The evolutionary conserved neurobiology of operant learning
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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)
5
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)
8
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)
13
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