CHAPTER 8 COMPARATIVE ANALYSIS OF ASSOCIATIVE LEARNING
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1 CHAPTER 8 COMPARATIVE ANALYSIS OF ASSOCIATIVE LEARNING
2 A tentative classification of learning phenomena Learning Phenomena General Specific Nonassociative Habituation Short term Long term? Sensitization Short term Long term? Associative/Cognitive Classical conditioning Instrumental conditioning Rule learning Social learning Human language Song learning Imprinting
3 Comparing classical and instrumental conditioning (a) Classical conditioning Stimulus 1 (CS) Stimulus 2 (US) Response 2 (CR) (b) Instrumental conditioning Stimulus 1 (S D ) Response 1 (R i ) Stimulus 2 (S R ) Response 1 (UR) Classical conditioning: a mechanism for predicting impending events. Response 2 (R c ) Instrumental conditioning: a mechanism for producing outcomes.
4 Comparative analysis of learning and cognition Comparative analysis: Basic concepts
5 Comparative analysis of learning and cognition Video Duration: 19:29 min
6 Key issues in the comparative analysis of learning and cognition Issue # 1: distinguish between learning phenomena and learning mechanisms Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Issue # 3: divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to contextual variables.
7 Key issues in the comparative analysis of learning and cognition Issue # 1: distinguish between learning phenomena and learning mechanisms Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Issue # 3: divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to contextual variables.
8 Issue # 1: distinguish between learning phenomena and learning mechanisms Mechanistic Level Example Modular Representation Psychological S S learning S S S S APPETITIVE FEAR Neurobiological Coincidence detectors CEREBELLUM AMYGDALA Neurochemical Cell-molecular Synaptic receptors Secondmessenger systems NMDA camp GABA ARACHIDONIC ACID SPECIFICITY
9 Key issues in the comparative analysis of learning and cognition Issue # 1: distinguish between learning phenomena and learning mechanisms Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Issue # 3: divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to contextual variables.
10 Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Avoidance learning in rats Lesions in the basolateral nucleus of the amygdala. Sham: simulated lesion. Two-way active avoidance training. Coover et al., 1973, J Comp Physiol Psychol, 85,
11 Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Avoidance learning in goldfish Master-yoked design Impaired by MP but no LP lesions Portavella et al., 2003, Physiol. Behav. Portavella et al., 2004, Neurosci. Let. MP: medial pallium (amygdala homologous) LP: lateral pallium (hippocampus homologous)
12 Homology or homoplasy? Some learning phenomena found in mammals and honeybees Overlearning extinction effect Partial delay of reinforcement extinction effect Overshadowing Potentiation Summation Within-compound associations Compound uniqueness Conditional discrimination Successive negative contrast Rats Partial reinforcement extinction effect Positive behavioral contrast Progressive improvement in spatial reversal learning Dimentional transfer Second-order conditioning Spontaneous recovery in extinction Latent inhibition US preexposure effect Escape and avoidance learning Honeybees >600 Mya (a planaria-like animal) Bitterman, 1988, Vertebrate-invertebrate comparisons.
13 Key issues in the comparative analysis of learning and cognition Issue # 1: distinguish between learning phenomena and learning mechanisms Issue # 2: homology of learning mechanisms is demonstrated when equivalent phenomena are shown to depend on the same processes at all levels of analysis. Issue # 3: divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to contextual variables.
14 Issue # 3: divergence in learning mechanisms is demonstrated when species differences in learning phenomena cannot be attributed to contextual variables. Determinants of performance Learning factors Signal/Response strength Contextual variables Sensory/Perception detection Motivation Motor control Acquired behavior Memory retrieval
15 Is this sufficient evidence for divergence? Rumbaugh & Pate, 1984, Evolution of cognition in primates.
16 Two views of divergence Ecological view: Learning mechanisms solve specific ecological problems Species vary in their ecology Thus, divergence in mechanisms should be widespread And associative selectivity should be common Examples: Taste aversion learning seems to violate equipotentiality Imprinting and song learning occur during a sensitive period Some species show selectivity for a specific sensory dimension
17 Two views of divergence General-process view: Most ecological niches involve causality, space, and time Learning mechanisms deal with such common dimensions Thus, divergence in mechanisms should be rare And similar phenomena should appear in very different lineages Examples: Contextual factors account for many examples of divergence Conditioning occurs in all animals with a CNS Conditioning may be involved in some otherwise specialized cases
18 Morphological divergence in Hawaiian honeycreepers Do learning mechanisms diverge like the beaks of these honeycreepers? Or are they as conservative as the eyes or the feathers of these honeycreepers?
19 Comparative analysis of learning and cognition Case study: Surprising nonreward
20 Phylogenetic relationships among fish, amphibians, and reptiles Osteichthyes (Goldfish) Amphibia (Toads) Aves (Pigeons) Reptilia (Turtles) Mammalia (Rats)
21 Classic learning theory Thorndike, 1898, Psychol Rev
22 Strengthening-weakening mechanism Stimulus Response Incentive Stimulus Response Nothing Thorndike, 1911, Animal ingelligence
23 Emotional (and cognitive) factors Tinklepaugh, 1928, Calif Pub Psychol Drawing by Katsuo & Chiharu Tomita
24 Successive negative contrast (SNC) Preshift Postshift Group Downshifted (9) Bran mash (6) Sunflower seeds Unshifted (9) Sunflower seeds (6) Sunflower seeds Sunflower seeds Sunflower seeds (for both groups) Bran mash SNC Elliott, 1928, Univ Cal Pub Psychol
25 Surprising nonreward Cues predict a larger or more preferred reward than that actually occurring. Surprising nonreward promotes two kinds of learning: Allocentric learning: about a change in the environment (cognitive). Egocentric learning: about the organism s reaction to that change (emotional). Papini, 2003, Brain Behav Evol
26 Reversed SNC in goldfish (not shown), toads, turtles, and pigeons Group Preshift Postshift Downshifted Large Small Unshifted Small Small Reversed SNC Reversed MREE Reversed PREE Papini, 2006, Jap J Anim Psychol
27 Phylogenetic relationships among fish, amphibians, and reptiles Osteichthyes (Goldfish) Amphibia (Toads) Aves (Pigeons) Reptilia (Turtles) Mammalia (Rats)???? Allocentric + Egocentric learning [SNC, MREE, PREE]?
28 Trials Trials Number of Errors Mammals (Rats) (c) Birds Pigeons (Pigeons) (b) Reptiles Turtles (Turtles) Trials S L-S ln Initial Latency Trial Blocks Mean Log Latency Trials S L L-S SNC (c) Pigeons 3.0 S L L-S (a) Amphibians Toads (Toads) Mean Log Latency Trials S L L-S ln Initial Latency Trial Blocks 2.5 S (b) Turtles 2.2 Reversed SNC S L L-S y S L L-S Mean Log Latency Bony Fish (Goldfish) L L-S Trial Blocks
29 Distribution of spaced-trial SNC and MREE in vertebrates Osteichthyes (Goldfish) Amphibia (Toads) Aves (Pigeons) Reptilia (Turtles) Mammalia (Rats) Allocentric + Egocentric learning [SNC] Allocentric learning (S-W learning mechanism) [Reversed SNC]
30 A tentative classification of learning phenomena Learning Phenomena General Specific Nonassociative Habituation Short term Long term? Sensitization Short term Long term? Associative/Cognitive Classical conditioning Instrumental conditioning Rule learning Social learning Human language Song learning Imprinting
Role of Reinforcement in Spaced-Trial Operant Learning in Pigeons (Columba livid)
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