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Learning and Memory 11/29/2009 •Learning – A long term change in behavior as a function of experiences. •Memory – The capacity to retain and retitrieve ppas t exper p iences. The Neurobiology of Learning and Memory Mary ET Boyle, Ph.D. Department of Cognitive Science UCSD 1 11/29/2009 Types of Learning Types of Learning: •Habituation Pavlovian Conditioning – A decrease in response following • Type of learning in repeated exposure to which a neutral a non-threatening stimulus is paired stilimulus. with a stimulus that elicits a reflex response until the • Sensitization neutral stimulus – An increase in elicits the reflex reactivity to a response by itself. stimulus following exposure to an intense event. 2 11/29/2009 Types of Learning: Types of Learning: Pavlovian Conditioning Operant Conditioning • Learning how to behave to obtain reinforcement 3 11/29/2009 Types of Learning: Operant Conditioning Reinforcers • activities that increase the frequency of the behavior PihPunishers • activities that decrease the frequency of the behavior 4 11/29/2009 Atkinson-Shiffrin model Baddeley’s rehearsal systems approach • Baddeley argued that memories go directly from the sensory register to long-term storage. November 2001 • American Psychologist 5 11/29/2009 Working Memory: A Modern Advance Dual task paradigm Baddeley Memorize arrangements of pieces on a chessboard while: Softly counting from 1 to 10 Phonological To tap a series of keys in a spatial pattern Visuospatial Produce a new random letter every second Decision-making Working memory—A system that permits the temporary storage and manipulation of information required for tasks such as comprehension, learning, and reasoning. 6 11/29/2009 Craik and Lockhart Dual task paradigm • The theory that memories differ in the extent to which they have been processed. Counting: no effect Tapping keys: impaired recall Producing letters: impaired recall November 2001 • American Psychologist 7 11/29/2009 Types of Memories: Endel Tulving 1972 • Episodic memory—The memory of temporally related events experienced at a particular time and place. • For example, “What were you doing on your birthday?” According to Tulving, animals • Semantic memory—The memory like his cat have no episodic of knowledge concerning the use memory so while they may of language and the rules, know many things, they do not remember past experiences the formulas, or algorithms for the wayyyj we do. They just know development of concepts or about them. solutions to problems. • For example, “How do you speak Japanese? Drawing by Ruth Tulving 8 11/29/2009 • Procedural memory (nondeclarative memory) Skill memory, or the memory o f highly practiced behavior. These memories may not be accessible to conscious awareness. 9 11/29/2009 • Declarative memory—Factual memory, or the memory of specific events. We are aware of declarative memories. 10 11/29/2009 The Memory Consolidation Process: Processes in LTM Hebb’s Cell Assemblies Encoding Cell assembly Processes used to store info rma tio n in me mo rry Reverberatory activity Storage Processes used to maintain Phase sequence information in memory Experience triggers Retrieval activity Processes used to get information back out of Activity is essential for LTM memory Physiological changes 11 11/29/2009 Is Reverberatory Activity Essential for Memory Storage? • Studies have used ECS (electroconvulsive shock) to break up the consolidation of memory. • CS reminders have brought back • ECS tends to produce retrograde memories apparently lost after amnesia supporting the idea that ECS indicating that reverberatory actiiivity is consolidation has occurred. necessary for memory consolidation. • Therefore, forms of brain intervention (e.g., ECS) affect •There are problems because ECS memory retrilhhieval rather than has aversive properties which consolidation. may inhibit responding. 12 11/29/2009 The Conditioning (Plasticity) of Neural Circuits • Thompson and associates – Studied conditioning of the nictitating membrane of the rabbit. – Used a corneal air puff ( UCS) and tone (CS) – Cerebellar interpositus nucleus was found to be site of the conditioned response 13 11/29/2009 How does the brain associate two events? Event 1 associate Event 2 Timing is the key!!! Neural Circuits in the Rabbits Nictitating Membrane Response 14 11/29/2009 two stimuli immediately stimulus separated in time follow one another (trace condition) Animals require many more Animals learn this association trials to learn the association easily. but eventually do it. 15 11/29/2009 Which structure is LPN: needed? Lateral Pontine Nucleus of the Cerebellum Hippocampus? 16 11/29/2009 Formation of trace The LPN is necessary for the memories requires the paired association. hippocampus!! • If the LPN was damaged – then pp(aired associations (CS-US))w would not be learned. • Remember: paired association means that there is no time bhbetween the presentat ifhCSion of the CS and the US. 17 11/29/2009 The Cellular Basis of Learning Sensitization in Aplysia & Memory californica • Kandel’s studies of learning in Aplysia californica (a shell-less marine mollusk) indicated both habituation and sensitization exist in this animal. 18 11/29/2009 Presynaptic Facilitation in • Normal classical conditioning occurs with a CS of a light touch to the Aplysia mantle or siphon paired with an electric shock to the tail. 19 11/29/2009 Structural Changes and Storing Experiences • Experience enhances Ca2+ ion entry into the hippocampus (Lynch, 1986). –Ca2+ ions activate the enzyme calpain which breaks down fodrin, a protein that coats the dendrites. This exposes more glutamate receptors to stimulation from other neurons, making the postsynaptic neuron more sensitive. – This may eventually cause changes in the terminal button and Lynch believes this may be the biological basis of learning and memory. 20 11/29/2009 The Anatomy of Learning & Memory The Anatomy of Learning and Memory • Squire and others have identified areas of the brain involved in declarative memory. – Medial temporal lobe—Hippocampus • Procedural or nondeclarative and surrounding cortical areas relay memory involves the neocortex information to diencephalic areas (mammillary nuclei, anterior thalamic and neostriatum. nuclei, and the mediodorsal thalamic • Classical conditioning of nucle i); it is p rocessed a nd re layed to reflexes depends on the the frontal lobe. cerebellum. – Frontal lobe—Medial temporal lobe structures and the medial thalamus jointly may be essential for the fiflformation of long-term memory; connections between these structures and the frontal lobe may provide a route by which memories can influence behavior. 21 11/29/2009 Key Brain Structures in Declarative Memory Storage and Retrieval 22 11/29/2009 The Case of HM Patient H.M. • DtthhiDamage to the hippocampus • Bilateral medial temporal • Bilateral medial temporal lobe lobectomy causes anterograde resection amnesia • PiPatient H HM.M. -surgery to treat severe epilepsy in the 1950’s – Normal intellectual ability – No personality change – CtlthiCannot learn anything new – Language was unimpaired, and reading and writing are still intact • However, HM’s language contains no words/phrases ldihiptilearned since his operation – For example, when asked to define “flower child”, he said it was a young person who grew flowers • STM unimpaired 23 11/29/2009 The Mirror The Case of HM: In His Tracing Words Task • “Every day is alone in itself, whatever enjoyment I’ve had, and whatever sorrow IIve’ve had… Right now, I’m wondering. Have I done or said anything amiss? You see, at this moment everyygthing looks clear to me, but what happened just before? That’s what worries me. It’s like waking from a dream; I just don’t remember” • Milner (1970, p37). 24 11/29/2009 Another famous patient The Importance of the Hippocampus • CliveWearing • Viral encephalitis—destroyed parts of his temporal lobe in his 40s • Lives totally within most recent 1-2 • Damage to hippocampus results minutes in memory deficits. – Remembers what just happened, forgets everything else – Deficits in learning a simple object – Inability to form new memories, discrimination constantly feels like he just woke up – Deficits in delayed nonmatching- • Describes his life as being “like death ” to-sample task • Musician, choral director – Case of R.B.—hippocampal – Wearing still recalls how to play the piano damage produced profound and conduct a choir – Has no recollection of having received a anterograde amnesia musical education – procedural memory is intact – http://www.youtube.com/watch?v=Vwigmktix2Y& feature=related 25 11/29/2009 Long-Term Potentiation The Importance of the in the Hippocampus Hippocampus • In the case of H.M., memories • Long-term acquired before surgery were potentiation—an retained suggesting that the increase in the hippocampus is involved in the amplitude and duration storage of declarative memory of EPSPs in response bibut is not t hihe site o f storage. tthtttilto the test stimulus. • Some researchers have found episodic encoding in the left frontal areas and episodic retrilihihflieval in the right frontal regions. 26 11/29/2009 Long-Term Potentiation in the Characteristics of LTP Hippocampus • Three pathways involved in LTP – Perforant fiber pathway – Mossy fiber pathway • A brief, sensitizing stimulus is sufficient to – Sc ha ffer co llatera l fiber pat hway prodLTPdttthtduce LTP; demonstrates that hippocampal neurons can change synaptic responsivity following a single event. • LTP-changed synaptic responsivity is confined to a specific
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