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Modelling Natural Action Selection PDF

569 Pages·2011·6.35 MB·English
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ModellingNaturalActionSelection Actionselectionisafundamentalprobleminbiologyandecology.Itrequiresdetermining availablealternatives,executingthosemostappropriate,andresolvingconflictsamong competinggoalsandpossibilities. Using advanced computational modelling, this book explores cutting-edge research into action selection in nature from a wide range of disciplines, from neuroscience tobehaviouralecology,andeventopoliticalscience.Itdeliversnewinsightsintoboth detailedandsystems-levelattributesofnaturalintelligenceanddemonstratesadvancesin methodologicalpractice.Contributionsfromleadingresearcherscoverissuesincluding whetherbiologicalactionselectionisoptimal,neuralsubstratesforactionselectionin thevertebratebrain,perceptualselectionindecisionmaking,andinteractionsbetween groupandindividualactionselection. Thismajorintegratedreviewofactionselectioninnaturecontainsabalanceofreview andoriginalresearchmaterial,consolidatingcurrentknowledgeintoavaluablereference forresearchers,whileillustratingpotentialpathsforfuturestudies. AnilK.SethisaReaderintheSchoolofInformaticsattheUniversityofSussex,anEPSRC Leadership Fellow, and Co-Director of the Sackler Centre for Consciousness Science. His research crosses the fields of computational neuroscience, consciousness science, andneurorobotics.Inadditiontocontributingnewinsightsintothemechanismsofaction selection,hehasdevelopednewwaystolinkbrainactivitytoconsciousexperienceand heiswellknownforhisresearchonthestatisticalanalysisofcausality. Tony J. Prescott is Professor of Cognitive Neuroscience at the University of Sheffield whereheteachescoursesoncomputationalneuroscienceandbiomimeticrobotics.His research lies within the biological and brain sciences, and concerns understanding the evolution, development, and function of natural intelligence. He is particularly well knownforhisworkonmodellingtheneuralsubstratesforactionselection,andbuilding robotmodelsofanimalandhumanbehaviour. JoannaJ.BrysonisaReaderintheDepartmentofComputerScienceattheUniversity ofBath.Sheconductsinterdisciplinaryresearchontheorigins,structure,andconstruc- tionofhumanandanimal-likeintelligence,andisinvolvedintopicsrangingfromthe evolution of altruistic communication to the ethical role of robots in our society. She is recognised for her work in systems AI and the design of action selection. Her most recentworkiscentredontheapplicationofmodellingtounderstandingtheevolutionof humansocialstructuresandculturallyderivedbehaviourmoregenerally. Modelling Natural Action Selection Editedby ANIL K. SETH UniversityofSussex TONY J. PRESCOTT UniversityofSheffield JOANNA J. BRYSON UniversityofBath CAMBRIDGEUNIVERSITYPRESS Cambridge,NewYork,Melbourne,Madrid,CapeTown, Singapore,Sa˜oPaulo,Delhi,Tokyo,MexicoCity CambridgeUniversityPress TheEdinburghBuilding,CambridgeCB28RU,UK PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork www.cambridge.org Informationonthistitle:www.cambridge.org/9781107000490 (cid:2)C CambridgeUniversityPress2012 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithoutthewritten permissionofCambridgeUniversityPress. Firstpublished2012 PrintedintheUnitedKingdomattheUniversityPress,Cambridge AcataloguerecordforthispublicationisavailablefromtheBritishLibrary LibraryofCongressCataloguinginPublicationdata Modellingnaturalactionselection/editedbyAnilK.Seth,TonyJ.Prescott,JoannaJ.Bryson. p. cm. Includesbibliographicalreferencesandindex. ISBN978-1-107-00049-0 1.Cognitiveneuroscience. 2.Neurobiology–Computersimulation. 3.Decisionmaking–Computer simulation. I.Seth,AnilK. II.Prescott,TonyJ. III.Bryson,JoannaJ. IV.Title. QP360.5.M63 2011 612.8(cid:3)233–dc23 2011027497 ISBN978-1-107-00049-0Hardback CambridgeUniversityPresshasnoresponsibilityforthepersistenceor accuracyofURLsforexternalorthird-partyinternetwebsitesreferredto inthispublication,anddoesnotguaranteethatanycontentonsuch websitesis,orwillremain,accurateorappropriate. Contents Foreword pageviii PeterM.Todd Preface xi Listofcontributors xii 1 Generalintroduction 1 AnilK.Seth,JoannaJ.Bryson,andTonyJ.Prescott PartI Rationalandoptimaldecisionmaking 2 IntroductiontoPartI:rationalandoptimaldecisionmaking 7 AnilK.Seth,TonyJ.Prescott,andJoannaJ.Bryson 3 Doweexpectnaturalselectiontoproducerationalbehaviour? 12 AlasdairI.Houston,JohnM.McNamara,andMarkD.Steer 4 Optimisedagent-basedmodellingofactionselection 37 AnilK.Seth 5 Compromisestrategiesforactionselection 61 FrederickL.Crabbe 6 Extendingabiologicallyinspiredmodelofchoice:multi-alternatives, nonlinearity,andvalue-basedmultidimensionalchoice 91 RafalBogacz,MariusUsher,JiaxiangZhang,andJamesL.McClelland 7 Bayesianapproachestomodellingactionselection 120 MaxBerniker,KunlinWei,andKonradKo¨rding 8 Post-retrievalinhibitioninsequentialmemorysearch 144 EddyJ.Davelaar PartII Computationalneurosciencemodels 9 IntroductiontoPartII:computationalneurosciencemodels 169 TonyJ.Prescott,JoannaJ.Bryson,andAnilK.Seth vi Contents 10 Actionselectionandrefinementinsubcorticalloopsthroughbasalganglia andcerebellum 176 JamesC.Houk 11 Corticalmechanismsofactionselection:theaffordancecompetition hypothesis 208 PaulCisek 12 Towardanexecutivewithoutahomunculus:computationalmodelsofthe prefrontalcortex/basalgangliasystem 239 ThomasE.Hazy,MichaelJ.Frank,andRandallC.O’Reilly 13 Hierarchicallyorganisedbehaviouranditsneuralfoundations: areinforcement-learningperspective 264 MatthewM.Botvinick,YaelNiv,andAndrewG.Barto 14 Themedialreticularformation:abrainstemsubstrateforsimpleaction selection? 300 MarkD.Humphries,KevinN.Gurney,andTonyJ.Prescott 15 Understandingdecision-makingdeficitsinneurologicalconditions:insights frommodelsofnaturalactionselection 330 MichaelJ.Frank,AnoukScheres,andScottJ.Sherman 16 Biologicallyconstrainedactionselectionimprovescognitivecontrolinamodel oftheStrooptask 363 TomStaffordandKevinN.Gurney 17 Mechanismsofchoiceintheprimatebrain:aquicklookatpositive feedback 390 JonathanM.Chambers,KevinN.Gurney,MarkD.Humphries,andTonyJ.Prescott PartIII Actionselectioninsocialcontexts 18 IntroductiontoPartIII:actionselectioninsocialcontexts 421 JoannaJ.Bryson,TonyJ.Prescott,andAnilK.Seth 19 Agent-basedmodelsasscientificmethodology:acasestudyanalysingthe DomWorldtheoryofprimatesocialstructureandfemaledominance 427 JoannaJ.Bryson,YasushiAndo,andHagenLehmann 20 Anagent-basedmodelofgroupdecisionmakinginbaboons 454 RussellA.Hill,BrianS.Logan,WilliamI.Sellers,andJulianZappala 21 Endogenousbirthanddeathofpoliticalpartiesindynamicparty competition 477 MichaelLaver,ErnestSergenti,andMichelSchilperoord Contents vii 22 Onoptimaldecisionmakinginbrainsandsocialinsectcolonies 500 JamesA.R.Marshall,RafalBogacz,AnnaDornhaus,RobertPlanque´,TimKovacs,andNigelR.Franks 23 State-dependentforagingrulesforsocialanimalsinselfishherds 523 SeanA.Rands,RichardA.Pettifor,J.MarcusRowcliffe,andGuyCowlishaw Index 538 Colourplatesaretobefoundbetweenpp.240and241. Foreword Theseapenleadsasimplelife.Afterfloatingonthedeep-seacurrentsasajuvenile,it settlesdownontoacomfortablepatchofsandandbeginsitslargelyimmobileadultlife, growingintoafeatheryshapeandswayinginthewaterwhileensnaringwhateveredible morselspassitsway.Ithardlymovesonitsown;itjustpassivelyfilterstheworldthat goesby.Nochoicesneedbemade,fortherearenoactionstotake.Assuch,theseapen willnotfeatureprominentlyinthisbook. For other more active (and more behaviourally interesting) species, life presents a streamofdecisionpoints,atwhichactionsmustbechosen:stayormoveon,ingestor passby,approachoravoid,waitorstrike,court/acceptordecline.Theseareallessentially formsoftheexploitation/explorationtrade-offthatorganismsmustbalancethroughout their lives, whenever resources are distributed in space or time and the individual can activelyseekthem.Thisistherealmofnaturalactionselection.Howdoorganismsdo it? Action selection mechanisms are decision mechanisms. Like the study of decision mechanisms used for making inferences – a common topic in research on (human) judgement and decision making – the study of action selection mechanisms aims to uncover what the mechanisms are that people and other animals use, how they work, and when and where they work well or do not work – that is, the conditions under whichtheydoordonotproduceadaptive (orrational)behaviour. Researchonhuman inference has revealed that there are multiple decision mechanisms that people can use in particular situations – the mind’s adaptive toolbox – and often several of these can produce similar outcomes. The chapters in this book reveal the range of possible action selection mechanisms that can be used as well. Some of the simple heuristics thathavebeenidentifiedforhumaninferenceshowthatdecisionscanbemadewithout fullycomparingdifferentalternativesonmultipleincommensuratedimensions(e.g.,by ignoringconflictingdimensionsandjustdecidingonthebasisofonefactor);someof themodelsinthisbookshowsimilarshortcutsorrulesofthumbthatcanworkinaction selection.Onedifferencebetweenstandarddecision-makingresearchandthatonnatural action selection is that, in the former, the possible alternatives that can be selected in aninferencetaskaregenerallyknownorspecifiedbytheresearcher,e.g.,whichcityis larger,DetroitorMilwaukee?Thepossibleactionsthatananimalmighttakeinnature may not always be explicitly identified and, without knowing precisely the range of possibilities,studyingthemechanismsthatselectamongthembecomesmoredifficult. Foreword ix In the human inference literature, the study of the contents of the adaptive toolbox has led to a further question beyond asking how people choose between alternatives in an inferential task: how do people choose what mechanism to use to make their inferences in a particular task? That is, how do people choose which tool to use from theadaptivetoolbox?Thisisanopenandchallengingquestion,withdifferentpossible answers;sometimestheenvironmentwilldeterminethechoice,dependingonwhattypes of information are available; other times, individuals may learn through experience or can be taught what decision mechanisms are good to employ in particular settings. If therearemultiplecompetingactionselectionmechanismsinagivensituation–notjust multiplecompetingactionsthemselves–thenthiskindofmeta-selectionquestionwill alsobeimportantinthestudyofnaturalactionselection. Specifyingtheactionselectionmechanismsthatinhabitthemind’sadaptivetoolbox meansspecifyinghowtheywork–buthowcanthisbedetermined?Thisisalsoamain themeofthisbook:exploringthemethodsthatarebeingemployedatthecuttingedgeof researchforunderstandingtheoperationofactionselectionmechanisms.Experiments and field observations are necessary for generating data on action selection in action, butthedatamustbeunderstoodintermsofatheory.Herethetheoriesareinstantiated asmodels,whichbecausetheyoftenoperateatmultiplelevels–cognitive,neural,and social – are complex enough that they are best implemented as computer simulations. Ideally,multiplecompetingmodelsthenspecifyfurtherdatatobecollectedtodistinguish betweenthem,andthenewdataconstrainthemodels(andthetheoreticalunderstanding) further,inanongoingfeedbackloop.Inthisbookweseemultiplestagesofthatprocess, andthecurrentstateofunderstandingtowhichithasled. This modelling approach can thus be characterised as ‘understanding by building’. But once the action selection models have been built and understood, we can also use themodelsasawaytoexplorehowtochangethingsinimportantapplicationareas.The applications,likethemodels,canbeatdifferentlevelsofdescription.Attheneurallevel, important health implications emerge from understanding clinical conditions such as Parkinson’sdiseaseandobsessive–compulsivedisorderasdisruptionsofactionselection. And by better knowing how natural action selection works at this level, we can gain betterunderstandingofwhatisgoingonwhenthingsgowrong–tweakingthemodels toachieve‘understandingbybreaking’–andinsightsintopossiblewaysofaddressing thoseproblems. Attheindividualandsociallevel,wemaywanttohelppeoplechoosebettercourses of actions for themselves and others – avoiding the third piece of chocolate cake, or promotingtheelectionofinnovativeleaders.Againbymodellingandunderstandingthe processes of action selection at these levels of description, we can develop, and test, ideasforhowtochangetheactionselectionprocessindesireddirections,beforetrying themoutintherealworld.Thesemethodscouldinvolvegivingpeopleandgroupsnew waysofselectingtheiractions–newtoolsforchoosingwhatandwhentoeat,ornew voting mechanisms, for instance – or could rely on changing the environmental cues thattheyreceive,to‘nudge’peopleintomakingdifferentselections.Andmodellingcan help us determine which approach may be more effective, changing environments or changingmindsornorms.

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Action selection is the task of doing the right thing at the right time. It requires the assessment of available alternatives, executing those most appropriate, and resolving conflicts among competing goals and possibilities. Using advanced computational modelling, this book explores cutting-edge re
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