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Chapter 13 Learning and Memory: Basic Mechanisms This multimedia product and its contents are protected under copyright law. The following are prohibited by law: • any public performance or display, including transmission of any image over a network; •preparation of any derivative work, including the extraction, in whole or in part, of any images; •any rental, lease, or lending of the program. 1 Copyright © 2004 Allyn and Bacon The Nature of Learning Learning: • The process by which experiences change our nervous system and hence our behaviors. Perceptual learning: • Learning to recognize a particular stimulus. 2 Copyright © 2004 Allyn and Bacon The Nature of Learning Classical conditioning: • A learning procedure; when a stimulus that initially produces no particular response is followed several times by an unconditioned stimulus that produces a defensive or appetitive response. 3 Copyright © 2004 Allyn and Bacon 4 Copyright © 2004 Allyn and Bacon The Nature of Learning Hebb rule: • The hypothesis proposed by Donald Hebb that the cellular basis of learning involves strengthening of a synapse that is repeatedly active when the postsynaptic neuron fires. 5 Copyright © 2004 Allyn and Bacon The Nature of Learning Instrumental learning: • A learning procedure whereby the effects of a particular behavior in a particular situation increase (reinforce) or decrease (punish) the probability of the behavior; also called operant conditioning. 6 Copyright © 2004 Allyn and Bacon 7 Copyright © 2004 Allyn and Bacon The Nature of Learning Reinforcing stimulus: • An appetitive stimulus that follows a particular behavior and thus makes the behavior become more frequent. Punishing stimulus: • An aversive stimulus that follows a particular behavior and thus makes the behavior become less frequent. Motor learning: • Learning to make a new response. 8 Copyright © 2004 Allyn and Bacon 9 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Long-term potentiation: • A long-term increase in the excitability of a neuron to a particular synaptic input caused by repeated high-frequency activity of that input. 10 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Hippocampal formation: • A forebrain structure of the temporal lobe, constituting an important part of the limbic system; includes the hippocampus proper (Ammon’s horn), dentate gyrus, and subiculum. 11 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Entorhinal cortex: • A region of the limbic cortex that provides the major source of input to the hippocampal formation. Granule cell: • A small, granular cell; those found in the dentate gyrus send axons to the field CA3 of the hippocampus. 12 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Dentate gyrus: • Part of the hippocampal formation; receives inputs from the entorhinal cortex and projects to the CA3 field of the hippocampus. Perforant path: • The system of axons that travel from cells in the entorhinal cortex to the dentate gyrus of the hippocampal formation. 13 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Field CA3: • Part of the hippocampus; receives inputs from the dentate gyrus and projects to field CA3. Pyramidal cell: • A category of large neurons with a pyramidal shape; found in the cerebral cortex and Ammon’s horn of the hippocampal formation. 14 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Induction of Long-Term Potentiation Field CA1: • Part of the hippocampus; receives inputs from field CA3 and projects out of the hippocampal formation via the subiculum. Population EPSP: • An evoked potential that represents the EPSPs of a population of neurons. 15 Copyright © 2004 Allyn and Bacon 16 Copyright © 2004 Allyn and Bacon 17 Copyright © 2004 Allyn and Bacon 18 Copyright © 2004 Allyn and Bacon 19 Copyright © 2004 Allyn and Bacon 20 Copyright © 2004 Allyn and Bacon 21 Copyright © 2004 Allyn and Bacon 22 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Role of NMDA Receptors Associative long-term potentiation: • A long-term potentiation in which concurrent stimulation of weak and strong synapses to a given neuron strengthens the weak one. NMDA receptor: • A specialized ionotropic glutamate receptor that controls a calcium channel that is normally blocked by Mg2 ions; involved in long-term potentiation. 23 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Role of NMDA Receptors AP5: • 2-Amino-5-phosphonopentanoate; a drug that blocks NMDA receptors. AMPA receptor: • An ionotropic glutamate receptor that controls a sodium channel; when it’s open, it produces EPSPs. 24 Copyright © 2004 Allyn and Bacon 25 Copyright © 2004 Allyn and Bacon 26 Copyright © 2004 Allyn and Bacon 27 Copyright © 2004 Allyn and Bacon 28 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Role of NMDA Receptors Dendritic spike: • An action potential that occurs in the dendrite of some types of pyramidal cells. 29 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Mechanisms of Synaptic Plasticity CaM-KII: • Type II calcium-calmodulin kinase, an enzyme that must be activated by calcium; may play a role in the establishment of long-term potentiation. Nitric oxide synthase: • An enzyme responsible for the production of nitric oxide. 30 Copyright © 2004 Allyn and Bacon Learning and Synaptic Plasticity Long-term Depression Long-term depression: • A long-term decrease in the excitability of a neuron to a particular synaptic input caused by stimulation of the terminal button while the postsynaptic membrane is hyperpolarized. 31 Copyright © 2004 Allyn and Bacon 32 Copyright © 2004 Allyn and Bacon Perceptual Learning Perceptual Short-term Memory Short-term memory: • Memory for a stimulus that has just been perceived. Delayed matching-to-sample task: • A task that requires the subject to indicate which of several stimuli has just been perceived. Paired-associate task • A task that requires the subject to learn to recognize pairs of stimuli. 33 Copyright © 2004 Allyn and Bacon 34 Copyright © 2004 Allyn and Bacon 35 Copyright © 2004 Allyn and Bacon Instrumental Conditioning and Motor Learning Reinforcement Neural Circuits Involved in Reinforcement Medial forebrain bundle (MFB): • A fiber bundle that runs in a rostral-caudal direction through the basal forebrain and lateral hypothalamus; electrical stimulation of these axons is reinforcing. 36 Copyright © 2004 Allyn and Bacon Instrumental Conditioning and Motor Learning Reinforcement Neural Circuits Involved in Reinforcement Ventral tegmental area (VTA): • A group of dopaminergic neurons in the ventral midbrain whose axons form the mesolimbic and mesocortical system; plays a critical role in reinforcement. 37 Copyright © 2004 Allyn and Bacon Instrumental Conditioning and Motor Learning Reinforcement Neural Circuits Involved in Reinforcement Nucleus accumbens: • A nucleus of the basal forebrain near the septum; receives dopamine-secreting terminal buttons from neurons of the ventral tegmental area and is thought to be involved in reinforcement and attention. 38 Copyright © 2004 Allyn and Bacon Instrumental Conditioning and Motor Learning Reinforcement Detecting Reinforcing Stimuli Conditioned reinforcer: • A previously neutral stimulus that has been paired with an appetitive stimulus, which then itself becomes capable of reinforcing a response. 39 Copyright © 2004 Allyn and Bacon Instrumental Conditioning and Motor Learning Reinforcement Detecting Reinforcing Stimuli Conditioned punisher: • A previously neutral stimulus that has been followed by an aversive stimulus, which then itself becomes capable of punishing a response. 40 Copyright © 2004 Allyn and Bacon 41 Copyright © 2004 Allyn and Bacon