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Brain and behavior correlates of intrinsic motivation and skill learning
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Brain and behavior correlates of intrinsic motivation and skill learning
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! ! !
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!
BRAIN&AND&BEHAVIOR&CORRELATES&OF&INTRINSIC&MOTIVATION&AND&SKILL&
LEARNING&
!
By!
!
Dorsa!Beroukhim!Kay !
!
!
A!Dissertation!Presented!to!the !
FACULTY!OF!THE!USC!GRADUATE!SCHOOL!
UNIVERSITY!OF!SOUTHERN!CALIFORNIA!
!
In!partial!fulfillment!of!the!requirements! for!the!degree !
DOCTOR!OF!PHILOSOPHY!
(NEUROSCIENCE)!
!
December!2017!
!
!
!
Copyright!2017 ! ! ! ! ! ! ! Dorsa!Beroukhim!Kay ! !
!
2!
!
ACKNOWLEDGMENTS!
!
The!experience!to!come!to!America!and!have!the!unique!opportunity!to!spend! six!
years!embracing!the!field!of!knowledge!and! philosophy !has!been!enriching,!both!
intellectually!and!spiritually.!I!feel!blessed!to!have!been!able!to!engage!in!research!and!
contribute!to!brain!science!in!such!an!exciting!time !of!new!discoveries!and!technologies !
in!neuroscience .!!
I!would!like!to!thank! my!advisor!Dr.!Carolee!Winstein!for!her!guidance! and!
mentorship!throughout!my!time!at!USC.!In!particular,!she!trained!me!as!a!scientist!to !
develop!my!thinking!and! execute!my!dissertation!study!from!experimental!design!to!
data!analysis!and!interpretation!of!my!results.!I!am!also!grateful!to!my!dissertation!
committee!members,!Dr.!John!Monterosso,!Dr.!Rebecca!Lewthwaite!and!Dr.!Jason!
Kutch ,!for!sharing!their!expertise,! engaging!in!discussions!with!me !and!for!their!valuable!
suggestions!regarding!my!dissertation.!!
I!would!also!like!to!thank!the!USC!CTSI!TL1!program,!the!James!H.!Zumberge!
Research!and!Innovation!Fund,!the!Neuroscience!Graduate!Program,!and!the!Division!
of!Biokinesiology!and!Physical!Therapy!at!University!of!Southern!California!for!their!
financial!support!of!this!research. !
I!appreciate!and!feel!so!lucky!for!my!lab!mates!at!the!Motor!Behavior!and!
Neurorehabilitation!Laboratory!and!in!the!Neuroscience!Graduate!Program!for!creating!
an!environment!for!research!and!collaboration!infused!with!friendship,! kindness!and!
support.!!
!
3!
!
A!big!thank!you!goes!to!my!husband ,!David!Kay,!for!giving!me!the!support,!
strength!and!encouragement!to!finish!my!PhD.!It!is!you!who!kept!me!motivated!about!
the!value!of!what!I!had!to!contribute,!even!when!I!felt!doubtful.!Thank!you! for!your!
unwavering!love!and!patience!as!I!completed!all!the!phases!of!my!project,!through!the!
juggle!of!building!and!raising!a!family.!I!love!you!and!you!are!my!shining!other!half.! !
My!mother!and!father ,!Juliet!and!Rahmatollah!Beroukhim, !deserve!a!great!level!
of!gratitude!for!their!support!and!encouragement!to!choose!this!journey!to!pursue!my!
PhD.!Maman,!thank!you!for!always!taking!time!to!make!sure!I!had!what!I!needed!to!
thrive!throughout!my!undergraduate!and!graduate!studies.!From!your!warm!and!
delicious!cooking!to!your!kind!care!of!Joshua,!I!truly!feel!like!a!part!of!this!degree!is!
yours.!Baba,!thank!you!for!encouraging !me!and!inspiring!me!to!pursue!a!career!in!
research!and!education.!Thank!you!for!always!listening!to!my!enthusiastic!ideas!and!
reflections!about!life!and!engaging!in!discussions!about!science!and!how!the!world!
works!from!when!I!was!young! girl.!Your!passion!to!learn!and!your!appreciation!for!
knowledge!have!driven!my!pursuit!for!truth!in!every!aspect!of!my!life.!!I!hope!to!continue!
to!make!you!both !proud!in!the!next!stage!of!my!career. !
A!special!thank!you!to!my!two!awesome!brothers,!David!and!Ariel!Beroukhim.!I!
feel!incredibly!fortunate!for!the!heartfelt!love!and!closeness!I!receive!from! you!in!every!
aspect!of!my!life,!and!specially!in!pursuing!my!career!path.!A!grand!merci!and!tashakor!
to!my!mamani!and!maman!bozorg,!my!two!grandmothers!who!I!am!proud!to!call!my!
family’s!matriarchs !and!my!aghajan!and!babajoon,!my!two!grandfathers!whose!memory !
bring!our!family!continued!blessings.!It!is!to!their!testament!that!they!raised!children!to!
!
4!
!
value!education!and!respect!the!time!investment!it!requires,!that!I!was!able!to!choose!to!
pursue!higher!education!in!an!academic!field!that!I!was!personally!passio nate!about.!
I!am!also!sincerely!grateful!for!the!encouragement!of!my!family!in!law,!and!in!
particular!my!mother!and!father!in!law,! Mrs.!Marilyn!and!Dr.!Martin!Kay,!for!their!loving!
support!that!made!completing!my!PhD!possible. !
!I!want!to!share!a!warm!thank !you!to!my!circle!of!friends!and!community!for!their!
love!and!cheer!throughout!the!past!six!years.!I!am!grateful!for!their!friendship!and!
companionship!in!every!stage!of!my!life!and! look!forward!to!continue!sharing !in!joys!and!
achievements !together. !
!
!
“The%happiness%of%the%journey%is%the%anticipation%of%the%destination.%
%The%happiness%of%the%destination%is%the%appreciation%of%the%journey.”%%
6%Rabbi%Dr.%Akiva%Tatz%
! !
!
5!
!
TABLE!OF!CONTENTS!
!
Acknowledgments! 2!
List!of!figures!and!tables! 6!
Abstract! 7!
Chapter! 1:!Overview! 8!
Chapter! 2:!How!does!motivation!modulate!behavior?!The!neural!basis!of!
motivation!related!benefits!on!motor!performance!and!learning:!A!systematic!
review!! 10!
!
Chapter! 3:!!Intrinsic!motivation!promotes!motor!skill!learning!through!
strengthening! of!cingulate!cortex!activity!during!feedback!processing!after!
good!performance!trials! 28!
!
!
Chapter! 4:!Summary!and!Conclusion! 64!
!
!
Appendix! !
SelfcEfficacy!Q uestionnaire!items! 67!
Intrinsic!Motivation!Inventory!items! 68!
Exit!Interview!Questionnaire!Items! 69!
Edinburgh!Handedness!Inventory ! 70!
Supplementary!Material! !
Room!Mean!Square!Error!(RMSE)!performance!outcome!measure:!
Methods!and!R esults! 71!
Exit!Interview!Questionnaire:!Methods!and!Results ! 72!
!
References! 73!
!
!
!
6!
!
LIST!OF!FIGURES!AND!TABLES!
Chapter&2:!How!does!motivation!modulate!behavior?!The!neural!basis!of!
motivation!related!benefits!on!motor!performance!and!learning:!A!
systematic!review& !
Figure!1.!PRISMA!outflow!of!systematic!review! 24!
Table!1.!Summarizing!methods!and!brain!results!for!impact!o f!motivation!
on!motor!behavior.! ! 26!
!
Chapter&3:!!Intrinsic!motivation!promotes!motor!skill!learning!through!
strengthening!of!cingulate!cortex!activity!during!feedback!processing!after!
good!performance!trials& !
Table!1.!Participant!characteristics ! 56!
Table!2.!Main!activation!at!the!time!of!tracking!task!performance ! 56!
Figure!1.!Study!Procedures,!Tracking!task!and!Trial!Sequence ! 57!
Figure!2.!Average!selfcefficacy!reports!across!time!points!and!intrinsic!
motivation!inventory!subscales!ratings! 58!
Figure!3.!Task!Performance! 59!
Figure!4.!Relationship!of!self cefficacy!change!and!motor!performance!at!
immediate!and!delayed!retention!for!motivation!and!control!group! 60!
Figure!5.!Brain!image!for!taskcrelated!activity!across!all!participants!
(combined!groups)!! 61!
Figure!6.!Brain!activity!for!control !group>motivation!group!contrast!during!
feedback!presentation,!modulated!by!time!lag!performance!!(associated!
with!long!time!lag) ! 62!
Figure!7.!Brain!activity!across!all!par ticipants!during!feedback!presentation ,!
modulated!by!time!lag!performance!!(associated!with!short!time!lag) !
! 63!
Figure!8.!Brain!activity!for!motivation!group>control!group!contrast!during!
feedback!presentation,!modulated!by!time!lag!performance!(associate d!
with!short!time!lag) !
! 63!
&
!
7!
!
ABSTRACT!
!
Motivationcrelated!boosts!in!motor!performance,!using!experimental!manipulations!that!
harness!extrinsic!sources!of!motivation,!have!been!demonstrated!to!activate!brain!
regions!within!the!dopaminergic!system.!Little!is!known!about!the!neural!networks!
driving!intrinsic!motivation!benefits!on!motor!behavior.!To!investigate!the!brain!
correlates!of!intrinsically!motivated!motor!skill!learning,!we!used!fMRI!to!examine!brain!
activity!during!acquisition!of!a!motor!skill,!using!a!pinch!force!tracking!task,!in!a!grou p!of!
young!adults!that!practiced!under!enhanced!motivation!or!neutral!conditions.!Motor!
learning!was!assessed!immediately!after!and!one!day!after!practice,!with!no cfeedback!
retention!test!trials.!Activation!in!the!cingulate!cortex!in!response!to!performanc e!
feedback!during!practice,!was! more!strongly!associated!with!good!performance!trials! in!
the!motivation!group!compared!to!the!contro l!group.!The!motivation!group! displayed!
better!motor!learning!compared!to!the!control!group,!as!indexed!by!reduced!temporal!
error!in!tracking!the!target!during!practice,!as!well!as!during!immediate!and!delayed!
retention!tests.!These!findings!provide!evidence!that! motor!skill!learning!is!positively!
modulated!by!motivational!conditions,!partly!mediated!by!changes!in!neural!activa tion!in!
response!to!feedback!in!areas!relevant!to!the!salience!network. !
!
!
!
!
!
!
8!
!
Chapter!1 :!Overview!
Brain&Behavior&Correlates&of&Intrinsic&Motivation&and&Motor&Skill&Learning& &
Motivation!colors!our!daily!life!experiences!and!affects!how!we!think,!feel!and!
behave.!The!study!of!motivation!has!generated!various!conceptual!models!to!explain!its!
relationship!and!impact!in!the!psychological,!cognitive!and!motor!domains! (Bandura,!
1977h!Dweck,!1986h!Ryan!and!Deci,!2000).!It!is!well!established!that !experimental!
methods!that! enhance !motivation!influence!the!energization,!direction!and!intensity!of!
behavior ,!with!a!majority!of!studies!showing!a!beneficial!effect!on!motor!skill!learning!
(Wulf!and!Lewthwaite,!2016) .!Investigations!of!the!neural!circuits!that!mediate!
motivational!influence!on!motor!skill!performance!are!beginning!to!shed!light!on!relevant!
brain!regions!and!processes.!Chapter! 2!is!a!systematic!review!of!studies!that!use!
neuroimaging!techniques!to!better!understand!how!motivation!affects!motor!behavior!
and!the!associated!brain!correlates.!Less!well!understood!is!how!support!of!
fundamental!psychological!needs!that!g enerates!motivation,!including!autonomy,!
competence!and!social!relatedness,! mediate!braincmotor!behavior!effects.! !
This!dissertation!uses! functional!magnetic!resonance!imaging!(fMRI) !to!
investigate!the !brain!and!behavioral!effects!of!instructions!that!enhance!a!learner’s!
expectation!of!performance!success!and!supports!competence,!an!experimental!
manipulation!used!to!increase!motivation.!Ch apter!3!describes!the!behavioral!findings!
from!the! dissertation!study!and!provides!evidence!for!the!associated!brain!networks!that!
are!involved!in!motivated!conditions!of!motor!skill!learning.!Chapter!4!provides!an!
overall!summary!of!the!results!from!Chapter!2 ccthe!systematic!review!and!Chapter!3 cc
!
9!
!
the!findings!from!thi s!dissertation!study!and!offers!future!directions!for!this!line!of!
research.! !
!
! !
!
10!
!
Chapter!2:!How!does!motivation!modulate!behavior?!The!neural!basis!of!motivation!
related!benefits!on!motor!performance!and!learning:!A!systematic!review !
Abstract&
Motivation!influences!behavior! across!a!spectrum!of!cognitive!and!motor!
domains.!Behavioral!studies!have!demonstrated!beneficial!effects!of!enhanced!
motivation!on!motor!performance!and!learning.!!What!is!not!well!understood!are!the!
brain!networks!that! underlie!or!drive!motivated!motor!behavior!and!promote!motor!skill!
learning.!We!conducted!a!systematic!r eview!of!published!studies!that!examined !
motivation,!used!a!motor!task!and!reported!braincrelated!findings!in!a!population!of!nonc
disabled!adults.!!We!applied!search!ter ms!to!the!PubMed!database,!resulting!in!181!
citations.!After!a!three!stage!screening!process,!nine!studies!were!selected.!Study!
characteristics,!including!methods!and!main!findings!were!extracted!and!described.!!
Results!from!this!review!show!a!critical!rol e!for!the!striatum!and!the!salience!network!in!
associated!neural!correlates!of!reward cenhanced!motor!behavior.!!Future!research!and!
implications!are!discussed.! !
Introduction&
Motivation!plays!a!profound!role!in!how!we!behave!in!our!everyday!life.!Whether!
we!are!learning!a!new!language,!choosing!the!route!we!take!to!work!or!picking!up!a!
new!sport,!our!motivation!for!each!experience!fuels!how!we!act!to!achieve!our!objective.!
For!the!purpose!of!this!review,!we!will!broadly!define!motivation!as!the!psychological !
drive!for!energization!and!intensity!of!behavior.! Sources!of!motivation!can!be!classified!
!
11!
!
as!extrinsic,!such!as!monetary!incentives!or!intrinsic,!such!as!a!sense!of!competence!or!
autonomy!(Wood!and!Bandura,!1989h!Ryan!and!Deci,!2000).!!
A!body!of!literature!has!demonstrated!an!impact!of!both!extrinsic!and!intrinsic!
motivation!on!behavior!using!a!range!of!motivational!manipulations!including!monetary!
reward!(Abe!et!al.,!2011h!Galea!et!al.,!2015),!performance!achievement,!instructions!
that!support!autonomy!and!competence! (Wulf!and!Lewthwaite,!2009h!Wulf!et!al.,!2012,!
2014),!social!comparison !(Lewthwaite!and!Wulf,!2010) ,!and!praise!(Sugawara!et!al.,!
2012).!The!majority!of!published!studies!have!demonstrated!an!enhancing!effect!of!
motivation!on!task!performance!and!learning.!!
The!study!of!the!neural!basis!for!the!positive!effects!of!motivation!on!general!
behavior!has!been!approached!from!numero us!directions,!distinguishing!intrinsic!and!
extrinsic!reward!processing! (Lutz!et!al.,!2012h!Widmer!et!al.,!2016),!appetitive!and!
aversive!(reward/punishment)!mechanisms! (Wrase!et!al.,!2007h!Wachter!et!al.,!2009) !
and!expectation!and!receipt!of!reward! (Knutson!et!al.,!2001).!Furthermore,! these!
studies!have!been!executed!in!various!behavioral!paradigms,!including!decision!
making,!cognitive!task!performance!and!motor!behavior,!which!is!the!focus!of!this!
review.!The!literature!specifically!investigating!the!brain!networks!and!processes!that!
modulate!the!impact!of!motivation!on!motor!behavior!improvements!shed!light!on!the!
associated!neural!mechanisms.! !
The!goal!of!this!article!is!to!review!findings!from!human!neuroimaging !studies!
that!examine!the!impact!of!motivation!on!brain!processes!during!motor!behavior.!In!the!
present!review,!we!discuss!implicated!neural!mechanisms!that!mediate!the!energizing!
!
12!
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effect!of!behavior!by!enhanced!motivation!and!propose!a!hypothesis!that!can!be!teste d!
to!further!our!understanding!of!the!role!of!neural!correlates!of!motivation!in!motor!skill!
behavior!in!health!and!disease. !
&
Methods!!!
Search&strategy&
We!did!an!electronic!search!of!1!online!database!(PubMed)!through!July!2017.!
We!searched!the!literature! to!include!studies!that!examined!the!effects!of!motivation!on!
motor!performance!and!reported!braincrelated!outcome!measure!in!humans.!We!
performed!a!search!with!the!following!terms!indicating!motivation:!intrinsic!motivation,!
extrinsic!motivation!or!rewardh!combined!with!the!following!terms!to!indicate!motor!
behavior:!behavioral!performance,!motor!performance,!skill!learning!or!motor!learningh!
and!the!f ollowing!terms!indicating!braincrelated!outcome!measures:!brain,!neural,!
cortex,!fMRI,!neuroimaging,!EEG,!or!TMS.!
&
Article&selection&
Inclusion&criteria&
We! included! all! studies! that ! are! published! articles! in! peer creviewed!
journals,! and! include! 1)! subjects! of! non cdisabled! adults,! 2)! have! a! primary!
outcome! measure! of! brain! activity! or! neural! process,! 2)! report! motor!
performance,!3)!report!motivation!that!has!been!manipulated!or!assessed ,!and!
are!present!in!PubMed.!
!
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13!
!
Exclusion&criteria&
We!excluded!articles!that!did!not!meet!the!inclusion!criteria,!unpublished!
studies,!dissertations,!abstracts,!and!review!articles.!
!
Screening&
Each!abstract!was!independently!screened!by!three!primary!reviewers!
(DBK,!OK,!NB)!to!identify!those!that!met!inclusion!and!exclusion!criteria!and!was!
rechecked!by!the!senior!author!(DBK).!The!selected!articles!were!then!read!in!
depth!by!the!sen ior!author!to!confirm!inclusion!criteria!were!met!and!examine!
reported!findings.!
!!
Data&extraction&
We!extracted!data!on!the!study!design,!number!of!participants,!study!population,!
motor!task!used,!manipulation!or!assessment!of!motivation!reported,!braincrelated!
measure!used,!methods!for!brain crelated!analysis,!and!brain!related!findings!reported.!
We!extracted!primary!outcome!results!(brain!activity/neural!processes)!from!all!articles.!
Other!relevant!findings!were!also!recorded.! !
!
Data&analysis&and&synthesis&
Information!extracted!from!the!selected!articles!is!synthesized!in!Table!1!and!
includes!motivational!manipulation!implemented,!motor!task!utilized,!brain!outcome!
measure!used,!time!period!analyzed!and!braincrelated!findings.!The!data!were!
!
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qualitatively!summarized!for!the!reported!brain!and!neural!processes!associated! with!
motivationcenhanced!motor!improvements.! !
Results:&
Flow&of&included&studies&
Figure!1!shows!the!flow!of!included!studies.! Based!on!initial!search!results!using!
the!key!terms,!we!identified!181!citations!and!screened!their!titles!and!abstracts!for!
inclusion!and!exclusion!criteria.!This!first!screen!identified!29!articles.!In!the!second!
stage!of!screening,!full!text!articles!we re!reviewed!using!the!same!inclusion!and!
exclusion!criteria!and!20!articles!were!excluded!because!they!did!not!include!a!motor!
task,!used!animal!models,!or!were!review!articles.!Finally,!9!studies!were!included!in!the!
systematic!review.!!
Study&characteristics&
The!details!of!the!study!characteristics!are!presented!in!Table!1.!The!nine!
selected!studies!were!published!between!2009!and!2017.!The!most!common!brain!
related!measure!was!fMRI,!except!for!two!studies!(one!structural!MRI!and!one!TMS!
study).!Five!out!of!the!nine!studies!used!a!within csubject!design.!Eight!studies!used!
monetary!reward!as!a!form!of!motivation.!Participants!in!almost!all!studies,!except!one,!
were!young!adults!between!the!ages!of!18 c39!and!included!men!and!women.!!
Description&and&summary&of&individual&studies&
Wachter!et!al.!used!monetary!gains!and!losses!to!investigate!the!differential!
brain!activations!of!reward!and!punishment!on!motor!performance!using!a!serial !
!
15!
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reaction!time!task!practiced!during!an!fMRI!session.!During!task!execution,!the!
rewarded!group!showed!specific!activity!in!the!dorsal!striatum,!in!the!nucleus!
accumbens,!amygdala,!and!prefrontal!cortex.!The!punished!group!showed!increased!
activation!during!the!punished!trials!in!the!insula!bilaterally!and!in!portions!of!prefrontal!
cortex.!The!authors!concluded!that!the!learning!effect!of!reward!is!mediated!through!the!
dorsal!striatum!and!the!performance!effect!of!punishment!through!the!insula! (Wachter!
et!al.,!2009).!
Schmidt!et!al.!used!monetary!reward!to!examine!the!effects!of!motivation!on!
motor!effort!using!a!hand!grip!force!task!executed!during!an!fMRI!session.!At!incentive!
presentation,!expected!reward!was!associated!with!activity!in!the!ventral!striatum,!
internal!capsule,!ventral!pallidum,!thalamus,!insula,!middle!temporal,!and!posterior!
cingulate!cortex.!Additionally,!incentive!effects!on!ventral!striatum!activity!could!predict!
subsequent!incentive!effects!on!behavioral!performance.!At!the!time!of!feedback!
display,!no!correlation!between!ventral!striatum!activity!and!monetary!reward!was!
found.!The!authors!propose d!that!activation!in!the!ventral!striatum!is!a!cause,!not!a!
result,!of!better!behavioral!performance !(Schmidt!et!al.,!2012) .!
Lutz!et!al.!used!a!visuomotor!force!production!task!to!examine!the!effects!of!
performance!feedback!with!or!without!monetary!reward!on!brain!activity!in!the!striatum,!
using!fMRI.!They!found!that!ventral!striatu m!activity!was!higher!in!response!to!feedback!
about!good!performance!compared!to!bad!performance!trials,!and!that!ventral!striatum!
activity!is!further!enhanced!when!good!performance!is!paired!with!monetary!reward.!
Moreover,!monetary!reward,!and!not!good!performance!alone,!was!associated!with!
higher!activity!in!the!dorsal!striatum.!These!findings!indicate!that!the!dorsal!striatum!is!
!
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particularly!sensitive!to!extrinsic!motivation,!as!indexed!by!monetary!reward,!whereas!
the!ventral!striatum!responds!to!intrins ic!motivation,!indexed!by!performance!
achievement! (Lutz!et!al.,!2012).!
Widmer!et!al.!used!an!arccpointing!motor!task!practiced!in!the!MRI !scanner!to!
investigate!if!knowledge!of!performance!after!good!trials,!previously!shown!to!enhance!
motor!skill!learning,!elicits!h igher!striatum!activity!as!compared!to!a!group!who!received!
feedback!after!a!random!selection!of!trials .!Additionally,!striatum!activity!was!assessed!
by!a!third!group! that!received!performance cdependent!monetary!(knowledge!of!results!
and!monetary!reward)!reward.!Using!an!ROI!analysis!of!the!striatum!during!the!
feedback!presentation!period,!the!authors!found!higher!ventral!striatum!activity!in!the!
group!that!received!knowledge!of!results!and!monetary!reward!compared!to!the!groups!
that!received!knowledge!of!p erformance!alone.!There!were!no!differences!found!
between!the!group!that!received!feedback!after!good!trials!and!the!group!that!received!
feedback!after!random!trials.!The!authors!conclude!that!conditions!that!are!rewarding!
and!elicit!dopamine!release!via!striatal!activation!augment!motor!skill!consolidation!and!
promote!motor!skill!learning!(Widmer!et!al.,!2016).!
Chib!et!al. (Chib!et!al.,!2014) !used!a!tracking!task!during!fMRI!scan s!to!assess!if!
potential!gain!or!loss!of!monetary!reward!differentially!influences!choking!effects,!the!
impairment!of!performance!due!to!large!incentives !for!good!performance.!They!
examined!ventral!striatum!activity!at!the!time!of!reward!incentive!cue!and!t ask!period.!!
At!the!time!of!incentive!presentation,!an!increase!in!activity!for!the!ventral!striatum,!
supplementary!motor!area,!parietal!cortex,!posterior!and!anterior!cingulate!cortex,!
occipital!cortex,!dorsolateral!prefrontal!cortex!and!insula!was! positively!correlated!with!
!
17!
!
the!magnitude!of!potential!gain!or!loss.!In!contrast,!at!the!time!of!the!motor!task,!
decreasing!activity!in!ventral!striatum,! supplementary!motor!area,!parietal!cortex,!
cingulate!cortex,!occipital!cortex,!dorsolateral!prefrontal!cortex ,!globus!pallidus,!!and!
primary!motor!cortex!was!positively!correlated!with!magnitude!of!potential!gain!or!loss.!
The!authors!conclude!the!ventral!striatum!is!encoding!the!degree!of!motivation !
(irrespective!of!gain!or!loss),!indexed!by!reward!incentive,!for!executing!an!associated!
action!(Chib!et!al.,!2014) .!!
Lee!et!al.!applied!monetary!reward!in!a!bimanual!tracking!task!using!fMRI!to!
assess!activity!in!the!motor!cortex,!supplementary!motor!area!and!caudate .!Specifically,!
they!examined !the!relationship!of!the!brain!activity!in!these!regions!with!the!collapse!of!
performance!with!high!reward!incentives!(choking).! During!reward!cue!presentation,!the!
authors!reported!significant!increases!in!activity!in!response!t o!increasing!reward!cue!
presentation!for!several!brain!regions,!including!motor!and!premotor!cortices,!striatum,!
parietal!cortex,!bilateral!dorsolateral!prefrontal!cortex,!anterior!cingulate!cortex,!medial!
orbitofrontal!cortex,!left!putamen,!and!a!posterior!portion!of!the!right!middle!temporal!
gyrus.!Moreover,!they!report! that! when!performance!is!impaired!due!to!high!reward!
incentives,!the!lateral!prefrontal!cortex!and!the!anterior!cingulate!cortex!are!overactive!
and!show!greater!functional!connectivity!wit h!the!motor!cortex,!implicating!the!role!of!
these!two!regions!in!impeding!performance! (Lee!and!Grafton,!2015).!
Belkhiria!et!al.!used!verbal!encouragement!during!a!hand!grip!force!task!and!
assessed!taskcrelated!activity!and!functional!connectivity!between! the!cerebellum!and!
the!following!regions:! primary!motor!cortex,!orbitofrontal!cortex,!dorsolateral!prefrontal!
cortex,!superior!temporal!gyrus,!red!nucleus!and!ventral!striatum.!The!authors!report!
!
18!
!
activations!specific!to!verbal!encouragement!in!the!right!an d!left!cerebellum,!right!
dorsolateral!prefrontal!cortex,!left!and!right!orbitofrontal!cortex ,!left!and!right !motor!
cortex,!superior!temporal!gryus!and!red!nucleus.!Results!from!this!study!further!
revealed!that!while!the!anterior!part!of!the!right!lobule!VI !of!Cerebellum!was!activated!
by!motor!execution,!its!posterior!part!was!specifically!activated!by!verbal!
encouragement.!Finally,!the!authors!suggest!that!functional!connectivity!between!the!
cerebellum!and!the!red!nucleus,!between!the!red!nucleus!and!prefro ntal!and!motor!
cortices,!as!well!as!between!the!prefrontal!cortex!and!ventral!striatum!imply!a!closed c
loop!that!drives!motor!performance!from!motivation!with!verbal!encouragement!
(Belkhiria!et!al.,!2017) .!
Dayan!et!al.!used!whole!brain!voxel cbased!morphometry ,!to!evaluate!if!the!extent!
to!which!monetary!reward!guides!subsequent!motor!behavior!on!a!trial!by!trial!basis!for!
a!procedural!skill,!using!a!sequential!explicit!visuomotor!task,!can!be!predicted!by!
structural!graycmatter!characteristics.!It!was!reported!that!individuals!wh o!were!more!
dependent!on!reward,!examined!by!the!degree!to!which!reward!or!performance!
feedback!at!any!trial!predicted!successful!performance!at!the! following!trial,!showed!
smaller!graycmatter!volume!in!the!right!and!left!superior!frontal!gyri,!a!region!implicated!
in!reward!anticipation!and!reward!based!decision!making.!Findings!from!this!study!
provides!evidence!that!anatomical!substrates!can!partly!ex plain!the!inter cindividual!
variability!for!responding!to!rewarding!outcomes!of!one’s!actions!(Dayan!et!al.,!2014).!
Wilkinson!et!al.!used!TMS!to!assess!if!monetary!reward!and!performance!
feedback!can!improve!motor!performance!on!a!serial!reaction!time!task!when!primary!
motor!cortex!is!disrupted!by!TMS.!Findings!from!this!study! showed!that!rewarding!
!
19!
!
feedback!was!able!to!reverse!motor!performance!impairments!due!to! primary!motor!
cortex!inhibition !(Wilkinson!et!al.,!2015).!
Discussion&
In!this!systematic!review,!we!identified!9!studies!that!examine!motivation!effects!
on!braincrelated!measures!associated!with!motor!skill!behavior.!Six!studies!showed!
striatum!activity!associated!with!reward crelated!motor!behavior.!Five!studies!reported!
frontal!cortex!activity!to!be!involved!with!motivation! and!action!processes.!Four!papers!
connected!the!motor!co rtex!to!rewarding!conditions!related!to!movement.!Three!studies!
reported!cingulate!cortex!and!the!insula!to!be!involved!with!reward!processing!related!to!
motor!behavior. !
The!neural!processes!suggested!by!these!studies!can!be! described!by!two!
general!lines!of!theory.!One!prevalent!idea!raised!by!studies!that!identify!striatum c
related!activity,!propose!the!role!of!dopaminergic!system!activation!(elicited!by!reward)!
in!facilitating!neural!processes!that!underlie!motor!performance!and!learning.!This!
theory!fits !in!with!animal!studies!that!demonstrate!that!dopamine!input!to!the!motor!
cortex!supports!cortical!plasticity,!which!is!important!for!skill!acquisition,!and!that!
midbrain!dopamine!projections!to!the!motor!cortex!are!essential!for!motor!learning!
(MolinacLuna!et!al.,!2009h!Hosp!et!al.,!2011).!!
A!second!notion!implicates!frontal!and!limbic!brain!regions!involved!with!driving!
motivational!effects!on!motor!actions.!This!idea!suggests!that!regions!such!as!the!
cingulate!cortex,!insula!and!frontal!cortex!play!an!execu tive!role!in!signaling!and!
integrating!reward!information!to!motor!control!circuits,!perhaps!by!exerting!influence!on!
!
20!
!
cognitive,!attentional!and!emotional!networks.!This!is!aligned!with!neuroimaging!
research!that!highlights!the!role!of!the!salience!network ,!which!comprises!the!insula!and!
dorsal!anterior!cingulate!cortex,!in!directing!attention!to!stimuli!that! are!relevant!to!the!
task!and!for!guiding!subsequent!behavior.!Behaviorally!salient!stimuli!can!encompass!a!
host!of!motivational!variables,!including!reward!incentives,!expected!outcomes!and!
feedback!relating!to!previous!actions!that!can!influence!motor!control!behavior!and!
motor!learning!processes!(Seeley!et!al.,!2007h!Ham!et!al.,!2013).!Taken!together,! the!
dopaminergic!system!and!salient!network!may!work!in!conjunction!with!one!another!to!
achieve!motivational!influences!on!behavior !(Menon,!2015).!!
Results!from!this!systematic!review!shows!that!while!there!are!common!nodes!of!
brain!regions!related!to!reward!in!the! context!of!motor!behavior,!variability!in!reported!
brain!regions!highlight! experimental!factors!that!may!be!relevant! in!how!motivation!
guides!the!movement.!C ognitive,!emotional!and!executive!function ing!mechanisms! may!
be!simultaneously!affected!by!motivation(Pessoa,!2009,!2010),!in!addition!to!or!in!
conjunction!with!guiding!the!motor!system.! !
Several!sources!of!variability!among!studies!may!contribute!to!differences!in!
braincrelated!findings!reported!in!this! systematic!review.!!The !variability!in!the!brain!
regions!that!are!engaged!in!motivated!motor!behavior!may!be!due!to!analysis!methods.!
Most!notably,!decisions!about!experimental!methods!used ,!such!whole!brain!or!ROI c
based!analysis,!how!the!ROI!is!defined!(landmarks,!atlases,!maps !etc.)!and!the!time!
period!evaluated!for!brain!activity!(during!task!movement,!at!reward!cue!presentation!or!
reward!feedback!presentation)!are!critical!in!consolidating!and!interpreting!the!reported!
findings.!!
!
21!
!
Moreover,!the!tasks!used!to!assess!motor!behav ior!differ!among!studies!in!the!
type!of!demand!required!from!the!task,!in!particular!on!the!cognitive cmotor!spectrum!of!
the!task!characteristics.!For!instance,!implicit!sequence!learning!and!explicit!procedural!
learning!have! been!shown!to!produce!different !behavioral!results!in!response!to!reward!
and!likely!may!engage!different!brain!networks!as!well!(Steel!et!al.,!2016).!
Another!important!consideration!is!t he!significance!of!assessing!participant!
motivation!to!check!for!the!effectiveness!of!the!manipulation!to!induce!the!intended!
motivation.!The!majorit y!of!studies!use!group!assignment!or!performance!improvement!
as!the!only! method !of!manipulation!check!and!h ave!not!used!validated!questionnaires!
or!other!direct!measures!of!motivation!to!further!confirm!their!assumption!for!
successfully! enhancing!motivation!in!participants.! !
!
Implications&and&future&directions&
&
Future!studies!can!further!investigate!the!discrete!and!shared!neural!
mechanisms!that!embody!motivated!motor!behavior.!The!results!of!this!review!generate!
a!hypothesis!that!can!be!tested!to!further!our!understanding!for!how!motivation!boosts!
motor!behavior .!Considering!the!behavioral!research!differentiating! the!effect!of!various!
aspects!of!motivation!(intrinsic!and!extrinsic ,!reward!and!punishment!etc.) !on!learning!
and!behavior! (Ryan!and!Deci,!2000),!these!forms!of!motivation!may!also!engage!
different!pieces!of!!common!neural!networks!in!the!context!of!motor!learning.!A!handful!
of!neuroimaging!studies!have!begun!to!address!this!distinction! (Lutz!et!al.,!2012h!
!
22!
!
Widmer!et!al.,!2016).!However,!validation!of!motivation!effects!from!implemented!
experimental!manipulations,!using!psychological!measures!of!motivation,!is!necessary!
to!better!evaluate!the!interpretation!of!findings!from!such!studies.! !
Further,!experimental!paradigms!that!harness!motor!tasks!and!motivational!
manipulations!that!mirror!real cword!circumstances!can!optimize!our!interpretation!and!
generalizability!of!findings!from!braincrelated!investigations.!For!instance,!motivation!
that!instills!a!heightened!expectation!about!performance!achievements!for!successfully!
performed!actions!can!reflect! situations!familiar!in!educational!and!classroom!settings,!
sports!training!and!parenting!scenarios.!Alternatively,!motivation!that!is!driven!by!
achieving!monetary!reward!may!reflect!economic!decision!making,!gambling!behavior!
and!workcrelated!incentives.!While!both!examples!leverage!motivation,!studies!using!
braincrelated!techniques!along!with!these!types!of!motivational!manipulations!in!the!
context!of!motor!skill!behavior!can!promote!insights!into!underlying!neural!networks!that!
drive!behavioral!gains.! !
Implications!from!this!line!of!research!can!be!translated!to!optimize!use!of!
motivation!to!boost!behavior!in!various!fields,!including!teaching!practices,!clinician!to!
patient!interactions!and!reward!value!systems!used!in!work!settings!and!marketing!
strategies.!In!particular,!empirical!evidence!for!neural!effects!of!motivation!on!behavior!
provides!support!for!integrating!motivational!strategies!within!different!learning!domains.!
Furthermore,!clinical!research!in!psychiatric! and!neurological!disorders!that! tha t!affect!
motivational!processing!can!benefit!from!a!deeper!understanding!of!the!relevant!brain!
correlates!for!impaired!cognitive!and!motor!behavior!due!to!disrupted!neural!networks.!
Finally,!this!understanding!can!fuel! future!research!to!improve !medical!treatments!and!
!
23!
!
rehabilitative!therapies!for!patients!affected!by!brain!damage!in!areas!associated!with!
motivationcaction!processing.! !
&
!
!
!
!
%
! &
!
24!
!
Figure&1.!PRISMA&outflow!of&systematic&review.!(preferred!reporting!items!for!
systematic!reviews!and!metacanalysis)!
&
&
&
& &
!
25!
!
Table&1.&Summarizing&methods&and&brain&results&for&impact&of&motivation&on&motor&behavior.!!
Author,&
year&
Study&Design& Analysis&Methods&
Brain@related&findings&
Motivational&
manipulation&
Motor&
Task&
Brain&
outcome&
measure&
Brain&analysis:&
Whole&brain&or&
ROI&analysis&etc.&
Time&period&
analyzed&
Wachter,!
2009!
Monetary!
reward!(gain)!or!
punishment!
(loss)!
Serial!
reaction!
time!task!
fMRI! Whole!brain!! Task!execution! The!rewarded!group!showed!activation!in!the!
dorsal!striatum,!in!the!nucleus!accumbens,!
and!amygdala,!as!well!as!in!prefrontal!cortex.!
The!punished!group!showed!increased!
activation!during!the!punished!trials!in!the!
insula!bilaterally!and!in!portions!of!prefrontal!
cortex.!!
!
Schmidt,!
2012!
Monetary!
reward!!
Hand!
grip!force!
task!
fMRI!
!
!
Psychophysiologic
al!interactions!
(PPI)!connectivity!
analysis!using!left!
ventral!striatum!as!
a!seed!!
!
!
Incentive!cue!
presentation,!effort!
exertion,!feedback!
presentation!
At!incentive!presentation,!expected!reward!
was!associated!with!activity!in!the!ventral!
striatum,!internal!capsule,!ventral!pallidum,!
thalamus,!insula,!middle!temporal!cortex,!and!
posterior!cingulate!cortex.!Additionally,!
incentive!effects!on!ventral!striatum!activity!
could!predict!subsequent!incentive!effects!on!
behavioral!performance.!At!the!time!of!
feedback!display,!no!correlation!between!
ventral!striatum!activity!and!monetary!reward!
was!found.!
!
Lutz,!2012! Performance!
feedback!trials!
with!or!without!
monetary!
reward!
!
Visuomot
or!force!
productio
n!task!!
fMRI! ROI!analysis!
!
Ventral!striatum!
(nucleus!
accumbens!and!
the!ventral!part!of!
the!caudate!and!
putamen)!&!dorsal!
striatum!(dorsal!
part!of!the!caudate!
and!the!putamen)!
Reward!cue!
presentation,!target!
presentation,!
feedback!
presentation!
!
*ROI%analysis%
reported%during%
feedback%
presentation%only%
Ventral!striatum!activity!was!higher!in!
response!to!feedback!about!good!
performance!compared!to!bad!performance!
trials,!and!that!ventral!striatum!activity!is!
further!enhanced!when!good!performance!is!
paired!with!monetary!reward.!Monetary!
reward!was!also!specifically!associated!with!
higher!activity!in!the!dorsal!striatum.!
!
26!
!
Widmer,!
2016!
Knowledge!of!
performance!
after!good!trials!
with!or!without!
monetary!
reward!
Arc!
pointing!
task!
!
Function
al!MRI!
ROI!analysisS!
ventral!and!dorsal!
striatum!
Feedback!
presentation!!
Higher!ventral!striatum!activity!reported!for!
knowledge!of!results!and!monetary!reward!
compared!to!a!group!that!received!knowledge!
of!performance!alone.!
Chib,!2014! Monetary!
reward!to!obtain!
potential!
monetary!gains!
(gain)!or!to!
avoid!losing!a!
monetary!
amount!(loss)!
Tracking!
task!
!
fMRI! Whole!brain! Reward!cue!
presentation!&!task!
execution!!
!
At!the!time!of!incentive!presentation,!
increasing!activity!in!ventral!striatum,!
supplementary!motor!area,!parietal!cortex,!
posterior!and!anterior!cingulate,!occipital!
cortex,!dorsolateral!prefrontal!cortex!and!
insula!positively!correlated!with!the!magnitude!
of!potential!gain/loss.!
!
At!the!time!of!the!motor!task,!decreasing!
activity!in!ventral!striatum,!occipital!cortex,!
parietal!cortex,!sup!motor!cortex,!globus!
pallidus,!cingulate!cortex,!dorsolateral!
prefrontal!cortex!and!primary!motor!cortex!
was!correlated!with!magnitude!of!incentive!
gain/loss.!!
!
!
Lee,!2015! Monetary!
reward!
Bimanual!
tracking!
task!!
.!
fMRI!! ROI!analysis!
!
ROI:!Motor!cortex,!
supplementary!
motor!area!and!
caudate!
Reward!cue!
presentation,!just!
prior!to!movement!
onset!
During!reward!cue!presentation,!there!were!
significant!increases!in!activity!in!response!to!
increasing!reward!cue!presentation,!for!motor!
and!premotor!cortices,!striatum,!and!parietal!
cortexS!bilateral!dorsolateral!prefrontal!cortex,!
anterior!cingulate!cortex,!medial!orbitofrontal!
cortex,!left!putamen,!and!a!posterior!portion!of!
the!right!middle!temporal!gyrus.!
!
!
27!
!
Belkhiria,!
2017!
Verbal!
encouragementS!
recorded!human!
voice!saying!
‘Go’!repeated!20!
times!during!
task!execution!
encourage!the!
subjects.’!
Hand!
grip!force!
task!!
fMRI! Whole!brain!
analysis!and!
psychophysiologic
al!interactions!
(PPI)!connectivity!
analysis!!
!
ROI:!Cerebellum,!
primary!motor!
cortex,!
orbitofrontal!
cortex,!
dorsolateral!
prefrontal!cortex,!
superior!temporal!
gyrus,!red!nucleus!
and!ventral!
striatum!
Task!execution! Activations!specific!to!verbal!encouragement!
reported!in!the!right!and!left!cerebellum,!right!
dorsolateral!prefrontal!cortex,!left!and!right!
orbitofrontal!cortex!and!right!and!left!motor!
cortex,!superior!temporal!gryus!and!red!
nucleus.!Anterior!part!of!the!right!lobule!VI!of!
cerebellum!was!activated!by!motor!execution,!
while!its!posterior!part!was!specifically!
activated!by!verbal!encouragement.!
Dayan,!
2014!
Performance!
feedback!with!or!
without!
monetary!
reward!!
Sequenti
al!explicit!
visuomot
or!task!
Structural!
MRI!
Whole!brain!voxel`
based!
morphometry!
NA! Individuals!who!were!more!dependent!on!
reward!(trial`by`trial!relationship!between!
reward!and!subsequent!performance)!had!
smaller!gray`matter!volume!in!right!and!left!
superior!frontal!gyri.!
!
Wilkinson,!
2015!
Monetary!
reward!and!
performance!
feedback!
(smiley!for!
successful!trials,!
sad!for!
unsuccessful!
trials)!
Serial!
reaction!
time!task!
Continuo
us!theta!
burst!
(cTBS)!
transcran
ial!
magnetic!
stimulatio
n!(TMS) !
NA! NA! Rewarding!feedback!reversed!motor!
performance!impairments!due!to!primary!
motor!cortex!inhibitions!with!TMS. !
!
!
28!
!
Chapter(3:(!Intrinsic(motivation(promotes(motor(skill(learning(through( strengthening(of(
cingulate(cortex(activity(during(feedback(processing(after(good(performance(trials (
(
Beroukhim(Kay,(D.,
1,3
(Monterosso,(J.,
1,2
(Lewthwaite,(R.,
3
(Kutch,(J.,
1,3
(Winstein(C.
1,3(
(
1
(Neuroscience(Graduate(Program,(Southern(California(Clinical(&(Translational(Science(
Institute,(
2(
Department(of(Psychology,(
3
Division(of(Biokinesiology(and(Physical(Therapy,(
The(Ostrow(School(of(Dentistry,(Un iversity(of(Southern(California(
!
Corresponding!author!
Carolee(J.(Winstein(
1540(Alcazar(St,(Los(Angeles,(CA(90033 (
winstein@usc.edu(
(323)(442Z 2903((
(
Number(of(pages:(43(
Number(of(figures:(6(
Number(of(tables:(2(
Abbreviated(title:!Neural(correlates(of(motivation(benefits(on(skill(learning (
!
29!
!
Abstract!
Intrinsic(motivation(can(energize(behavior(and(has(also(been(shown(to(enhance(
motor(skill(performance(and(learning.(To(investigate(the(neural(mechanisms(that(
mediate!the(influence(of(motivation(on(motor(learning,(we(used(functional(magnetic(
resonance(imaging((fMRI)(to(scan(healthy(partici pants(during(practice(of(a(challenging(
spatiotemporal(tracking(task,(under(either(a(neutral(condition((control(group)(or(an(
enhanced(motivation(condition((motivation(group).(Motor(learning,(expressed(as(spatial(
and(temporal(accuracy(for(learning(the(tracki ng(pattern,(was(assessed(immediately(after(
practice(and(one(day(later( using(performance(on(retention(tests.(Motor(learning(was(
compared(between(groups(using(an(independent(samples(t Z test.(Brain(activity(during(
practice,(which(correlated(with(the(motor(task(execution(period(and(nonZ movement(
performance(feedback(interval(as(modulated(by(corresponding(trial(performance,(was(
compared(between(groups(using(an(independent(samples(t Z test.((
The(motivation(group(showed(better(motor(skill(learning,(as(indexed(by(highe r(
temporal(accuracy(performance,(but(had(similar(spatial(accuracy(performance(during(
immediate(and(delayed(retention(tests,(as(compared(to(the(control(group.(Activation(in(
the(cingulate(cortex,(distinguished(the(groups(during(the(performance(feedback(inter val,(
with(the(motivation(group(showing(stronger(association(of(activity(with(performance(
trials(of(higher(temporal(accuracy( compared(to(the(control(group.(The(cingulate(cortex(is(
a(key(region(in(the(salience(network(and(implicated(in(performance(monitoring.(
Our(results(demonstrate(that(enhanced(intrinsic(motivation(improves(motor(
learning(and(modulates(brain(activity(associated(with(the(processing(of(feedback(
!
30!
!
information.(This(suggests(that(motivation( enhanced(sensitivity(to(performance(
feedback(after(good(trials(during(skill(acquisition(for(the(group(that(practiced(under(
motivated(conditions,(compared(to(that(for(the(control(group. (
(
Introduction!
! Motivation(can(have(a(powerful(influence(on(learning(and(behavior.(Evidence(
from(several(lines(of(research(have(demonstrated(its(effect(in(various(learning(
paradigms(and(environments,(including(education,(decision Z making(and(movement(
science((Dweck,(1986_(Wood(and(Bandura,(1989_(Ryan(and(Deci,(2000_(Wulf(and(
Lewthwaite,(2016).(Motivation(can(be(classified( in(several(ways,(including,(extrinsic(and(
intrinsic(forms.(Extrinsic(motivation(refers(to(being(driven(to(do(something(because(it(
leads(to(an(independent(outcome,(such(as (working(for(monetary(incentives(or(studying(
for(a(good(grade.(Intrinsic(motivation,(on(t he(other(hand(refers(to(being(moved(to(doing(
something(because(it(is(inherently(interesting(or(valuable(and (perhaps(satisfies(
psychological(needs(for(competence,(autonomy(and(social(relatedness( (Ryan(and(Deci,(
2000)(
( There(is(considerable(evidence(that(motivation((i.e.(intrinsic(and(extrinsic)(can(
benefit(motor(learning.(Behavioral(studies(have(shown(the(influence(of(motivation(on(
learning(and(performance(of(motor(skills(using(various(manipulations(to(induce(
motivation,(including(monetary(reward((Abe(et(al.,(2011_(Galea(et(al.,(2015) ,(social(
comparison((Lewthwaite(and(Wulf,(2010),(social(reward((Sugawara(et(al.,(2012)((
performance(feedback((Chiviacowsky(and(Wulf,(2007)(and(instructions(that(enhance(
!
31!
!
expectancies(and(support(competence(( (Wulf(and(Lewthwaite,(2009_(Wulf(et(al.,(2012,(
2014)(
( Findings(from(human(neuroimaging(studies(have(revealed(several(brain(areas(
that(may(be(working(together(to(explain(the(positive(behaviora l(effects.(This(body(of(
work(has(highlighted(the(corticoZ striatal(circuit(including(the(ventral(tegmental(area,(the(
striatum(and(prefrontal(cortex(as(crucial(for(reward Z related(motor(behavior((Dayan(and(
Balleine,(2002_(Wickens(et(al.,(2003) .(Functional(magnetic(resonance(imaging((fMRI)(
studies(have(primarily(investigated(the(relationship(of(motivation(and(motor(behavior(
using(monetary(reward((Pessiglione(et(al.,(2007_(Wrase(et(al.,(2007_(Schmidt(et(al.,(
2009,(2012_(Wachter(et(al.,(2009),(and(some(forms(of(intrinsic(motivation( (Lutz(et(al.,(
2012_(Widmer(et(al.,(2016).((For(example,(a(study(by(Widmer( and(colleagues(found(that(
providing(monetary(reward(after(good(performance(trials(during(task(practice(leads(to(
higher(ventral(striatal(activation(and(is(associated(with(better(consolidation(of(the(motor(
skill,(compared(to(a(group(that( receives(performance(feedback(alone.(The(authors(
propose(that(rewarding(conditions(that(increase(dopaminergic(signals(during(task(
practice(facilitate(successful(motor(skill(learning( (Widmer(et(al.,(2016).((
( Evidence(from(animal(studies(have(further(supported(the(role(of(the(
dopaminergic(system(in(explaining(the(benefits(of(extrinsic(reward(on(motor(skill(
acquisition((MolinaZ Luna(et(al.,(2009_(Hosp(et(al.,(2011_(Hosp(and(Luft,(2013) .(In(an(
experiment(with(rodents(training(on(a(forelimb(reachin g(skill(to(retrieve(a(food(pellet(
(reward),(dopaminergic(projections(from(the(ventral(tegmental(area((VTA)(of(the(
midbrain(to(the(motor(cortex((M1)(were(shown(to(be(necessary(for(successful(motor(skill(
learning.(Previous(research(has(described(the(role(of( dopaminergic(VTA(neurons(in(
!
32!
!
encoding(reward((Schultz,(2007).(Taken(together,(dopaminergic(input(to( M1(may(be(
providing(motivationZ related(information(that(supports(cortical(synaptic(plasticity,(a(
mechanism(implicated(in(motor(skill(learning( (RioultZ Pedotti(et(al.,(2000),(leading(to(
effective(learning(processes(for(motor(skill(acquisition( (Hosp(and(Luft,(2013).(Whether(
intrinsic(motivation(influences(brain(processes(related(to(human(motor(skill(learning(in(
the(same(way(as(previously(reported(aspects(of(motivation((i .e.,(extrinsic)(remains(to(be(
revealed.(
( The(objective(of(this(study(is(to( investigate(the(neurobiological(basis(for(the(
impact(of(intrinsic(motivation,(induced(by(enhanced(expectancies(of(performance(
success(and(support(of(competence,(on(motor(sk ill(learning.(We(used(a(between Z
subject(design(to(examine(effects(of(enhanced(intrins ic(motivation(on(brain(activity,(
using(fMRI,(during(concurrent(motor(performance(on(a( spatiotemporal(tracking(task.(We(
assessed(effects(of(our(motivational(manipulation(on(motor(learning,(evaluated(by(
spatial(and(temporal(accuracy(performance(on(the(same(d ay(and(next(day(retention( and(
transfer(tests,(and(intrinsic(motivation,(using(validated(measures,(including(selfZ efficacy(
and(intrinsic(motivation(inventory(subscales(for(interest/enjoyment,(perceived(
competence(and(effort/importance.(The(overall(hypothesis(was(that(intrinsic(motivation(
would(engage(the(striatum(during(task(execution,(similar(to(extrinsic(motivation ,(and(
enhance(motor(cortex(activity(leading(to(improved(motor(performance(and(learning.(( (
Methods!
Participants.!43(participants(were(recruited(into(the(study.(Three(individuals(were(
excluded(from(analysis(because(of(incomplete(behavioral(and(fMRI(data(due(to(
!
33!
!
scheduling(conflicts(for(the(neuroimaging(scan.(40(subjects((25(female,(mean(age(
(SD)=23.83((4.63)(years)(participated((Table(1).(Al l(participants(were(rightZ handed,(able(
to(participate(in(an(MRI(and(had(normal(or(corrected(to(normal(vision.(Hand(preference(
was(assessed(using(the(Edinburgh(Handedness(Inventory((Oldfield,(1971)(confirming(
that(all(participants(were(rightZ hand(dominant.(Participants(were(recruited(from(the(
university(community(and(had(no(prior(experience(with(the(motor(task.(All(participants(
gave(informed(consent(and(the(study(was(performed(with( USC(Institutional(Review(
Board(approval.((The(sample(size(for(the(behavioral(analysis(was(based(on(a(previous(
study(that(used(a(similar(motivational(manipulation( (Wulf(et(al.,(2012).(To(detect(
differences(in(motor(learning,(we(used(an(effect(size(of(0.18,(an(alpha(of(0.05(and(
power(of(0.8.(G*power(calculations(resulted(in(a(total(sample(size(of(20(per( group.(A(
formal(power(analysis(was(not(implemented(a (priori(to(identify(the(sample(size(for(the(
fMRI(analysis.(The(sample(size(for(the(fMRI(analysis(was(estimated(to(be(similar(to(
other(studies(in(this(field.(
(
Experimental!design.!We(used(a(betweenZ subject(design(to(examine(effects(of(
enhanced(intrinsic(motivation(on(brain(activity(and(concurrent(motor(performance .!!
Participants(lay(supine(in(the(MRI(and(held(a(pressure(sensor((TSD110 Z MRI,(BioPac)(
between(their(thumb(and(index(finger.(The(pressure(sensor( was(connected(to(an(air(
tube,(which(was(led(to(the(MRI(control(room,(where(it(was(connected(to(a(transducer(
converting(air(pressure(to(voltage.(Pinching(of(the(pad(result ed(in(generation(of(a(
differential(voltage(signal.(The(recorded(signal(was(used(to(provide(participants(with(
!
34!
!
visual(feedback(of(the(cursor(movement.(The(cursor(movement(and(moving(target(
presentation(were(displayed(on(a(projector(screen,(which(participants(were(able(to(see(
with(a(mirror(positioned(above(their(eyes.( (
The(study(consisted(of(four(phases((Familiarization ,(Baseline,(
Training/Immediate(Retention(and(Delayed(Retention/Transfer),(as(illustrated(in(Figure(
1.(During(the(fMRI(scan,(participants(practiced(a(tracking(tas k(over(6(blocks(of(12(trials(
each,(with(performance(feedback(presented(after(every(trial.(The(first(block((Baseline)(
for(both(groups(was(practiced(under(a(neutral(condition(to(assess (baseline(
performance,(followed(by(5(practice(blocks((Training)(with(neutr al(statements((control(
group)(or(neutral(and(motivational(statements((motivation(group).(Prior(to(the(fMRI(
session,(participants(were(familiarized(with(the(task(inside(a(mock(scanner(
(Familiarization).(Participants(were(assessed(with(a(test(block(immediate ly(after(the(
training(session((Immediate(retention)(in(the(MRI(and(1(day(later((Delayed(retention(and(
Transfer(session)(inside(the(mock(scanner.(All(test(blocks((immediate/delayed(retention(
and(transfer)(included(5(trials(without(performance(feedback.(Thre e(different(
spatiotemporal(tracking(patterns(of(equivalent(difficulty(were(used(to(optimize(
comparability(of(tracking(pattern(and(non Z pattern(specific(learning(for(the(phases(
(familiarization(period:(pattern(A,(baseline(period:(pattern(B,(and(training(period:(pattern(
C),(respectfully.(All(tracking(patterns(had(equivalent(amplitude(range(and(number(of(
reversals.!
!
!
35!
!
Motivational!manipulation.(After(the(baseline(block(in(the(scanner,(the(motivation(
group(received(the(following(statements,(to(induce(enhanced(intr insic(motivation,(prior(
to(each(training(block(followed(by(‘Well(done’(at(the(end(of(the(training(period.(
Motivation(statements(were(modified(from(previous(behavioral(studies(and(were(
designed(around(three(criteria:(to(enhance(self Z efficacy(expectation,(induce(a(positive(
(growth(or(incremental)(concept(of(ability(mindset((abilities(can(be(developed(through(
effort(and(persistence)((Dweck,(1986)(and(support(perceived(competence,(using(
positive(comments/praise(about(the(participant’s(performance.(Statements(included(the(
following(prior(to(each(training(block !1)(Keep!in!mind!that!at!the!beginning!it!is!common!
to!undershoot!or!overshoot!the!target,!but!this!is!the!type!of!task!that!you!get!better!at!
with!practice.!2)!Alright!!Your!improvement!across!the!past!trials!is!reflecting!your!
learning!and!getting!the!hang!of!it.!3)!Very!good!!Based!on!your!performance!so!far,!it!
looks!like!you!are!going!to!continue!to!improve!across!the!coming!trials.!4)!Alright!!Nice!
work!!5)!Great!job!!
!
Manipulation!check.!(SelfZ efficacy(was(assessed(before(and(after(the(training(phase.(
At(the(end(of(the(study,(participants(completed( the(Intrinsic(Motivation(Inventory(
(adapted(to(the(study(task)(and(rated(their(interest/enjoyment,(perceived(competence(
and(the(effort/importance(they(attributed(to(the(task.( (
!
Self ?efficacy!questionnaire.(A(6Z item(questionnaire(assessed(taskZ specific(selfZ
efficacy(on(a(Likert(scale.(All(participants(were(asked(to(rate(how(confident(they(were(to(
!
36!
!
keep(the(red(box((cursor)(in(the(white(target(box(for(2,(4,(6,(8,(10(and(12(seconds.(The(
scale(ranged(from(0((not(confident(at(all)(to(10((extremely(confident).(The(self Z efficacy(
questionnaire(was(given(after(the(familiarization,(before(and(after(the(training(session ,(
and(before(delayed(retention(trials(on(day(2.(See(appendix(for(questionnaire(items. (
( Cronbach’s(alpha(was(used(as(a(measure(of(internal(consistency(for(the(six(
items(of(the(selfZ efficacy(scale.(The(Cronbach(alpha(for(the(four(time(points :(
familiarization((0.799),(before(training((0.759),(after(training((0.778)(and(before(delayed(
retention((0.713),(indicated(an(acceptable(level(of(internal(consistency.(
Intrinsic!Motivation!Inventory.(An(adapted(version(of(the(perceived(competence,(
effort/importance,(and(task(interest/enjoyment(subscales(of(the(Intrinsic(Motivation(
Inventory(was(administered(after(delayed(retention(trials(on(day(2.(Items(on(the(
questionnaire(were(shuffled(in(order(of(the(items(presented (for(participants.(See(
appendix(for(questionnaire(items.(
!
Control!condition.!Neutral(statements((‘Just(checking(in,(we(will(go(ahead(and(
continue(with(the(task(trials(now.’)(were(provided(to(all(participants(between(tr aining(
runs(to(ensure(that(social(contact(and(communication(between(the(groups(was(
comparable.(
(
Behavioral!task.(The(goal(of(the(tracking(task(was(to(keep(the(cursor(inside(a(target(
box,(which(was(moving(vertically(across ((up(and(down)(the(screen,(for(as(long(as(
!
37!
!
possible(in(a(12Z second(trial(period.(Participants(controlled(the(cursor(movement(by(
squeezing(an(instrumented(pneumatic(pad,(between(the(thumb(and(index(finger.(
Performance((time(on(target)(feedback(was(presented(for(1(second(after(each(trial(
followed(by(a(12Z second(rest(before(the(next(trial(began.( (
(
Task!performance!measures!
Task(performance(was(assessed(during(practice(and(retention(in(2(ways.( Time(
on(target(was(used(as(one(measure,(which(was(part(of(the(explicit(task(goal,(provided(
as(performance(feedback(during(practice(and(used(for(the(self Z efficacy(questionnaire.(
For(a(second(measure,(we(decomposed(the(participant’s(cursor(trajectory(into(
spatial(and(temporal(error(components(to(determine(the(effect(of(motivational(
statements(on(these(two(aspects(of(the(motor(skill(being(learned,(to(modulate(force(
according(to(a(spatioZ temporal(pattern.(In(a(timeZ series(analysis,(the(participant’s(cursor(
trajectory(was(iteratively(shifted(in(time(toward(the(target(onset(until (a(maximum(
correlation(between(participant(trajectory(and(target(was(achieved(for(each(12(second(
trial.(Correlation(coefficients(represent(the(spatial(accuracy(of(tracking(performance.(
The(number(of(samples,(transformed(to(milliseconds(that(the(participant’s(trajectory(was(
shifted(to(achieve(the(maximum(spatial(accuracy(reflects(the(time(lag(or(temporal(
error/accuracy(of(tracking.((
Time!on!target.(The(time(that(the(cursor(was(entirely(inside(the(target((the(
minimum(of(the(cursor(was(greater(the(minimum(of(the(target(and(the(maximum(
!
38!
!
of(the(cursor(was(less(than(the(maximum(of(the(target)(was(calculated(for(each(
trial.(Average(time(on(target(was(calculated(separately(for(each(block.( (
Spatial!accuracy:! Maximum!correlation!coefficient.!The(maximum(correlation(
coefficient(between(the(target(trajectory(and(the(participant’s(cursor(trajectory,(
after(adjusting(for(time(lag,(was(calculated(for( each(trial.(Average(correlation(was(
calculated(separately(for(each(block.(
(
Temporal!accuracy:!Time!lag!of!tracking.!The(shift(in(time(that(was(associated(
with(the(maximum(correlation(between(the(target(trajectory(and( participant’s(
cursor(trajectory(was(calculated(for(each(trial.(Average(time(lag(was(calculated(
separately(for(each(block.(
!
Behavioral!data!analysis.((
( To(assess(performance,(time(on(target,(maximum(correlation(between(the(
participant’s(cursor(trajectory(and(target(trajectory (and(the(time(lag(that(was(used(to(
achieve(the(maximum(correlation(was(evaluated(for(each(practice(block.(Data(were(then(
corrected(for(outlier(trials((time(lag>(average(time(lag(of(the(corresponding(block(of(trials(
+/Z (2(standard(deviations).((
!
Image!acquisition.!We(acquired(T2Z weighted(echo(spiral(images(with(blood(oxygen(
level(dependent((BOLD)(contrast(using(a( 3(Tesla(scanner((GE(Signa(Excite)(with(an(8 Z
!
39!
!
channel(head(coil.(Each(volume(comprised( 37,(3mm(thick(axial(slices,(using(the(
following(parameters:(TR(=(2.5(seconds,(TE(=(34.5(flip(angle(=(90_ (FOV(=(22,(64(x(64(
matrix.(For(each(participant,(a(total(number(of(131 (volumes(were(obtained.((Additionally,(
we(acquired(a(T1Z weighted(structural(image(at(the(start(of(the(scan(session,(before(
performing(the(tracking(task.((
(
fMRI!analysis.(Analysis(of(fMRI(data(was(conducted(using(FSL ((FMRIB(Software(
Library).(The(fMRI(data(for(each(participant(was(preprocessed(using(the(FMRIB(
(Functional(Magnetic(Resonance(Imaging(of(the(Brain)(Expert(Analysis(Tool((FEAT,(
http://fsl.fmrib.ox.ac.uk/fsl/fslwiki/).(Preprocessing(included(skull(extraction(with(the(FSL(
brain(extraction(tool((BET),(motion(correction(and(spatial(smoothing(using(a(Gaussian(
kernel(with(fullZ width(halfZ maximum(of(5(mm.(For(first(level(data(analysis(of(each(
participant’s(individual(runs((block),(we(designed(a(general(linear(model(that(included(
two(regressors.(A(regressor(for(onset(and(duration(of(each(task(period(was(defined(for(
each(trial,(corresponding(to(task(planning(and(execution.(The(performance(feedback(
presentation(was(modeled(as(a(separate(regressor(with(onset(and(duration(
corresponding(to(the(1(sec(feedback(period(following(each(task(trial,(with(parametric(
modulation(of(the(feedback(regressor(by(temporal(accuracy(for(each(corresponding(trial.(
HigherZ level(analysis(was(used(to(combine(the(5(training(blocks(using(a(one(sample(t Z
test(for(each(participant.(A(one Z sample(tZ test(was(used(to(assess(task Z related(activity(
across(both(groups.(An(independent(samples(tZ test(was(used(to(compare(brain(activity(
1)(related(to(task(or(2)(related(to(fee dback(processing(between(groups((contrast(1:(
associated(with(long(time(lag_(contrast(2:(associated(with(short(time(lag) .(The(groupZ
!
40!
!
level(images(were(thresholded(with(clusterZ based(correction(for(multiple(comparisons(
with(Z>2.3(and(p<0.05.((
Five(participants(were(excluded(from(the(fMRI(analysis((motivation(group:(1(
missing(data,(1(congenital(neurological(condition_(control(group:(3(missing(data),(leaving(
n=18(in(the(motivation(group(and(n=17(in(the(control(group(represented(in(the(reported(
fMRI(results.(
(
Statistical!analysis!of!behavioral!and!questionnaire!data.!! All(statistical(tests(were(
conducted(using(SPSS.!
Task(performance((time(on(target,(spatial,(and(temporal(accuracy)(was(analyzed(
in(a(2((groups:(motivation(and(control)(X(5((training(block)(analysis(of(variance((ANOVA)(
with(repeated(measures(on(the(training(blocks(factor,(controlling(for(baseline(
performance.(To(assess(learning,(tracking(pattern(C(performance(during(immediate(and(
delayed(retention(trials(was(used.(To(assess(generalizability(of(skill(learned,(
performance(during(transfer(trials(for(pattern(A(and(B(were(used.(An(independent(t Z test(
was(used(to(compare(measures(between(groups(f or(retention(and(transfer(trials.(
(SelfZ efficacy(was(analyzed(in(a(2((groups:(motivation(and(control)(X(2((time(
point:(post(training(and(preZ retention)(repeated(measures(ANOVA(to(c ompare(ratings(
between(groups.(To(examine(the(relationship( between(1)(selfZ efficacy(changes(and(task(
performance(gains((start(to(end(of(training),(and(2)(self Z efficacy(changes(and(measures(
of(motor(learning,(assessed(at(immediate(and(delayed(retention,(Spearman(correlations(
!
41!
!
were(examined.(The(difference(in(correlation(coefficients (between(groups(was(assessed(
for(statistical(significance(by(applying(a(Fisher(r Z toZ z(transformation.((
(An(independent(tZ test(was(used(to(compare(ratings(between(groups(for(each(
subscale(of(the(intrinsic(motivation(inventory.(The(relationship(between(intri nsic(
motivation(reports((for(the(3(subscales(of(intrinsic(motivation(inventory:(
interest/enjoyment,(perceived(competence(and(effort/importance)(and (measures(of(
motor(learning,(performance(at(immediate(and(delayed(retention,(was(examined(using(
Spearman(correlations.!!
Results!
Behavioral!results!!
!
! Motivation!measure!ratings!in!the!two!groups!
SelfIefficacy!ratings!
( Figure(2A(shows(that(selfZ efficacy(increased(across(the(four(time(points(of(
the(study(for(both(groups,(f(1.863,61.464)=44.062,(p<0.001.(At(post( training(and(
pre(retention(time(points,(the(motivation(group(show ed(a(trend(for(higher(reported(
selfZ efficacy(compared(to(the(control(group,(however(when(controlling(for(pre(
training(selfZ efficacy(ratings,(the(main(effect(of(group(was(not(significant,(
f(1,32)=1.919,(p=0.176.(
(
!
42!
!
Intrinsic!motivation!ratings!
( As(figure(2B(shows,(interest/enjoyment(ratings(and(perceived(competence(
ratings(were(comparable(between(groups,(t(35)=0.995(p=0.327(and(t(35)=0.952(
p=.348,(respectively.(Effort/important(ratings(were( marginally(higher(in(the(
motivation(group(compared(to(the(control(group,(approaching (conventional(
significance(levels,(t(35)=1.791(p=.082.((
(
Task!performance:!Motivation!improves!motor!skill!learning!
Motor!performance!!
( Figure(3(shows(both(groups(demonstrate(si milar(time(on(target,(spatial(
accuracy((cross(correlation)(and(temporal(accuracy((time(lag)(during(the(baseline(
block(and(all(participants(improved(across(training(blocks(on(day(1.(The(main(
effect(of(training(block(approached(significance(for(time(on(target(performance,( f(
(2.045)=3.005,((p=0.055(and(tracking(cross(correlation,( f((2.497)=2.342,(p=0.090.(
There(was(no(main(effect(of(training(block(for(time(lag( performance,(f(
(2.155)=1.872,(p=0.158.(
( There(was(no(statistically(reliable(effect(of(group(during(training(for(time(on(
target(performance,(f((1,(34)=0.532,(p=0.471(or(tracking(accuracy,(f((1,35)=0.156(
p=0.695.(There(was(no(Group(x(Block(interaction(for(time(on(target(or(tracking(
accuracy(performance.((
!
43!
!
( Notably,(there(was(a(reduction(in(temporal(error((time(lag(of(tracking)(for(
both(groups,(with(the(motivation(group(showing(a(significantly(greater(temporal(
error(reduction.(This(group(difference(was(reliable(as(revealed(by(a(statistically(
significant(group(main(effect(of(time(lag(performance(during(training,(
f(1,34)=5.017,(p=.032.(There(was(no(Group(x(Block(interaction (for(time(lag(
performance,(f((2.155)=2.273,(p=0.106.(
(
Motor!learning(!
( Figure(3(shows(that(the(motivation(group(presents(higher(time(on(target(
performance(at(immediate(retention,(though(this(was(not( statistically(significant,(
t(31.506)=1.9,(p=.067,(and(neither(was(delayed(retention,(F(28.27)=1.577,(
p=0.126.(The(groups(show(comparable(spatial(tracking(accuracy(for(immediate(
and(delayed(retention,(t(37)=0.216,((p=0.830(and(t(35)=( Z 1.042,(p=0.313,(
respectively.(In(sharp(contrast(to(time(on(target(and(spatial(accuracy ,(the(
motivation(group(had(significantly(higher(temporal(accuracy(at(immediate(
retention,(t(Z 3.452)=37,(p=0.001,(and(delayed(retention,(t(29.737)=(Z 2.803,(
p=0.009(than(did(the(control(group.( (
!
Transfer!effects!
( On(day(2(transfer(tests,(the(motivation(group(showed(higher(time(on(target(
performance(compared(to(the(control(group(for(transfer(1(( pattern(B)(
!
44!
!
(t(35)=2.169,((p=.037)(but(not(transfer(2((pattern(A)(t(35)=1.016,((p=0.316.(
Spatial(accuracy(was(comparable(for(both(groups(for(both(t ransfer(tests,(t(35)=Z
0.493,(p=0.625(and((t(35)=Z 0.605,(p=0.549,(transfer(1(and(2,(respectively. (In(
contrast,(the(motivation(group(showed(significantly( greater(temporal(accuracy(
compared(to(the(control(group(for(transfer(1( but(not(for(transfer(2,(t(29.38)=Z
2.051,(p=.049(and(t(35)=Z 1.903,(p=.065,(respectively.(((
!
Relationship!of!motivation!and!task!performance:!Motivation!strengthens!
the!link!between!self?efficacy!and!motor!skill!learning!
( Across(the(groups,(there(was(a(significant(correlation(between(change(in(
selfZ efficacy((pre(to(post(training)(and(gain(in(time(on(target(performance(
(baseline(to(last(training(block),(r=0.510,(p=0.022(and(this(relationship(was(
similar(between(groups((M:(r=.594,(p=0.006,(C:(r=0.637,(p=0.004).((
( More(importantly,(and(shown(in( Figure(4,(there(was(a(statistically(
significant(correlation(between(change(in(self Z efficacy(preZ toZ post(training(and(
time(on(target(performance(at(immediate(and(delayed(retention,( for(the(
motivation(group,(but(not(for(the(control(group((Immediate(retention,(M:(R=0.492,(
p=0.028,(C:(R=0.124,(p=0.602_(Delayed(retention(M:(R=0.535,(p=0.022,(C:(
R=0.019(p=0.939).(The(correlation(coefficient(between(selfZ efficacy(change(and(
delayed(retention(was(higher(for(the(motivation(group(compared(to(the(control(
group,(approaching(statistical(significance,(p=0.0594.!
!
45!
!
( (Moreover,(there(was(a(statistically(significant(correlation(between(self Z
efficacy(reported(post(training(on(day(1(and(time(on(target(performance(at(
delayed(retention(for(the(motivation(group(but(not(for(the(control(group((M:(
R=0.634,(p=.005_(C:(R=0.415,(p=.087).( !
( There(was(no(statistical(significant(correlation(between(time(on(target(
performance(at(immediate(retention(and(reported(effort(( R=0.405,(p=0.095),(
interest((R=0.437,(p=0.070)(for(either(group((R=0.137,(p=0.577_(R=0.173,(
p=0.480).(The(two(groups(had(a(similar(correlat ion(between(immediate(retention(
performance(and(reported(competence((M:(R=0.505,(p=.033_(C:(R=0.444,(
p=.057).(There(was(no(significant(correlation(between(IMI(subscales(and(delayed(
retention(performance(for(either(group.( (
!
Neuroimaging!results:!Motivation!strengthens!brain!activity!in!the!cingulate!
cortex,!during!feedback!interval,!associated!with!short!time!lag!performance!!
( To(determine(task(related(brain(activity,(we(combined(both(groups(after(
combining(the(5(training(runs.(As(demonstrated(in(figure(5,(n eural(correlates(of(motor(
task(execution(were(principally(observed(in(the(premotor,(motor(and(supplementary(
motor(cortex,(putamen,(cingulate(cortex,(parietal(cortex,(cerebellum(and(occipital(cortex,(
consistent(with(previously(reported(visuoZ motor(task(related(activity((Vaillancourt(et(al.,(
2003_(FloyerZ Lea(and(Matthews,(2004_(Hatakenaka(et(al.,(2007) .(All(reported(significant(
activations((Table(2)(have(a(minimum(of(100(voxels,(with(a(threshold(of(p<(0.05.( (
( To(identify(brain(activity(that(mediates(the(effect(of(enhanced(motivation(on(motor(
!
46!
!
performance(processes,(we(examined(where(the(two(groups(differed(during(task(
execution.(We(used(the(mask(of(the('taskZ related(activity'(from(our(combined(group(
analysis(and(specific(relevant(regions(of(interest,(including(M1,(premotor(cortex,(
putamen,(cingulate(cortex(and(insula,(and(found(no(group (differences(during(the(task(
period.((
( To(assess(regions(that(were(as sociated(with(measures(of(motivation,(an(
independent(samples(tZ test(was(used(to(compare(task Z related(activity(between(groups,(
with(changes(in(selfZ efficacy(as(a(covariate_(again(there(were(no(group(differences.(
( To(examine(if(the(groups(differed(in(brain(a ctivity(as(a(function(of(motivation,( an(
independent(samples(tZ test(was(used(to(compare(task Z related(activity(between(groups,(
with(changes(in(selfZ efficacy((pre(to(post(training)(as(a(covariate.(We(did(not(observe(
any(group(differences(during(task(period(for(this(analysis. (
( To(identify(neural(networks(that(modulate(feedback(processing(during(motor(
learning(by(enhanced(motivation,(we(compared(between(groups(in(activation(that(was(
parametrically(modulated(by(temporal(accuracy(of(the(preceding(trial(during(feedback(
presentation.(Across(groups,(relatively(long(time(lag(performance(of(the(corresponding(
trial(was(not(significantly(associated(with(areas(of(brain(activation(during(the(feedback(
interval.(However,(comparing(between(groups,(the(control(group(revealed(activity(in(the(
cingulate(cortex(and(insula(more(strongly(associated(with(long(time(lag(performance(
compared(to(the(motivation(group,(as(shown(in(figure(6 .(Across(groups,(during(the(
feedback(interval,(relatively(short(time(lag(performance(of(the(corresponding(trial(was(
associated(with(brain(activity(in(the(posterior(and(anterior(cingulate(cortex,(caudate(and(
!
47!
!
nucleus(accumbens((figure(7).(Comparing(between(groups,(the(motivation(group(
showed(brain(activity(in(the(ant erior(cingulate(cortex(that(was(more(strongly(associated(
with(short(time(lag(performance(compared(to(the(control(group ((figure(8).(
Discussion!!
( Our(study(design(allowed(an(investigation(of(the(effects(of(enhanced(motivation(
at(three(levels(of(analysis,(inc luding(motor(skill(performance(and(learning,(motivation(
selfZ reports(and(brain(activity((fMRI),(during(practice(of(a(challenging(motor(skill(task.(
Results(demonstrate(that(motivational(statements(that(enhance(expectation(of(
performance(success(and(support(competence(improve(motor(skill(learning(of(a(
spatiotemporal(tracking(task(by(improving(temporal(accuracy(of(tracking.(Moreover,(
practice(under(motivational(conditions(showed(a(higher(association(between(activation(
in(response(to(performance(feedback(for(the(cingulate(cortex,(a(region(known(to(play(a(
role(in(control(of(attention(and(implicated(in(the(salience(netw ork.(
( We(demonstrated(that(our(experimental(paradigm(was(successful(in(activating(
brain(regions(related(to(motor(control(of( a(challenging(visuoZ motor(tasks.(Contrary(to(our(
hypothesis,(no(significant(group(differences(were(found(during(the(task(execution(
period.(
(
Relationship!of!motivation!and!motor!learning!behavior!
( The(novel(behavioral(finding(is(that(motivational(statements(that(enhance(self Z
efficacy(promote(skill(learning(on(a(tracking(task(revealed(by( improved(temporal(
!
48!
!
accuracy(and(increased(task(goal(performance((time(on(target).(The(largest(benefit(of(
enhanced(motivation(was(indicated(by(shorter(tracking(time(lag.(Time(lag(during(
immediate(and(delayed(retention(was(approximately(90(ms( for(the(motivation(group,(
which(can(be(described(as(an(anticipatory(or(predictive(response .(The(skill(to(predict(the(
upcoming(position(of(the(target( demonstrates(movement(planning(for(the(sequence(of(
movements(necessary(to(move(the(cursor(along (a(repeated(track.(Shorter(time(lag(for(
tracking(the(target(pattern(for(the(motivation(group(compared(to( the(control(group(
suggests(that(enhanced(intrinsic(motivation(promotes( motor(learning(through(feedZ
forward(processes(for(achieving(a(task(goal(and(may(reflect(heighten ed(preparation,(
anticipation(of(the(target(pattern(and(readiness(to(move((i.e.(faster(responses). (
( Our(behavioral(finding(is(in(line(with(previous(studies(reporting(greater(automatic(
movement(control(in(a(balancing(task(from(instructions(that(enhanced(learner’s(
competence((Wulf(et(al.,(2012).(Furthermore,(a(recent(theory,(OPTIMAL((Optimizing(
Performance(through(Intrinsic(Motivation(and(Attention(for(Learning),(proposes(that(
intrinsic(motivation(contributes(to(skill(learning(by(stre ngthening(goal(action(coupling.(
Taken(together,(enhanced(motivation(in(our(study(may(have(improved(implicit(skill(
learning(by(promoting(focus(on(the(task(goal,( reducing(attention(to(selfZ regulatory(
processes(that(may(interfere(with(learning(and(optimizing( the(link(between(the(objective(
of(the(task(and(the(sequence(of(actions(necessary(to(achieve(the(goal( (Wulf(and(
Lewthwaite,(2016)(.(This(effect(may(be(critical(in(motor(skills(that(require(motor(control(
such(as(playing(a(musical(instrument(or(operating(a(medical(tool,(where(learning(can(be(
improved(by(incorporating(motivational(instructions(during(training.( (
!
49!
!
( We(found(that(the(selfZ efficacy(increase(with(training(was(associated(with(
corresponding(improvements(in(performance((first(to(last(training(block),(and(was(
comparable(between(groups.(While(the(relationship(of(selfZ efficacy(and(performance(
gains(during(training(was(similar(for(both(groups,(the(significant(correlation(between(
selfZ efficacy(increase(and(time(on(target(performance(at(immediate(and(delayed(
retention(for(the(motivation(group,(but(not(for(the(control(group,(is(important.(It(suggests(
that(the(motivation(manipulation(strengthened(and(sustained(the(impact(of(self Z efficacy(
benefit(on(short(term(learning( (immediate(retention)(and(long(term(learning((i.e.(
consolidation),((delayed(retention(performance).(This(result(is(consistent(with(studies(
using(monetary(reward(that(display(enhanced(performance(on(24(hour(retention(tests(
using(similar(motor(tasks((Abe(et(al.,(2011_(Widmer(et(al.,(2016) .(
( The(association(between(performance(immediately(after(training((immediate(
retention)(and(effort(and(interest(reported(on(the(second(day( for(the(motivation(group(
but(not(for(the(control(group(is(important.(One(interpretation(is(that(the(motivational(
manipulation(induces(a(higher(value(and(importance(to(the(task,( which(in(turn(leads(to(
better(task(performance.(This(result(is(consistent(with(previous(reports(showing(a(
positive(relationship(between(high(selfZ efficacy(and(expended(effort((Bandura,(1986).(
!
Brain!activity!modulated!by!motivation!and!skill!learning!
( We(demonstrate(that(the(augmented(motor(learning(benefits(in(the(enhanced(
motivation(group(is(associated(with(changes(in(neural(activity(during(performance(
feedback(processing.(Specifically,(the(motivation(group(shows(activity(in(the(anterior(
!
50!
!
cingulate(cortex(more(strongly(associated(with( greater(temporal(accuracy(of(the(
corresponding(trial(compared(to(the(control(group. (
( Our(fMRI(results(are(aligned(with(the(interpretation(of(our(behavioral(results(
showing(reduced(temporal(error(with(enhanced(intrinsic(motivation,(suggesting(
enhanced(task(preparation(and(greater(attention(to(the(task(goal.(EventZ related(brain(
potential((ERP)(and(fMRI(studies(have(provided(evidence(that(the(anterior(cingulate(
cortex((ACC)(is(implicated(in(monitoring(performance(by(detecting(errors( (Carter,(1998_(
Critchley(et(al.,(2005)(and(controlling(attention(to(taskZ relevant(stimuli.( (
Furthermore,(neuroimaging(studies(demonstrate(that(ACC(is(a(key(region(
involved(with(emotion(processing(and( it(can(be(modulated(by(affect(and(motivation(
(Bush(et(al.,(2000).(Specifically,(cognitive(and(affective(subdivisions(of(the(ACC(have(
distinct(anatomical(connections(consistent(with(its(reported(functions.(Whereas(the(
former(has(connections(with(areas(of(the(lateral(prefrontal(cortex,(premotor(and(
supplementary(motor(areas(and(hippocampus,(and(is(implicated(in(modulating(attention(
and(motor(control_(the(affective(subdivision(is(connected(to(the(nucleus(accumbens,(
insula(and(orbitofrontal(cortex,(and(may(also(play(a(role(in(evaluating(and(regulating(
motivational(and(emotional(responses((Bush(et(al.,(2000).(Our(results(display(activation(
specifically(in(the(cognitive(subdivision(of(the(anterior(cingulate(cortex,(consistent(with(
the(reported(anatomical(projections,(indicating(a(contribution(to(executive(functions(
related(to(monitoring(performance(and(regulating(associated(behavioral(responses. (
( Our(results(provide(evidence(for(a(mechanism(of(intrinsic(motivation( that(
modulates(motor(learning(by(enhancing(brain(activity(during(the(feedback(interval(after(
!
51!
!
trials(that(have(been(performed(with( greater(temporal(accuracy.(This(is(supported(by(a(
theoretical(model(where(the(cingulate(cortex(and(insula(work(together,(in(conjunction(
with(connections(to(the(motor(cortices,(to(identify(salience(and(guide(forthcoming(
behavior((Menon(and(Uddin,(2010_(Menon,(2015) .(Our(findings(are(consistent(with(
another(study(examining(biomarkers(for(enhanced(selfZ efficacy(on(motor(performance.(
Themanson(and(colleagues(used(EEG(to(demonstrate(that(higher(selfZ efficacy(was(
associated(with(better(performance(on(a(flanker(task.(Further,(this(relationship(was(
mediated(by(higher(p30(amplitude,( also(linked(to(anterior(cingulate(activity(and( used(to(
index(taskZ relevant(attentional(control((Themanson(and(Rosen,(2015).(((
( Given(the(role(of(the(cingulate(cortex(and(insula(in(the(salience(network,(it(is(
possible(that(motivation(modulates(salience(attributed(to(performance(feedback(and(
augments(attention(to(the(overall(task(goal.( For(instance,(compared(to(neutral(
conditions,(enhanced(motivation(conditions(may(amplify(the(salience(given(to(trial(by(
trial(feedback(associated(with(task(execution(that(indicates(better(performance( and(
emphasizes(overall(performance(improvement(during(the(practice(blocks.(
This(is(in(keeping(with(previous(research(in(motor(learning(showing(that(the(
frequency(of(knowledge(of(results(can(h amper(motor(learning(if(provided(in(excess,(
possibly(by(causing(a(dependency(on(the(feedback(and( blocking(the(use(of(inherent(
feedback(from(the(produced(movement.((Winstein(and(Schmidt,(1990_(Schmidt,(1991) .(
Moreover,(Chiviacowsky(and(colleagues(have(d emonstrated(a(beneficial(effect(on(motor(
learning(for(practice(conditions(that(provide(performance(feedback(after(good(trials(
compared(to(poor(practice(trials( (Chiviacowsky(and(Wulf,(2007).(While(the(frequency(of(
knowledge(of(results((time(on(target(performance(feedback)(for(our(study(was(the(same(
!
52!
!
for(both(groups((following(every(practice(trial),(the(attention(to(and(dependency(on(
feedback(may(have(been(modulated(by(our(motivational(manipulat ion(by(emphasizing(
salience(to(feedback(after(more(successful(trials(and( attenuating(the(possible(
detrimental(effects(of(frequent(feedback(on(skill(learning. ((
( Moreover,(the(motivation(manipulation(may(impact(how(individuals(interpret(
feedback.(Instructions(and(comments(to(provide(a(positive(perspective(for(what(to(
expect(of(task(performance(throughout(practice((errors/poor(performance(is(common(in(
the(beginning(of(practice(and(performance(improves(over(time), (may(influence(the(
motivation(group(to(interpret(their(feedback(as(more(salient(when(they(are(performing(
well.(Thus,(performanceZ based(feedback(may(play(a(more(effective(role(in(guiding(the(
motivation(group’s(confidence(in(the(ability(and(expectation( to(perform(well(on(the(task. (
Compared(to(the(control(group,(participants(in(the(motivation(group(may( adopt(a(goal(
oriented(mindset(where(they(use(the(feedback(to(perform(better(and(improve(on(the(
proceeding(trials,(facilitated(by(greater(allocated(attention(available(for(task(planning,(
preparation(and(successful(motor(execution.((
( Findings(from(this(study(complement(previously(reported(neural(correlates(of(
motivation(and(motor(behavior,(suggested(by(studies(using(monetary(reward(or(food((in(
animal(studies)(to(induce(motivation.(It(is(possible(that( different(aspects(of(motivation(
function(to(achieve(improved(motor(skill(learning(behavior (by(recruiting(different(
components(of(the(salience(network,(consisting(of(brain(regions(implicated(in(the(reward(
system(such(as(the(striatum.(While(explicit(trial(by(trial(rew ard(using(money(or(food(may(
more(heavily(utilize(the(dopaminergic(system(via(the(striatum(to(enhance(motor(system(
processing(during(learning,(aspects(of(motivation,(such(as(enhanced(selfZ efficacy(
!
53!
!
expectation(for(success,(may(modulate(motor(learning(and(beh avior(by(optimizing(
attentional(resources(towards(task(salient(stimuli(and(augmenting( focus(devoted(to(
accomplish(task(goals.((
( Furthermore,(studies(that(have(examined(the(brain(processes(associated(with(
intrinsic(motivational(effects(on(cognitive(task(par adigms,(reveal(additional(insights(into(
associated(neural(mechanisms(of(motivation(and(behavior.(For(instance,(Depasque(and(
Tricomi((2015)(studied(the(impact(of(enhanced(intrinsic(motivation,(using(motivational(
interviewing(techniques(to(enhance(task(value ,(on(a(word(association(learning(task.(
They(found(that(neural(response(to(performance(feedback(in(the(posterior(cingulate(
cortex(was(associated(with(greater(gain(in(performance(after(the(motivational(
manipulation(compared(to(a(control(condition( (DePasque(and(Tricomi,(2015).(Another(
study(examined(the(effect(of(verbal(reward,(by(praising(one’s(competence,(on(a(picture(
puzzle(task,(where(participants(were(presented(with (2(art(pictures(and(were(instructed(
to(find(subtle(differences(between(the(two(presentations( (Albrecht(et(al.,(2014).(Albercht(
and(colleagues(found(higher(activation(in(the(right(lateral(prefrontal(cortex(during(task(
execution(compared(to(the(control(group(that(received(no(reward.(Taken( together,(these(
studies(suggest(areas(of(the(salience(network( that(may(be(engaged(by(motivation(to(
produce(improved(behavior.(
( In(our(experiment,(we(did(not(find(any(between(group(differences(in(task Z related(
activation.(While(this(is(contrary(to(our(hypothesis,(this(result(is(consistent(with(other(
neuroimaging(studies(of(motivation(that(have(reported(ac tivity(associated(during(
performanceZ dependent(reward(presentation((cue(or(outcome(of(reward),(which(are(
separate(from(the(task(execution(period.((
!
54!
!
( We(acknowledge(that(several(questions(remain(to(be(clarified. (It(is(not(well(
understood(how(different(aspects(of(the(salience(network(interact (with(one(another((i.e.,(
cortical(and(subcortical(regions)(to(produce(behavioral(outcomes.(Future(studies(can(
further(examine(the(interplay(of(differen t(aspects(of(motivation,(such(as(autonomy(
support,(in(motor(learning(to(investigate(the(brain(correlates(that(are(common (to(and(
differentiate(the(multifaceted(effects(of(motivation.(Furthermore,(future(neuroimaging(
studies(using(eventZ related(paradigms(can(further(examine(brain(activity(related(to(
specific(aspects(of(task(execution,(including(pre Z movement(planning,(that(may(provide(
additional(insight(into(how(motivation(influences( motor(planning(related(to(improved(
performance.(
Limitations!and!Conclusions!
( Our(study(has(limitations(that(should(be(considered.(We(utilized(an(fMRI(block(
design,(which(comes(with(the(disadvantage(of (potential(habituation,(anticipation(and(
other(strategy(effects(across(the(scanning(period.(Having(a(control(group(minimized(the(
possibility(for(habituation(and(anticipation(during(the(scanning(period(to(influence( the(
between(group(comparisons.(Nevertheless,(further(studies(can(better(address(this(
limitation(using(eventZ related(neuroimaging(paradigms.(Another(limitation(of(our(study(
regards(the(interpretation(of(neural(activity,(indicated(by(fMRI(Blood Z Oxygen(Dependent(
(BOLD)(signal.(While(BOLD(signal(impli es(activation(in(the(brain(region(of(interest,(we(
are(limited(in(interpreting(if(that(activation(represents(inhibition(or(excitation.( (
( Additionally,(to(enhance(the(integrity(and(authenticity(of(the(motivational(
manipulation,(we(did(not(have(a(baseline(mea sure(for(the(subscales(of(intrinsic(
!
55!
!
motivation(inventory.(While(we(recruited(study(subjects(from(the(university(community,(
consisting(of(students(and(employees,(in(both(our(experimental(and(control(groups,(we(
cannot(rule(out(the(variability(of(trait(and(in terZ subject(differences(in(intrinsic(motivation(
that(participants(came(in(with.(Moreover,(we(used(a(target(tracking(task(for(our(motor(
skill(learning(paradigm.(It(is(possible(that(other(motor(tasks(with(different(task(demands(
and(experimental(setups(may(be(affected(differently(to(manipulation(of(enhanced(
motivation.(Overall,(the(generalizability(of(our(findings(to(effects(of(intrinsic(motivation(in(
other(populations(and(on(learning(of(other(motor(and(non Z motor(skills(is(limited.((
( In(conclusion,(our(results(demonstrate(that(intrinsic(motivation(during(practice(of(
a(motor(task(benefits(motor(skill(learning(and(engages(neural(networks(responsible(for(
attention(regulation(and(performance(monitoring.(In(particular,(enhanced(intrinsic(
motivation(strengthens(brain(activity(in(the(cingulate(cortex(during(feedback(processing(
associated(with(greater(temporal(accuracy,(which(is(accompanied(by(better(motor(task(
performance(on(short(term(and(long(term(assessments(of(skill(learning.(Moreover,(
motor(learning(is(positively(correlated(with(the(degree(of(change(in( selfZ efficacy(
specifically(when(expectation(of(performance(success(and(competence(is(enhanced(
with(our(motivational(manipulation.(This(leads(us(to(conclude(that(conditions(that(
enhance(expectancy(of(performance(amplify(intrinsic(motivation(and(benefit(motor(skill(
consolidation(by(modulating(neural(networks(involve d(with(control(of(attention(and(
learning(processes.((
(
( (
!
56!
!
Table!1.!Participant!characteristics!
(
(
Overall!! Motivation! Control!
N! 40( 20( 20(
Age!(SD)! ! 23.83((4.63)(( 23.35((5.36)( 24.3((3.84)(
Sex,!male/female ! 15/25( 9/11( 6/14(
(
(
(
(
Table!2.!Main!activation!at!the!time!of!tracking!task!performance((task(period)(
(
Brain!region! Side !
Number!
of!voxels!
Beta!value! Z!score!
MNI!coordinates!
(x,!y,!zS!mm)!
Sensorimotor(Cortex( R,L( 28571( ( 8.72( Z 6,Z 4,58(
((Motor(Cortex( L( 2452( 0.4795( 8.387( Z 44,Z 12,62(
((Premotor(Cortex( L( 5645( 0.5898( 8.716( Z 26,Z 6,76(
Visual(Cortex( R,L( 11280( ( 7.72( 50,Z 64,(8(
Putamen( L( 750( 0.2242( 7.93( Z 24,2,8(
Anterior(Cingulate( R,(L( 917( 0.2648( 8.018( Z 4,0,50(
Insular(Cortex( R,(L( 734( 0.2091( 6.487! 34,18,10!
(
The(listed(clusters(showed(significant(activation((p<0.05)(in(relation(to(task(planning(and(
execution(across(trials.(Effect(sizes(of(BOLD(signal(increase(during(task(period(are(reported.(
Effect(size(expressed(as(mean(beta(values.(Z(values(indicate(the(local(maxima(at(the(ROI(
center.(
! !
!
57!
!
Figure!1.!Study!Procedures,!Tracking !Task!and!Trial!Sequence !
(
Motivational!manipulation!!
1)(Keep(in(mind(that(at(the(beginning(it(is(common(to(undershoot(or(overshoot(
the(target,(but(this(is(the(type(of(task(that(you(get(better(at(with(practice. !!
2)(Alright!(Your(improvement(across(the(past(trials(is(reflecting(your(learning(and(
getting(the(hang(of(it.!!
3)(Very(good!(Based(on(your(performance(so(far,(it(looks(like(you(are(g oing(to(
continue(to(improve(across(the(coming(trials.( (
4)(Alright!(Nice(work!!!
5)(Great(job!(
6)(Well(done!(
(
!
58!
!
Figure!2.!Average!self?efficacy!reports!across!time!points!and!Intrinsic!Motivation!
Inventory!subscales!ratings(
A.!
!
B.!
!
!
!
59!
!
Figure!3.!Task!Performance!(Motivation(=(red,(Control(=(blue)(
(
A.!
!
B.!
(
! (
(
!
!
60!
!
Figure!4.!Relationship!of!self?efficacy!change!and!motor!performance!at(immediate(and(
delayed(retention(for(motivation((red)(and(control((blue)(groups(
A.! Immediate!retention!performance.!Two(participants(had(missing(data(and(were(
excluded(from(this(graph.((1(missing(preZ training(time(point,(and(1(missing(postZ training(
time(point,(were(removed(from(correlation(analysis) ((
!
(
B.! Delayed!retention!performance.!Three(participants(had(missing(data(and(were(
excluded(from(this(graph.((3(missing(delayed(retention(time (point,(were(removed(from(
correlation(analysis)(
(
(
!
!
61!
!
Figure!5.!Brain!image!for!task?related!activity!across!all!participants!(combined!groups)!!
!
!
(
(
(
! !
2.3
4.3
Pre$motor(and(supplementary(
Motor(cortex
Thalamus
Visual(
cortex
Cerebellum
M1(
S1(
Putamen
!
62!
!
Figure!6.!Brain!activity!for!control!group>motivation!group!contrast!during!feedback!
presentation,!modulated!by!time!lag!performance!(associated!with!long!time!lag)!
! !
Anterior(
Cingulate(
Cortex
Insula
!
63!
!
Figure!7.!Brain!activity!across!all!participants!during!feedback!presentation,!modulated!
by!time!lag!performance!(associated!with!short!time!lag)!
!
!
Figure!8.!Brain!activity!for!motivation!group>control!group!contrast!during!feedback!
presentation,!modulated!by!time!lag!performance!(associated!with!short!time!lag)!
!
74
2.3 3.6
Posterior.
Cingulate.
Cortex
Anterior.
Cingulate.Cortex
Caudate.and.
Nucleus.Accumbens
75
2.3 3.1
Anterior.
Cingulate.Cortex
Motivation(group((>(Control(group
!
64!
!
Chapter(4:(Summary(and(Conclusion((
(
Summary!of!main!results!and!general!discussion !
The(systematic(review(of(neuroimaging(research( examining(brain(correlates(of(
motivation(and(motor(performance(revealed(common(brain(regions(implicated(in(
processing(of(reward(information,(monitoring(of(executive(function(and(signaling(of(
salient(stimuli.(From(the(nine(studies(that(were(identified(within(our(inclusion(criteria,(six(
reported(striatum(activity,(five(reported(frontal(cortex(associations(and(three(reported(
cingulate(cortex(and(insula(involvement.(
The(reports(of(striatum(and(frontal(cortex(activity(provide(evidence(for(the(role(of(
the(corticoZ striatal(circuit(in(facilitating(reward(enhanced(motor(learning.(This(
mechanism(is(further(supported(by(animal(work(that(demonstrates(that(dopaminergic(
projections(to(the(motor(cortex,(perhaps(activated(by(the(cortico Z striatal(loop,(are(
essential(for(motor(skill(learning( (MolinaZ Luna(et(al.,(2009_(Hosp(et(al.,(2011) .(Moreover,(
the(reports(of(the(cingulate(cortex(and(insula(in(the(context(of(motivated(motor(behavior(
highlight(these(brain(regions,(implicated(in(salience(processing(and(attention(allocation,(
as(important(to(integrating(motivation(related(information(to(motor(control(systems(
(Seeley(et(al.,(2007_(Ham(et(al.,(2013) .((
! Findings(from(this(dissertation(study(extend(our(current(knowledge(from(previous(
neuroimaging(work(in(understanding(neural(mechanism( of(rewardZ related(motor(
learning.(Using(a(between(subject(design,(we(investigated(the(effects(of(enhanced(
intrinsic(motivation(on(motor(skill(learning(and(brain(activity((fMRI)(during(practice(of(a(
!
65!
!
challenging(spatiotemporal(motor(task.(We(show(that(motor( skill(learning,(assessed(by(
temporal(accuracy,(is(enhanced(in(the(motivation(group(compared(to(the(control(group.(
Importantly,(our(neuroimaging(results(suggest(that(different(neural(processes(occur(
under(motivation(and(neutral(practice(conditions.(Specific ally,(activation(in(the(cingulate(
cortex(during(the(performance(feedback(interval(is(more(strongly(associated(with(good(
time(lag(performance(trials(in(the(motivation(group(compared(to(the(control(group.( (
The(present(research(adds(to(our(understanding(of(mo tivated(motor(learning(and(
the(associated(neural(correlates.(Notably,(results(from(this(study(demonstrate(that(
motivation(enhances(motor(skill(learning(and(modulates(neural(activity(in(the(cingulate(
cortex(associated(with(processing(of(feedback(during(skil l(acquisition.(Strengthened(
association(of(cingulate(cortex(activity(and(good(time(lag(performance(in(the(motivation(
group(as(compared(to(the(control(group,(suggests(that(motivation( enhances(sensitivity(
to(performance(feedback(after(successful(trials(during(practice(and(promotes(motor(skill(
learning.(
(Finally,(our(results(fit(in(with(a(recent(theoretical(model(of(motivation(and(motor(
learning,(OPTIMAL((Optimizing(Performance(through(Intrinsic(Motivation(and(Attention(
for(Learning).(OPTIMAL(proposes(that( intrinsic(motivation(promotes(motor(skill(learning(
by(strengthening(the(link(between(goals(and(actions ((Wulf(and(Lewthwaite,(2016).(The(
main(behavioral(and(neuroimaging(results(fr om(our(study(suggest(that(enhanced(
intrinsic(motivation(may(facilitate(successful(skill(learning(by(directing(attention(and(
salience(to(the(task(goal,(which(promotes(preparation(and(planning(for(upcoming(
movements,(by(modulating(brain(networks(that(control (attention(to(taskZ relevant(
feedback.(This(mechanism,(which(is(partially(rooted(in(modulation(of(brain(activity(in(the(
!
66!
!
cingulate(cortex,(may(enhance(task(preparation(and(motor(execution(during(skill(
acquisition(to(boost(motor(learning(and(consolidation. (
Implications!and!future!directions(
( This(work(has(implications(for(the(role(motivation(can(play(in( the(context(of(skill(
learning,(spanning(the(domains(of(education,(parenting,(sports(training(and(medical(
rehabilitation.(Specifically,(incorporating(strategies(that(enhance(intrinsic(motivation,(
namely(by(supporting(autonomy,(competence(and(social(relatedness,(can(benefit(
behavioral(performance(and(promote(effective(learning(in(healthy(and(disabled(
populations.(Additionally,(neurobiological(evidence(that(motivational(strategies(improve(
task(preparation(and(skill(learning,(perhaps(through(enhanced(attention(monitoring,(
provides(support(to(integrate(cognitive(training(or(meditation(to(help(individuals(more(
effectively(control(focus(of(attention(to(improve (behavioral(performance(or(alleviate(
behavioral(deficits(in(individuals(with(motor(impairments. (
( Moreover,(this(research(can(further(our(understanding(of(disorders(that(are(
impacted(by(disrupted(motivational(functioning(and(associated(neural(networks.( For(
instance,(patients(with(Parkinson’s(disease(who(have(impaired(functioning(of(the(
dopamine(system,(may(be(able(to(leverage(other(intact(networks(that(engage(
motivation,(including(the(cingulate(cortex.(Utilizing(treatments(and(therapies(that(
enhance(intrinsic(motivation(can(harness(alternative(neural(mechanisms(for(optimizing(
learning(and(behavior,(in(both(the(cognitive(and(movement(domain. (
Future(empirical(and(clinical(research(can(extend(this(work(by(investigating(the(
interplay(between(dopaminergic(reward(processes(and(other(brain(circuits(that(integrate(
motivation(in(guiding(motor(actions(and(skill(behaviors.( (
!
67!
!
Appendix(
SelfZ efficacy(questionnaire(
!
Research!Study!Questionnaire !
(
Please!indicate!how!much!you!agree!with!the!statement.!
Please(rate(how(confident(you(are(that(you(would(be(able(to(keep(the(red(box(in(the(white(target(
box(for(2(seconds?(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident)(
Please(rate(how(confident(you(are(that(you(would(be(able(to(keep(the(red(box(in(the(white( target(
box(for(4(seconds?(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident)(
Please(rate(how(confident(you(are(that(you(would(be(able(to(keep(the(red(box(in(the(white(target(
box(for(6(seconds?(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident)(
Please(rate(how(confident(you(are(that(you(would(be(able(to(keep(the(red(box(in(the(white(target(
box(for(8(seconds?(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident)(
Please(rate(how(confident(you(are(t hat(you(would(be(able(to(keep(the(red(box(in(the(white(target(
box(for(10(seconds?(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident(
Please(rate(how(confident(you(are(that(you(would(be(able(to(keep(the(red(box(in(the(white(target(
box(for(12(seconds?(
(
0( 1( 2( 3( 4( 5( 6( 7( 8( 9( 10(
(Not(confident(at(all)(( ( ( ( ( (Extremely(confident)(
! !
!
68!
!
Intrinsic(Motivation(Inventory(items(
Your(feedback(is(important(to(us.(Please(share(about(your(experience(by(responding(to(the(
questions(below.(Thank(you(
For(each(of(the(following(statements,(please(indicate(how(true(it(is(for(you,(using(the(
following(scale:(
(( ( ( 1( 2( 3( 4( 5( 6( 7(
(Not(at(all(true)(( ( ( ( (((((Very(true)((
1.( I(think(I(am(pretty(good(at(this(activity. (
2.( While(I(was(doing(this(activity,(I(was(thinking(about(how(much (I(enjoyed(it.(
3.( I(didn't(put(much(energy(into(this. (
4.( I(didn't(try(very(hard(to(do(well(at(this(activity. (
5.( I(put(a(lot(of(effort(into(this. (
6.( It(was(important(to(me(to(do(well(at(this(task. (
7.( I(was(pretty(skilled(at(this(activity. (
8.( I(tried(very(hard(on(this(activit y.(
9.( After(working(at(this(activity(for(a(while,(I(felt(pretty(competent. (
10.(This(task(was(fun(to(do. (
11.(I(thought(this(was(a(boring(task. (
12.(This(was(an(activity(that(I(couldn't(do(very(well. (
13.(I(thought(this(task(was(quite(enjoyable. (
14.(I(think(I(did(pretty(well(at(this (activity,(compared(to(other(participants.(
15.(I(am(satisfied(with(my(performance(at(this(task. (
16.(This(task(did(not(hold(my(attention(at(all. (
17.(I(enjoyed(this(task(very(much. (
18.(I(would(describe(this(task(as(very(interesting. (
Mindset(questions(
(Please(indicate(how(true(the(following(statements(are(for(you,(using(the(following(scale: (
(( ( ( 1( 2( 3( 4( 5( 6( 7(
(Not(at(all(true)(( ( ( ( (((((Very(true)((
1.( With(practice,(one(can(significantly(improve(capability(to(control(fingers(to(track(a(target. (
2.( One(has(a(certain(inherent(amount(of( capability(to(control(fingers(to(track(a(target. (
How(would(you(rate(task(difficulty(on(a(scale(of(1 Z 7((1=easy,(7=hard)(
(
1( 2( 3( 4( 5( 6( 7(
(((((((( (Easy)(( ( ( ( (((( (((((((( (Hard)((
(
(
(
!
69!
!
Exit(Interview(Questionnaire(Items(
(
How(was(your(experience(with(the(task? (
(
What(did(you(think(about(any(of(the(instructions(or(comments(you(heard(while(you(were(
in(the(scanner?(
(
(
How(was(your(control(in(moving(the(cursor?(Did(you(have(any(problems(with(the(task(or(
the(performing(the(task?(
(
Do(you(have(any(other(comments(or(feedback( you(would(like(to(share(with(us? (
(
(
How(was(your(experience(while(you(were(in(the(scanner? (
(
(
Was(the(questionnaire(understandable?((yes(or(no) (
(
Does(this(task(remind(you(of(any(physical(activity(that(you(do?(If(so,(please(describe. (
(
Did(you(come(up(with(any (strategies(that(were(more(helpful(than(others?(If(so,(please(
describe.(
(
(
(
( (
!
70!
!
Edinburgh(Handedness(Inventory(
Your(participant(ID:( ( ( (
Please(indicate(with(a(one((1)(your(preference(in(using(your(left(or(right(hand(in(the(following(
tasks.(
Where(the(preference(is(so(strong(you(would(never(use(the(other(hand,(unless(absol utely(forced(
to,(put(a(two((2).(
If(you(are(indifferent,(put(a(one(in(each(column(((1((|((1). (
Some(of(the(activities(require(both(hands.(In(these(cases,(the(part(of(the(task(or(object(for(which(
hand(preference(is(wanted(is(indicated(in(parentheses. (
Task(/(Object( Left(Hand( Right(Hand(
1.(Writing( ( (
2.(Drawing( ( (
3.(Throwing( ( (
4.(Scissors( ( (
5.(Toothbrush( ( (
6.(Knife((without(fork)( ( (
7.(Spoon( ( (
8.(Broom((upper(hand)( ( (
9.(Striking(a(Match((match)( ( (
10.((Opening(a(Box((lid) ( ( (
Total(checks:( LH(=(( RH(=((
Cumulative(Total( CT(=(LH(+(RH(=( (
Difference( D(=(RH(–(LH(=((
Result( R(=((D(/(CT)(×(100(=((
Interpretation:(
(Left(Handed:(R(<(Z 40)(
(Ambidextrous:(Z 40(≤(R(≤(+40)(
(Right(Handed:(R(>(+40)( (
!
71!
!
!
Supplementary(Material(
Root(Mean(Square(Error((RMSE)(performance(outcome(measure:(Method(and(Results (
!
Root!Mean!Square!Error!(RMSE). ((The(average(between(the(center(of(the(cursor(and(
the(center(of(the(target(was(calculated(for(each(trial.(Average(RMSE(was(calculated(for(
each(block(separately.(
(
Both(groups(demonstrated(improvements(in(RMSE(across(training(blocks(and(there(
was(no(statistically(reliable(effect(of(group(for (RMSE.(
!
(
(
!
!
72!
!
Exit(Interview(Questionnaire:(Methods(and(Results(
Exit!Interview!questionnaire.!!At(the(end(of(the(study,(participants(were(asked(a(series(of(
questions(inquiring(their(perceived(difficulty(and(concept(of( ability(mindset(for(the(task(in(
additional(to(their(personal(experience(during(the(study. !!
!
Both(groups(showed(comparable(reported(ratings(on(perceived(difficulty,(growth(mindset(and(
fixed(mindset.(
A.! Ratings!for!perceived!difficulty!
(
B.! Ratings!for!growth!and!fixed!mindset!
(
(
(
!
!
73!
!
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Creator
Beroukhim Kay, Dorsa
(author)
Core Title
Brain and behavior correlates of intrinsic motivation and skill learning
School
College of Letters, Arts and Sciences
Degree
Doctor of Philosophy
Degree Program
Neuroscience
Publication Date
09/25/2019
Defense Date
08/24/2017
Publisher
University of Southern California
(original),
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(digital)
Tag
brain,fMRI,Motivation,motor behavior,motor learning,motor performance,OAI-PMH Harvest,skill learning
Language
English
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Electronically uploaded by the author
(provenance)
Advisor
Monterosso, John (
committee chair
), Kutch, Jason (
committee member
), Lewthwaite, Rebecca (
committee member
), Winstein, Carolee (
committee member
)
Creator Email
dberoukh@usc.edu,dorsa26@gmail.com
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etd-BeroukhimK-5767.pdf (filename),usctheses-c40-435483 (legacy record id)
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etd-BeroukhimK-5767.pdf
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435483
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Beroukhim Kay, Dorsa
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texts
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University of Southern California
(contributing entity),
University of Southern California Dissertations and Theses
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The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the a...
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Tags
brain
fMRI
motor behavior
motor learning
motor performance
skill learning