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AnUpsidetoRewardSensitivity:TheHippocampusSupportsEnhancedReinforcementLearninginAdolescence:Neuron

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AnUpsidetoRewardSensitivity:TheHippocampusSupportsEnhanced
ReinforcementLearninginAdolescence
JulietY.Davidow
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,KarinFoerde,AdrianaGalvn,DaphnaShohamy 7,

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DOI:http://dx.doi.org/10.1016/j.neuron.2016.08.031 |

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Highlights

Newevidenceforaroleforthehippocampusinreinforcementlearningin
adolescents

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Figure1

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Enhancedcooperationbetweenmultiplelearningsystemsintheadolescent
brain

Associationsbetweenlearningandmemoryinbehaviorandbrainfor
adolescents

Summary
Adolescentsarenotoriousforengaginginrewardseekingbehaviors,atendency
attributedtoheightenedactivityinthebrainsrewardsystemsduring
adolescence.Ithasbeensuggestedthatrewardsensitivityinadolescencemight
beadaptive,butevidenceofanadaptiverolehasbeenscarce.Usinga
probabilisticreinforcementlearningtaskcombinedwithreinforcementlearning
modelsandfMRI,wefoundthatadolescentsshowedbetterreinforcement
learningandastrongerlinkbetweenreinforcementlearningandepisodicmemory
forrewardingoutcomes.Thisbehavioralbenefitwasrelatedtoheightened
predictionerrorrelatedBOLDactivityinthehippocampusandtostronger
functionalconnectivitybetweenthehippocampusandthestriatumatthetimeof
reinforcement.Thesefindingsrevealanimportantroleforthehippocampusin

Figure1

BehavioralTasktoAssessTrialbyTrialIncrementalLearningand
EpisodicMemory
(A)Learningphase:oneachtrial,acentrallypresentedcue
appearedbelowtwotargets.Participantspressedabuttonto
predictwhichflowerabutterflywouldlandonandreceived
probabilisticreinforcementalongwithatrialuniquepictureofa
commonplaceobject.
(B)Memorytest:participantssawapictureofanobject,judged
whetherthepicturewasoldornew,andthenratedtheirlevelof
confidenceinthatchoice.
ViewLargeImage|ViewHiResImage|DownloadPowerPoint
Slide

reinforcementlearninginadolescenceandsuggestthatrewardsensitivityin
adolescenceisrelatedtoadaptivedifferencesinhowadolescentslearnfrom
experience.

Introduction
Adolescentsarehighlysensitivetoreward(Andersenetal.,1997,Brenhouse
etal.,2008,Galvnetal.,2006,SomervilleandCasey,2010,vanDuijvenvoorde
etal.,2014),whichhasbeenlinkedtotheemergenceofmaladaptivebehaviors
(BrenhouseandAndersen,2011,Galvn,2013,Spear,2000).Ithasbeen

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suggestedthatthisrewardsensitivitymayalsobeadaptivebypromoting
learningandexploration,whicharecriticalfortransitioningtoindependence
(Casey,2015,Spear,2000).However,evidenceforenhancedlearningin
adolescenceandassociatedneuralmechanismshaveremainedelusive.We
soughttotestthehypothesisthatadolescentswouldbebetterthanadultsat
learningfromreinforcementandthatthisbenefitwouldberelatedtoenhanced
activityinbrainregionsthatsupportlearningandmemory,particularlythe
striatumandthehippocampus.
Advancesinunderstandingneuralmechanismsofreinforcementlearningin
adultshaveleveragedcomputationalreinforcementlearningmodelstoquantify
trialbytriallearningsignalsinthebrain(Dawetal.,2005,Dawetal.,2011,
ODohertyetal.,2003).Suchmodelshighlighttheimportantroleofprediction
errors(PEs),whichreflecttheextenttowhichreinforcementreceivedonagiven
trialdeviatesfromwhatisexpected.Byreflectingtrialbytrialdeviationsbetween
predictionsandoutcomes,predictionerrorsprovidealearningsignalthatupdates
subsequentbehavior.fMRIstudiesinadultsandadolescentshaveshownthat
predictionerrorscorrelatewithbloodoxygenleveldependent(BOLD)activityin
thestriatum(e.g.,Christakouetal.,2013,Cohenetal.,2010,Hareetal.,2008,
ODohertyetal.,2003,vandenBosetal.,2012).Despitesomereportsof
enhancedstriatalactivityinadolescents,reportsofdevelopmentaldifferencesin
predictionerrorrelatedstriatalactivityaremixed(Christakouetal.,2013,Cohen
etal.,2010,vandenBosetal.,2012),andsofar,nonehaveshownalink
betweenenhancedstriatalBOLDactivityinadolescentsandenhancedlearning.
Thissuggeststhat,totheextentthatadolescentsrewardsensitivitycouldbe
relatedtobenefitsforlearning,thesemaybeaccountedforbyotherbrain
systems.
Anaturalbraincandidateregionforsupportingreinforcementlearningin
adolescenceisthehippocampus,knownforitsroleinlongtermepisodic
memory(e.g.,Davachi,2006,Gabrieli,1998,Squireetal.,2004).The
hippocampusalsocontributestorewardrelatedbehaviors,including
reinforcementlearning,rewardguidedmotivation,andvaluebaseddecision
Summary
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making.Studiesinadultsshowthatthehippocampusandthestriatuminteract

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cooperativelytosupportbothepisodicencodingandreinforcementlearning
(Adcocketal.,2006,Bunzecketal.,2010,WimmerandShohamy,2012).These

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findingssuggestthatrewardsensitivityinadolescencecouldberelatedto
enhancedhippocampalactivity,tobetterreinforcementlearning,andtobetter

Figure1

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episodicmemoryforrewardingevents.But,sofar,theroleofthehippocampus
inreinforcementlearninginadolescencehasnotbeenstudied.
WeusedalearningtaskincombinationwithfMRIandreinforcementlearning
modelstoaddressthisgap.Wehypothesizedthat,comparedtoadults,(1)
adolescentswouldbebetteratlearningfromreinforcingoutcomes(2)
adolescentswouldshowagreaterrelationbetweenreinforcementlearningand
episodicmemoryforrewardingeventsduringlearningand(3)thesedifferences
inlearningwouldberelatedtoenhancedactivityinthehippocampusand
strongercouplingbetweenthehippocampusandthestriatum.
Participantslearnedincrementally,basedontrialbytrialreinforcement,to
associatecueswithoutcomes(Figure1A).Theassociationbetweencuesand
outcomeswasprobabilistic,requiringcontinualuseofreinforcementtoupdate
choices.Reinforcementwassimplythewordcorrectorincorrectandwasnot
motivatedbymonetaryincentivestoavoidconfoundsrelatedtothemotivational
significanceofmonetaryrewardacrossagegroups.Totestepisodicmemoryfor
reinforcementevents,weincludedauniquepictureofanobjectthatwas

Figure1

BehavioralTasktoAssessTrialbyTrialIncrementalLearningand
EpisodicMemory
(A)Learningphase:oneachtrial,acentrallypresentedcue
appearedbelowtwotargets.Participantspressedabuttonto
predictwhichflowerabutterflywouldlandonandreceived
probabilisticreinforcementalongwithatrialuniquepictureofa
commonplaceobject.
(B)Memorytest:participantssawapictureofanobject,judged
whetherthepicturewasoldornew,andthenratedtheirlevelof
confidenceinthatchoice.
ViewLargeImage|ViewHiResImage|DownloadPowerPoint
Slide

incidentaltothereinforcementitselfineachoutcome(Figure1B).Thisdesign
allowedustomeasure(1)incrementallearningbasedontrialbytrial
reinforcement,(2)episodicmemoryforreinforcementevents,whicharepositive
versusnegative,and(3)theroleofthehippocampusandthestriatuminboth
formsoflearning.

Results

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EnhancedReinforcement
LearninginAdolescents
Wetestedwhetheradolescents(n=41,1317yearsold)differedfromadults
(n=31,2030yearsold)atlearningfromreinforcements,comparing(1)overall
performanceand(2)estimatedlearningratesfromthereinforcementlearning
model.Learningperformancewasquantifiedasthepercentoftrialsforwhich
participantsrespondedwiththeoutcomemostoftenassociatedwithagivencue
(e.g.,Poldracketal.,2001,Shohamyetal.,2004).Arepeatedmeasures(RM)
ANOVA(blockgroup)revealedthatbothagegroupsshowedsignificant
learning,but,consistentwithourprediction,adolescentslearningexceededthat
ofadults(Figure2Amaineffectofblock:F3,210=20.2,p=0.000blockgroup
interaction:F3,210=4.04,p=0.008).Similarresultswerefoundforoptimal
choicebytrial(mixedeffectregression,maineffectoftrial:z=7.13,p=0.000
grouptrialinteraction:z=2.97,p=0.003),andwealsofoundabetterfitof
theinteractionmodel(2=8.2,p=0.004)afterpenalizingformodelcomplexity
(Akaike,1974).
Tofurthercharacterizetrialbytrialresponses,weappliedastandard
reinforcementlearningmodeltoeachparticipantschoicedata(Equation1in
SupplementalExperimentalProcedures ).Wechosetofitacanonicalmodel,

whichrepresentsastandardclassofmodelsusedextensivelyinstudiesofbrain
correlatesofrewardpredictionerrorsinadults(seeDawetal.,2011).We
estimatedalearningrateparameterforeachparticipant(),whichreflectsthe
extenttowhichfeedbackoneachtrialisusedtoupdatelaterchoices.Here,a
lowerlearningrateisbetterbecausetheprobabilisticassociationsbetweencues
andtargetsarefixedalowerlearningratesuggeststhatlearningisguidedby
accumulatingevidenceoveragreaternumberoftrialsratherthanshifting
behaviorbasedontheoutcomeofanysingletrial(e.g.,Daw,2011).
Importantly,themodelprovidedagoodfittotheobservedbehavioracrossboth
groups(onewayttestcomparinganullmodel,t71=39.70,p=0.000,Akaikes
InformationCriterion[AIC]usedtopenalizemodelcomplexity),andthemodel
fitsdidnotdifferbetweenthem(independentsamplesttest,t
Summary
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References
70=1.35,p=0.2).

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Consistentwiththeiroverallbetterlearning,adolescentshadalowerlearningrate

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thanadults(t70=3.0,p=0.004Figure2B),indicatingmoreincremental
learning.Moreover,acrossgroups,therewasasignificantnegativecorrelation

Images/Data

betweenlearningrateandimprovedperformanceonthetask(r70=0.43,p=
0.000FigureS1 A),indicatingthatlowerlearningrateswereindeedrelatedto

Figure1

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betterperformance.Reactiontimesdecreasedovertimeforbothgroups,withno
differencesbetweenthem,suggestingthatdifferencesinlearningarenotdueto
generaldifferencesinresponsestotaskdemands(FigureS1 B).

MemoryPositivityBiasin
AdolescentsandAdults
Wefirstassessedepisodicmemoryforthetrialuniqueobjectsthatwere
presentedduringlearning,separatingtrialsbywhethersubjectshadbeenshown
positive(correct)versusnegative(incorrect)outcomes.Wefoundasignificant
effectofreinforcement(RMANOVA,F1,70=24.6,p=0.000noeffectofgroup,
F1,70=1.6,p=0.2nointeraction,F1,70=1.2,p=0.3Figure2CSupplemental
InformationTableS1 ),indicatingthatbothgroupsshowedapositivitybias
bettermemoryforpositive,ratherthannegative,reinforcementevents.

TrialbyTrialPrediction
ErrorsAreAssociatedwith
EpisodicMemoryin
Adolescents,butNotin
Adults

Figure1

BehavioralTasktoAssessTrialbyTrialIncrementalLearningand
EpisodicMemory
(A)Learningphase:oneachtrial,acentrallypresentedcue
appearedbelowtwotargets.Participantspressedabuttonto
predictwhichflowerabutterflywouldlandonandreceived
probabilisticreinforcementalongwithatrialuniquepictureofa
commonplaceobject.
(B)Memorytest:participantssawapictureofanobject,judged
whetherthepicturewasoldornew,andthenratedtheirlevelof
confidenceinthatchoice.
ViewLargeImage|ViewHiResImage|DownloadPowerPoint
Slide

Wenexttestedwhetherreinforcementlearningmeasureswererelatedto
episodicmemoryusingmodelderivedestimatesoftrialbytrialpredictionerrors
()(Equation1inSupplementalExperimentalProcedures ).Predictionerrors
provideanestimateofhowsurprisingeachtrialsoutcomewas,whichweused
asawithinparticipantregressorforbothbehavioralandbrainimaginganalysis.
Wefoundthatpredictionerrorswererelatedtomemoryaccuracyandthatthis
effectsignificantlyinteractedwithgroup(mixedeffectregressioninteraction:PE
group,z=2.4,p=0.02nomaineffectofPE,p=0.2orgroup,p=0.7).This

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interactionreflectedasignificantrelationshipbetweenpredictionerrorand
memoryamongtheadolescents(z=5.2,p=0.000Figure2D),butnotthe
adults(z=1.3,p=0.2).Thus,inadolescents,butnotadults,episodicmemory
foroutcomeswasrelatedtopredictionerrors.Asimilareffectwasfoundforthe
relationshipbetweenreinforcementlearningandthepositivitybiasinepisodic
memoryacrossparticipants(FiguresS1 CandS1D).

PredictionErrorSignalsin
theHippocampusin
Adolescents
Asubsetof25adolescentsand22adultsunderwentfMRIwhileperformingthe
learningtask(behavioraleffectsinthefMRIgroupwerethesameasinthefull
behavioralsampleseeFiguresS1 ES1I).Tointerrogatethebrainsystems
underlyingdifferencesinbehaviorbetweengroups,weregressedprediction
errorsagainstBOLDactivitywithineachparticipantandcomparedthegroupsin
regionsofaprioriinterestinthehippocampusandthestriatum(forwholebrain
results,seeTableS2 foranalysesofvalueintheventromedialprefrontalcortex
[vmPFC],seeSupplementalInformation).
WefoundthatpredictionerrorswerecorrelatedwithBOLDactivityinthestriatum
inbothgroups,withnosignificantdifferencesbetweenthem(seeFigureS2 D
TableS2 ).Inthehippocampus,bycontrast,adolescentshadsignificantlygreater

predictionerrorrelatedBOLDactivitythanadults(Figure3CFigureS2 C).
Giventhebehaviorallinkbetweenreinforcementsandmemoryinthe
adolescents,weinvestigatedwhetherepisodicmemorywasrelatedtofunctional
connectivitybetweenthehippocampusandthestriatum.Weuseda
psychophysiologicalinteraction(PPI)analysiswiththetimeseriesfroma
hippocampalseedasthephysiologicalvariable(Figure4A)andreinforcement
valenceoftheoutcomeevent(correct>incorrect)asthepsychologicalvariable.
Wefoundsignificantconnectivitybetweenthehippocampusandtheputamenin
adolescents(butnotadults)thatwasgreaterforcorrectthanincorrectoutcomes
(z=2.68,familywiseerror(FWE)p<0.01,155voxels,peak[16,10,6]
Summary

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Figure4B).Wethenextractedtheinteractionvalueforeachparticipantfromthe

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PPIandcorrelatedthismeasureoflearningrelatedconnectivitywithan
independentwithinparticipantbehavioralmeasureofmemorypositivitybias
(Figure4C).Wefoundasignificantcorrelationbetweenconnectivityduring
learningandtheextenttowhichmemoriesforpositivereinforcementeventswere

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Figure1

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enhancedfortheadolescents(r=0.62,p=0.000),butnottheadults(r=0.05,
p=0.84),andasignificantdifferenceinthecorrelationsbetweenthegroups
(comparisonofFisherztransformedcorrelationcoefficientsz=2.16,p=0.03).

Discussion
Thenegativeimplicationsofrewardsensitivityinadolescentshavebeenwell
documented,butmuchlessisknownaboutthepossibleadaptivesidefor
learning.Ourresultsshowthatadolescentswerebetteratlearningfrom
outcomes,outperformingadults.Wealsofoundthatinadolescents,butnot
adults,trialbytrialreinforcementlearningisrelatedtoepisodicmemoryfor
reinforcementevents,suchthatmemorywasbetterforsurprisinglypositive
versusnegativeoutcomes.Thesebehavioralbenefitswererelatedtoheightened
predictionerrorrelatedBOLDactivityinthehippocampusandtostronger
functionalconnectivitybetweenthehippocampusandthestriatumatthetimeof
reinforcement.Finally,onlyintheadolescents,functionalconnectivitybetween
theselearningsystemswasrelatedtotheextentofbiastowardbettermemory
forpositivereinforcementevents.

Figure1

BehavioralTasktoAssessTrialbyTrialIncrementalLearningand
EpisodicMemory
(A)Learningphase:oneachtrial,acentrallypresentedcue
appearedbelowtwotargets.Participantspressedabuttonto
predictwhichflowerabutterflywouldlandonandreceived
probabilisticreinforcementalongwithatrialuniquepictureofa
commonplaceobject.
(B)Memorytest:participantssawapictureofanobject,judged
whetherthepicturewasoldornew,andthenratedtheirlevelof
confidenceinthatchoice.
ViewLargeImage|ViewHiResImage|DownloadPowerPoint
Slide

Thisisthefirstdemonstrationofaroleforthehippocampusinreinforcement
learninginadolescents.Ourresultsimplythat,asadolescentsnavigatethrough
newlifeexperiences,learningfromreinforcementislinkedtohowepisodic
memoriesareshapedandtotheextenttowhichtheyarebiasedtowardencoding
moreofthegoodthanthebad.Thisfeatureoflearningisimportanttoconsiderin
relationtodecisionmakingbecauseitspeakstothesortsofbiasesthat
adolescentsmayencounterwhentheydrawonpriorexperiencetoinformcurrent
decisions.

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AnUpsidetoRewardSensitivity:TheHippocampusSupportsEnhancedReinforcementLearninginAdolescence:Neuron
Itisimportanttonotethattheadolescentsinourstudywerenotbetteratall
typesoflearningrather,thebenefitswereselectivetoreinforcementbased
updatingandrewardrelatedmemory.Overall,episodicmemoryinthe
adolescentswasnotbetterthaninadults,andtherewerenodifferencesbetween
thegroupsinmemoryforjustpositiveorjustnegativelearningevents.Instead,
thegroupsdifferedspecificallyinthestrengthoftheinteractionbetweenthese
twoformsoflearning.
Thesefindingssuggestthat,inadolescents,thereislessdifferentiationbetween
differentformsoflearningandtheirneuralsubstrateswhencomparedwith
adults.Onepossibleinterpretationofthisfindingisthatitmayberelated,inpart,
totheknowndelayindevelopmentofprefrontalcontrolmechanismsin
adolescence(SomervilleandCasey,2010forreview).Althoughitisnotknown
preciselyhowthearbitrationbetweendifferentlearninganddecisionsystems
takesplaceintheadultbrain,ithasbeensuggestedthattheprefrontalcortex
mayplayanimportantrole(Dawetal.,2005,PoldrackandPackard,2003).
Indeed,aninfluentialmodelofadolescentdecisionmakingpositsadynamic
imbalancebetweenappetitivemotivationalbrainsystems,includingthestriatum,
andinhibitorycontrolsystemsintheprefrontalcortex(Galvn,2013,Somerville
andCasey,2010).Ourfindingsextendthisframeworkandshowthatthestriatum
maybejustonelearningsystem,alongwiththehippocampus,thathasrelatively
greaterinfluenceduringadolescence.Specifically,ourfindingssuggestthatthe
functionaldevelopmentofmidbraindopaminergicrewardsystemsandtheir
connectivitywiththestriatumandthehippocampusinadolescenceispositioned
toaffectbothstrengtheningofrewardguidedhabitsandactions,aswellas
episodicmemoryformotivationalevents.Futurestudieswillneedtoassessthe
roleofcontrolandflexibilitytoidentifywhetherprefrontalsystemsregulatethe
interactionsbetweenthestriatumandthehippocampus.
Thecurrentstudyaimedtoevaluatethelinkbetweenreinforcementlearningand
episodicmemorybyconcurrentlypresentingincidentaltrialuniquestimuliwith
reinforcement.Animportantdirectionforfutureresearchwillbetodetermine
whetherthesefindingsextendtogoaldirectedepisodicencoding.Inadults,
Summary
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striatalactivityhasbeenshowntorelatetogoaldirectedmodulationofepisodic

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memory(Hanetal.,2010).Priorworkinadolescentshasshowngreater
sensitivitytorewardpredictivecuesinthestriatum(Galvnetal.,2006).

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Togetherwiththecurrentfindings,thissuggeststhatgoaldirectedcue
processinginadolescentsmayelicitgreatercooperationbetweenthe

Figure1

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hippocampusandthestriatumandbettergoaldirectedencoding.Thispossibility
remainstobetested.
Anotherimportantquestionishowsubregionsofthestriatumcontributeto
learningandinteractwiththehippocampus.Wefoundpredictionerrorrelated
BOLDactivityintheventralstriatum,ashasbeenshownrepeatedly(e.g.,Bartra
etal.,2013,ClitheroandRangel,2014).Thisregionisconnectedwiththe
hippocampus(e.g.,HaberandKnutson,2010)andinteractswithitfunctionally
(e.g.,KahnandShohamy,2013).However,ourfunctionalconnectivityanalysis
revealedactivityinaseparateregionintheputamenthatcorrelatedwiththe
hippocampus.Researchinadultsidentifiedasimilarregiondisplaying
connectivitywiththehippocampusduringcuevaluelearning(Wimmeretal.,
2014).WhilefunctionalconnectivityinBOLDdatadoesnotnecessarilyreflect
anatomicalconnectivity,thesefindingsraiseimportantquestionsforfuturework
abouttheinteractingcircuitssupportingreinforcementlearningandepisodic
memoryinadolescence.
Ourfindingsaregenerallyconsistentwithstudiesofepisodicmemoryin

Figure1

BehavioralTasktoAssessTrialbyTrialIncrementalLearningand
EpisodicMemory
(A)Learningphase:oneachtrial,acentrallypresentedcue
appearedbelowtwotargets.Participantspressedabuttonto
predictwhichflowerabutterflywouldlandonandreceived
probabilisticreinforcementalongwithatrialuniquepictureofa
commonplaceobject.
(B)Memorytest:participantssawapictureofanobject,judged
whetherthepicturewasoldornew,andthenratedtheirlevelof
confidenceinthatchoice.
ViewLargeImage|ViewHiResImage|DownloadPowerPoint
Slide

development(Ghettietal.,2010,Ofenetal.,2012).Previousresearchhas
shownthatadolescentshaveadultlikerecognitionmemory(Ghettietal.,2010),
whereasyoungerchildrenhaveworseepisodicencoding(Ghettietal.,2010)and
retrieval(DeMasteretal.,2014,Lloydetal.,2009).Findingsregarding
developmentalchangesinthehippocampushavebeenmixed.Studiesofitem
recognitionreportnodifferencesinthehippocampusduringdevelopment(Ofen
etal.,2007).Butotherstudiesindicatethatchangesinthehippocampusdo
continueintoadolescence(Daughertyetal.,2016,Leeetal.,2014)andare

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relatedtodifferencesinassociativememoryperformanceinadolescents(Ghetti
etal.,2010,DeMasteretal.,2014).
Manynewexperiencesoccurduringadolescence.Someworksuggeststhat,at
leastwhenlookingbackfromadulthood,adolescenceisatimeinwhich
particularlypowerfulandpositivememoriesareformed(HaqueandHasking,
2010,RubinandBerntsen,2003,Thomsenetal.,2011).Ofcourse,adolescence
isalsoatimewhenpsychopathologymaybegintoemerge(Caseyetal.,2015,
Ernstetal.,2009,PadmanabhanandLuna,2014).Bothperspectivesemphasize
theimportanceoflearningfromexperiencesduringthistimeofdevelopment.The
heightenedsensitivityofstriatallearningsystemsmayputrewardseeking
actionsintooverdrivebutcanalsoconferabenefitinlearningfrompredictable,
butvariable,outcomes,asweshowhere.Ourfindingsdemonstratethatthis
reinforcementsensitivityhasimplicationsforwhatkindsofmemoriesareformed
inadolescenceandhowthesememoriesdrivebehavior.

AuthorContributions
J.Y.D.,K.F.,A.G.,andD.S.designedthestudyJ.Y.D.andA.G.collectedthe
dataJ.Y.D.analyzedthedataunderthesupervisionofD.S.,A.G.,andK.F.and
J.Y.D.,K.F.,A.G.,andD.S.wrotethemanuscript.

Acknowledgments
TheauthorsthankKatherineDuncan,BradDoll,andRaphaelGerratyfor
assistancewithdataanalysisandLucyOwen,ElizabethPierce,KathyDo,
EmilyBarkleyLevenson,DianeGoldenberg,KaitlynBreiner,andElicaRahdar
forhelpingwithdatacollection.ThisresearchwassupportedinpartbyThe
NationalScienceFoundation(DGE1144155toJ.Y.D.,BCS0963750toA.G.,
andCareerAward0955494toD.S.).

Supplemental
Information
Summary
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DocumentS1.SupplementalExperimentalProcedures,FiguresS1andS2,and

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TablesS1andS2

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Received:December23,2015Receivedinrevisedform:July10,2016Accepted:August10,2016Published:October5,2016

2016ElsevierInc.PublishedbyElsevierInc.Allrightsreserved.

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