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

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no

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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



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



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%



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'



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



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



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



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



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



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



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



Ch. Ih. I

echanisms of sensitiveechanisms of sensitiveperiod plasticityeriod plasticity



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



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



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



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, !!!"



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



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



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



7uring sensitive period7uring sensitive period



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



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



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



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



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



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



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



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



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



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



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#



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



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



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*!


● overall pattern of resultsoverall pattern of results



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


● studies revealing visualstudies revealing visual



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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heory$heory$ %iving ontrol !ystems Publishing4%iving ontrol !ystems Publishing4

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Meta-Chemical Control of Brain and Behaviour .pdf