Ch. Ih. I
3
- axon elaboration and synapse- axon elaboration and synapseformationformation
.- axon and synapse elimination.- axon and synapse elimination
0shown to alter circuit0shown to alter circuitarchitecture during sensitivearchitecture during sensitive
periods-periods-1- synapse consolidation1- synapse consolidation
* Hebbian rule - activity of a* Hebbian rule - activity of atentative, presynaptic elementtentative, presynaptic element
consistently anticipates (contributesconsistently anticipates (contributes
to driving) the activity of ato driving) the activity of a
postsynaptic neuron, that synapse ispostsynaptic neuron, that synapse is
stabilized and strengthened stabilized and strengthened
distribution of stabili3eddistribution of stabili3edsynapses shapes growth patternssynapses shapes growth patterns
of axons and dendritesof axons and dendrites
echanisms of sensitiveechanisms of sensitiveperiod plasticityeriod plasticity
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Ch. Ih. I
echanisms of sensitiveechanisms of sensitiveperiod plasticityeriod plasticity
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Ch. Ih. I
!
5Hypothesis # experience during a5Hypothesis # experience during asensitive period potentiatessensitive period potentiates
specific synapses which arespecific synapses which arestructurally stabili3ed by thestructurally stabili3ed by theinsertion of particular (inds ofinsertion of particular (inds of
A*sA*s
synapses become invulnerable tosynapses become invulnerable toelimination, even if functionalelimination, even if functionalefficacy was to drop to 3eroefficacy was to drop to 3ero
persistence suggests they becomepersistence suggests they become
relatively invulnerable torelatively invulnerable to
elimination, because they haveelimination, because they havebeen consolidated by a particularbeen consolidated by a particular
(ind of A*(ind of A*
echanisms ofechanisms ofarchitectural chan'erchitectural chan'eunderlyin' Sensitivenderlyin' Sensitive
periodseriods
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Ch. Ih. I
&
has a uni/ue advantage inhas a uni/ue advantage in
shaping the connectivity of ashaping the connectivity of a
circuitcircuit
"ntense and repeated activation"ntense and repeated activationalters the circuit conditionsalters the circuit conditions
dramaticallydramatically
Although may have a uni/uelyAlthough may have a uni/uelypotent effect in shaping patternspotent effect in shaping patterns
of connectivity, subse/uentof connectivity, subse/uentexperience has the ability toexperience has the ability to
cause further structural andcause further structural andfunctional changes as long as thefunctional changes as long as the
sensitive period remainssensitive period remainsopenopen
4niue advanta'e ofniue advanta'e ofinitial experiencenitial experience
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Ch. Ih. I
(
4niue advanta'e ofniue advanta'e ofinitial experiencenitial experience
The pattern of connectivityThe pattern of connectivityinstructed by experienceinstructed by experience
becomes more sharply definedbecomes more sharply definedand highly preferred, evenand highly preferred, eventhough the pattern may bethough the pattern may be
atypicalatypical
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Ch. Ih. I
*
Secondaryecondary
experiencexperience
instructs a new pattern of connectivity,instructs a new pattern of connectivity,
extra energy must be expended toextra energy must be expended to
overcome the influences that stabilize theovercome the influences that stabilize the
initial patterninitial pattern
Repeated experience that instructs newRepeated experience that instructs new
pattern of connectivity refines andpattern of connectivity refines and
stabilizes the new pattern, creating a newstabilizes the new pattern, creating a new
low point in the landscapelow point in the landscape
humans are able to learn multiple languageshumans are able to learn multiple languages
with equal facility during a sensitive periodwith equal facility during a sensitive period
(Doupe & Kuhl, !!!"(Doupe & Kuhl, !!!"
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Ch. Ih. I
,
6666stability landscape&&#stability landscape&&#
functional implications of thefunctional implications of thecellular and molecular eventscellular and molecular eventsfor the future performance offor the future performance of
a circuita circuit
represents the range ofrepresents the range ofpossible connectivity patternspossible connectivity patterns
circuit might ac/uire andcircuit might ac/uire anddegree to which anydegree to which any
particular pattern is preferredparticular pattern is preferred
5Stability landscape6 as atability landscape6 as ametaphor for sensitiveetaphor for sensitive
period plasticityeriod plasticity
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Ch. Ih. I
.
Endin' ofndin' of
sensitive periodsensitive periods
change may still occur 0as longchange may still occur 0as long
as not a critical period- but extraas not a critical period- but extraenergy re/uired for a circuit toenergy re/uired for a circuit to
maintain a less stable pattern ofmaintain a less stable pattern ofconnectivityconnectivity
critical period ends # alternativecritical period ends # alternativepatterns of connectivity are nopatterns of connectivity are no
longer possible due to thelonger possible due to theproperties of the circuit&sproperties of the circuit&s
stability landscapestability landscape
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Ch. Ih. I
/
# 0as a function of developmental# 0as a function of developmentalstage- involve circuits that havestage- involve circuits that havethe potential to learn multiple,the potential to learn multiple,
stable patterns of connectivitystable patterns of connectivityduring the sensitive periodduring the sensitive period
that end rapidly involve circuitsthat end rapidly involve circuitswith strong innatewith strong innate
predispositions to be shaped bypredispositions to be shaped bycertain (inds of stimulicertain (inds of stimuli
These circuits learn to respond toThese circuits learn to respond toa particular stimulus # once thea particular stimulus # once thecircuit ac/uires selectivity forcircuit ac/uires selectivity for
that stimulus, subse/uentthat stimulus, subse/uentexperience has little or no effectexperience has little or no effect
Endin' ofndin' ofsensitive periodsensitive periods
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7uring sensitive period7uring sensitive period
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Ch. Ih. I
!3
7uring sensitive period,7uring sensitive period,particular (inds of experienceparticular (inds of experience
shape the connectivity of ashape the connectivity of a
circuit in fundamental ways,circuit in fundamental ways,
causing certain patterns ofcausing certain patterns ofconnectivity to becomeconnectivity to become
energetically preferred orenergetically preferred ormechanistically specifiedmechanistically specified
ritical periods # a subset ofritical periods # a subset ofsensitive periods for which thesensitive periods for which the
instructive influence ofinstructive influence of
experience is essential forexperience is essential for
typical circuit performancetypical circuit performance0effects of experience on0effects of experience on
performance are irreversible-performance are irreversible-
In 2etrospectn 2etrospect
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Ch. IIh. II
!
● +illiam T4 08ill- Powers began+illiam T4 08ill- Powers beganin early 9:;s by applyingin early 9:;s by applyingcontrol engineering andcontrol engineering and
natural science to the sub<ectnatural science to the sub<ectof psychologyof psychology
● ontrol theory explains howontrol theory explains howorganisms can control whatorganisms can control what
happens to themhappens to them
● explains what a goal is, howexplains what a goal is, howgoals relate to behavior, howgoals relate to behavior, how
behavior affects perceptions,behavior affects perceptions,how perceptions define realityhow perceptions define realitywe live in and move and havewe live in and move and have
our beingour beingerceptual control erceptual control
in theoryn theory
● behavior "! orderlybehavior "! orderly
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Ch. IIh. II
!!
● behavior "! orderlybehavior "! orderly
● O)!=$)! of output forcesO)!=$)! of output forcesare shaped by organism intoare shaped by organism into
highly regular and reliablyhighly regular and reliablyrepeatable states and patternsrepeatable states and patterns
● "*PO>TA)T2 how much you (now"*PO>TA)T2 how much you (nowabout the outcomes that are underabout the outcomes that are under
controlcontrol
● confusion is all in the eye of theconfusion is all in the eye of thebeholderbeholder
● ach person controls oneach person controls one
contribution to the pattern that allcontribution to the pattern that allperceive, in such a way as toperceive, in such a way as to
preserve the higher#level patternpreserve the higher#level patternas each person desires to see itas each person desires to see it
+n the obviousn the obvious
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Ch. IIh. II
!&
● once we understood theonce we understood theorderliness of simple acts and theirorderliness of simple acts and their
immediate conse/uences, weimmediate conse/uences, weshould be able to go on andshould be able to go on and
understand more general patternsunderstand more general patterns
● +e must explore all levels # what+e must explore all levels # whatwe find at each level ma(es sensewe find at each level ma(es senseonly in the context of the othersonly in the context of the others
● PT2PT2● focuses on how we loo( at andfocuses on how we loo( at and
experience things, and the wayexperience things, and the waythese perceptions are comparedthese perceptions are compared
with experiences we wantwith experiences we want● explains how thoughts becomeexplains how thoughts becomeactions and feelings and whyactions and feelings and why
stimuli appear to cause responsesstimuli appear to cause responses
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Ch. IIh. II
!(
● ProposesProposes # nervous system is# nervous system ismade up of large number ofmade up of large number of
control systems in acontrol systems in a
hierarchical arrangement, eachhierarchical arrangement, eacha simple circuit of neuronsa simple circuit of neuronswhich /uic(ly and efficientlywhich /uic(ly and efficientlycan perform the way we docan perform the way we do
●
OffersOffers # explanation# explanation of howof howpurposeful behavior wor(s andpurposeful behavior wor(s and
what it accomplisheswhat it accomplishes
● xplainsxplains # behavior from inside# behavior from insideperspective of controllingperspective of controllingorganism rather than fromorganism rather than from
outside perspective ofoutside perspective ofobserverobserver
People as livin' thin's7eople as livin' thin's7he story
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Ch. IIh. II
!*
● ontrol of input by means ofontrol of input by means ofoutput # defining characteristicoutput # defining characteristic
distinguishing living things fromdistinguishing living things from
inanimate ob<ectsinanimate ob<ects● ssence 0of PT method- # tossence 0of PT method- # to
apply a force, stimulus orapply a force, stimulus or
disturbance to an ob<ect anddisturbance to an ob<ect and
observe the result or reactionobserve the result or reaction
● "f a force?stimulus is applied to"f a force?stimulus is applied toa variable that a living organisma variable that a living organismperceives and is controlling, theperceives and is controlling, the
organism producesorganism producescountervailing forces tocountervailing forces to
maintain that variable in statesmaintain that variable in statesthat it prefersthat it prefers
eople as livin' thin's7eople as livin' thin's7he story
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Ch. IIh. II
!,
5 a theory of human5 a theory of human
communication is an extensioncommunication is an extension
of our efforts to explain brainof our efforts to explain brainand behaviorand behavior
our beliefs about the brainour beliefs about the brainshould have direct implicationsshould have direct implications
for our characteri3ation offor our characteri3ation oflanguage 0vice versa-language 0vice versa-
Hierarchical Perceptual ontrolHierarchical Perceptual ontrol
Theory 0HPT- one of–
Theory 0HPT- one of–
theories addressing the brain#theories addressing the brain#language connection islanguage connection is8an'ua'e7 control ofan'ua'e7 control of
erception
Powers& insight was toPowers& insight was to
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Ch. IIh. II
!.
Powers insight was toPowers insight was to
emphasi3e the role of behavior inemphasi3e the role of behavior inperceptual controlperceptual control
language is <ust as purposeful aslanguage is <ust as purposeful asother behaviorsother behaviors
# %anguage forms the basis for# %anguage forms the basis for
most of our complexmost of our complex
interactions44allows for possibilityinteractions44allows for possibilityof resolving disputes andof resolving disputes and
conflicts without resorting toconflicts without resorting toviolenceviolence
@@higher level variability inhigher level variability inmorphology and syntax may bemorphology and syntax may be
crucial, even life#threateningcrucial, even life#threatening0+hat did you call meBC-0+hat did you call meBC-
8an'ua'e7 control ofan'ua'e7 control oferception
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Ch. IIh. II
!/
HPT offers a recast ofHPT offers a recast ofimportant /uestions aboutimportant /uestions about
individual linguistic behaviourindividual linguistic behaviour
# "f there is a hierarchy to"f there is a hierarchy toperception in general, then theperception in general, then thesame principles may apply tosame principles may apply to
language as welllanguage as well
# "f behavior is the control of# "f behavior is the control of
perception, then linguisticperception, then linguistic
behavior may be viewed asbehavior may be viewed ascontrol of perception as wellcontrol of perception as well
8an'ua'e7 control ofan'ua'e7 control oferception
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Ch. IIh. II
!1
Any discussion of developmentAny discussion of developmentalso raises the /uestion ofalso raises the /uestion of
critical periods2 is there one orcritical periods2 is there one or
more such periods for humanmore such periods for humanlanguageBlanguageB
A true critical period forA true critical period for@language would involve@language would involve
virtually the entire brainCvirtually the entire brainC
Alternative # multiple critical orAlternative # multiple critical orsensitive periods for2sensitive periods for2
- configuration#related- configuration#relatedperceptual developmentperceptual development
.- transition#related perceptual.- transition#related perceptualdevelopment etc4development etc4erceptual control erceptual control
in theoryn theory
%inguistic behaviors result from%inguistic behaviors result from
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Ch. IIh. II
&3
%inguistic behaviors result from%inguistic behaviors result fromthe interplay of various contextualthe interplay of various contextual
and neurological factorsand neurological factors0in PT terms, reference levels and0in PT terms, reference levels and
disturbances to them-disturbances to them-
The semantics for ob<ects or forThe semantics for ob<ects or forconcepts we use changes as weconcepts we use changes as we
gain relevant experiencesgain relevant experiences
our DimplicitE understanding ofour DimplicitE understanding oflanguage grows and changeslanguage grows and changes
through our experiencesthrough our experiences
These experiences form the basisThese experiences form the basisfor our decisions regarding suchfor our decisions regarding such
things as word choice 0categories-things as word choice 0categories-or lexical ordering 0se/uence-4or lexical ordering 0se/uence-4
an'ua'e and sub9ectivean'ua'e and sub9ectiveperceptionserceptions
# move beyond @word## move beyond @word#
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Ch. IIh. II
&
move beyond word move beyond wordmanipulation to completelymanipulation to completely
understand connection betweenunderstand connection between
language and behaviorlanguage and behavior
primary interest in effect weprimary interest in effect wedesire to produce upon thedesire to produce upon the
listener and so are usually notlistener and so are usually not
attentive to the processes byattentive to the processes by
which we produce the effectwhich we produce the effect00Pillsbury and *eaderF 8a(htin andPillsbury and *eaderF 8a(htin and
Gygots(y-Gygots(y-
44@purpose, @volition, @will,44@purpose, @volition, @will,@motive, 44whatever this guiding@motive, 44whatever this guiding
force may be, is ultimateforce may be, is ultimatedictator of the whole process ofdictator of the whole process of
speech 07un(el 9I-speech 07un(el 9I-
8an'ua'e and sub9ectivean'ua'e and sub9ectiveperceptionserceptions
● neural processing in low#levelneural processing in low#leveli lti b d l t d b
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Ch. IIIh. III
&!
cortices can be modulated bycortices can be modulated byinteractions between senses atinteractions between senses at
specific, early poststimulus timespecific, early poststimulus timeperiodsperiods
● early onset of cortical responses inearly onset of cortical responses inhumans to auditory relative tohumans to auditory relative to
visual stimulation ma(es itvisual stimulation ma(es itconceivable that auditory processesconceivable that auditory processes
can impact feedforward visualcan impact feedforward visualprocesses within nominally visualprocesses within nominally visualcortices 0!chroeder et al4, .;;-4cortices 0!chroeder et al4, .;;-4
● evidence of auditory#drivenevidence of auditory#drivenresponses within inferior parietalresponses within inferior parietal
cortex of humans 0!chroeder et al4,cortex of humans 0!chroeder et al4,.;;- that represent a potential.;;- that represent a potential
basis for modulation of earlybasis for modulation of earlysensory processing within visualsensory processing within visual
cortexcortex
Perceptual control erceptual control in practicen practice
● auditory inputs particularlyauditory inputs particularly
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Ch. IIIh. III
&&
well poised to impact visualwell poised to impact visual
processing not only at theprocessing not only at the
lowest cortical levels butlowest cortical levels but
potentially at poststimuluspotentially at poststimuluslatencies 0as short as .; ms-latencies 0as short as .; ms-
● 4g4 occipital T*! has4g4 occipital T*! has
opposing effects on visualopposing effects on visualand auditory stimulusand auditory stimulusdetection over identical timedetection over identical time
periodsperiods
●
external auditory stimuliexternal auditory stimulienhance visual cortexenhance visual cortex
excitability as measured viaexcitability as measured viaT*!T*!
Perceptual control erceptual control in practicen practice
● overall pattern of resultsoverall pattern of results
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Ch. IIIh. III
&(
indicates that there is aindicates that there is a
period during whichperiod during which
behavioral conse/uences ofbehavioral conse/uences of
occipital T*! depend on theoccipital T*! depend on thesensory modality of externalsensory modality of external
inputsinputs
●
results support the existence ofresults support the existence ofearly critical period for producingearly critical period for producing
facilitative multisensoryfacilitative multisensory
interactions and suggestinteractions and suggest
auditory and visual inputs areauditory and visual inputs are
converging ithin the occipitalconverging ithin the occipital
cortex cortex
Perceptual control erceptual control in practicen practice
● studies revealing visualstudies revealing visual
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Ch. IIIh. III
&*
studies revealing visualgsuppression by occipital T*!suppression by occipital T*!
0Amassian et al4, 9I9-0Amassian et al4, 9I9-
● showed that stimulation of ashowed that stimulation of agiven area can induce opposinggiven area can induce opposing
0disruptive or facilitative-0disruptive or facilitative-
effects depending on tas( oreffects depending on tas( orbehavioral context 0+alsh et al4,behavioral context 0+alsh et al4,99IF Jrosbras and Paus, .;;.,99IF Jrosbras and Paus, .;;.,
.;;1-.;;1-
● results imply that visualresults imply that visual
perception and brain plasticityperception and brain plasticity
can be heavily influenced by acan be heavily influenced by a'targeted' auditory stimulus,'targeted' auditory stimulus,
0especially under occipital T*!-0especially under occipital T*!-erceptual control erceptual control in practicen practice
)h :$;)h :$;
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)han: ; )han: ; Very Muchly Very Muchly
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