The evolutionary conserved neurobiology of operant learning

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The evolutionary conserved neurobiology of operant learning Björn Brembs Universität Regensburg http://brembs.net

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The evolutionary conserved neurobiology of operant learningBjrn BrembsUniversitt Regensburghttp://brembs.net

Pavlovian/Classical Conditioning

Tone: Conditioned stimulus CSFood: Unconditioned Stimulus US

Canonical Synaptic Plasticity

> 530 Mya

CREBCRE

Long-termmemory

cAMPPKA

CytoplasmNucleus

NtDNArutabaga

Modified from: "Comparative psychology: Evolution and development of behavior" (2nd Edition, Psychology Press, 2008)

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Operant/Instrumental Learning

CSUSBehaviorReinforcement

Canonical Synaptic Plasticity

> 530 Mya

CREBCRE

Long-termmemory

cAMPPKA

CytoplasmNucleus

NtDNArutabaga

Modified from: "Comparative psychology: Evolution and development of behavior" (2nd Edition, Psychology Press, 2008)

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Operant Learning

CS?US?

Drosophila Turning

Drosophila Turning

Behavioral variability in a constant stimulus situation: Actions, not responses

Outcomes

operantclassicalselfworld

Canonical Synaptic Plasticity

> 530 Mya

CREBCRE

Long-termmemory

cAMPPKA

CytoplasmNucleus

NtDNArutabaga

Modified from: "Comparative psychology: Evolution and development of behavior" (2nd Edition, Psychology Press, 2008)

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Action Outcome Evaluation

Brembs & Plendl (2008)

rut-AC dependent Synaptic PlasticityProtein Kinase Cworldselfrut2080 wtbrut2080 wtb HS noHSPKCi PKCi HS noHSPKCi PKCi

Intrinsic PlasticityPKC but not rut-AC involved in Aplysia self-learning

Aplysia californicaF. Lorenzetti, D. Baxter, J. Byrne (2008): Neuron 59, 815-828

Cerebellar LTDPKC involved in self-motion learning

Rochefort C, Arabo A, Andr M, Poucet B, Save E, Rondi-Reig L. (2011) Science 334(6054):385-389

Two Components

PKC and Self-Learning

> 530 MyaAplysia(Mollusks)Drosophila(Arthropods)mice(Chordates)

Where is PKC required?

Colomb & Brembs, 2016

Operant learningAction Outcome Evaluation

Generate spontaneous actions (trying out)

Shape behavior by evaluating sensory feedback

Action Outcome EvaluationFoxP2Vocal learning

FoxP is Necessary for Self-Learning

worldselfselfWTFoxPRNAi ctrl.

> 530 MyaAplysia(Mollusks)Drosophila(Arthropods)Humans/mice/birds(Chordates)

Self-Learning

???

cAMPPKA

CytoplasmNucleus

NtDNAType II cyclasePKCFoxP

Multiple MemoriesWhy have more than one learning system?

F1 2005

30Flightsimulator, turned into a learning device

PI = (tc - th) / (tc + th)

31Standardexperiment (n=22). As it was already supposed from the spontaneous preference experiment, the fly can discriminate the both pattern types from each other. Furthermore, during test, it can recognize and identify that particular pattern, which previously was associated with the heat punishment. This means, that the fly must have a visual memory. During the recognition process, the actual visual information has to match with a memory template, which was formed during the training procedure. Even the spontaneous pattern preference recquires the comparison between actual visual information and a memory template, since the fly can see only one panorama orientation at any point of time.

Learning by DoingBrembs & Heisenberg (2000) L & M

Master: operantReplay: classical

Learning by Doing

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturning

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturningWTWT 16minMBImpairedOKOK

Self-Learning Test

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturningcolorWTWT 16minMBImpairedOKOK

Self-Learning TestWTWT 16minMBOKImpairedImpaired

World-Learning Test

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturningcolorWT 8minWT 16minMBImpairedOKOK

Self-Learning TestWT 8minWT 16minMBOKImpairedImpaired

World-Learning Testhabit formation

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturningcolorWT 8minWT 16minFoxP 16minImpairedOKImpaired

Self-Learning TestWT 8minWT 16minMBOKImpairedImpaired

World-Learning Test

MBs Gating Habit Formation

TrainingTest

composite:turning + colorturningcolorWT 8minWT 16minMB 8minImpairedOKOK

Self-Learning TestWT 8minWT 16minMB 8minOKImpairedImpaired

World-Learning TestBrembs, 2009MB247 or 17D driving TNT

Multiple Interacting Learning SystemsBrembs (2009) Curr BiolColomb & Brembs (2010) Commun Integr BiolBrembs (2010) Proc Roy SocBrembs (2011) Behav Proc

Degrees of PredictabilitySpontaneousActivity independent of sensory inpute.g. explorationMixedComplex stimuli/variable actionsBehavioral setEndogenous evoked responsesClosed loopEvokedFixed stimulus/response at fixed latencye.g. reflex, habitStimulus-response couplingPredictability

Degrees of EfficiencySpontaneousActivity independent of sensory inpute.g. explorationMixedComplex stimuli/variable actionsBehavioral setEndogenous evoked responsesClosed loopEvokedFixed stimulus/response at fixed latencye.g. reflex, habitStimulus-response couplingPredictability/Efficiency

Evolutionary ChallengeFlexibilityEfficiency

Ecological ChallengeExplorationExploitation

Psychological ChallengeGoal-directedHabitual

KahnemannThinking SlowThinking Fast