1. Neurobiology of Learning and Memory Stephan Anagnostaras

2. My view on learning and memory For this class: Field is broad and the breadth is important, so we will follow good examples to keep depth Several different fields cover learning and memory. My view is there should be one field and we would all be in the learning and memory building. Course will come from an eclectic perspective

3. Several modern disciplines study neurobiology of learning and memory Psychology Behavioral Neuroscience Cognitive Neuroscience/Neuropsychology Physiology Synaptic Plasticity Molecular Genetics reverse, forward, and epigenetic approaches Biology Neuroethology of birdsong learning

4. Learning and Memory Why learning and memory? Learning - current views on this term are from the field of learning theory/behavioral psychology. A relatively permanent change in behavior as the result of experience - BF Skinner Memory - current views from in information processing theory (cognitive psychology and computer science). The capacity to store and retrieve of information (comp sci) The faculty of the mind by which it retains the knowledge of previous thoughts, impressions, or events. (cognitive psych)

5. Learning versus Memory Historically there was deep animosity between behavioral & cognitive psychologists. Why? Problem relates to view of associationism. Behaviorism - the cornerstone is behavior because it is observable. Goal is to find lawful relationships between external events & behavior Cognitive psychology - a key tenet is the concept of an internal representation. This is how knowledge is stored, and operations can occur between representations. Can usually only be inferred or known through introspection.

6. Origins of the study of L&M The study of learning is closely related to the beginning of experimental psychology (around 1900) and, before that epistemology (philosophical study of how we have knowledge) Nature-Nurture Debate Renee Descartes (1641) Nativism - knowledge is innatively given - e.g., knowledge of God, perfection, infinity Rationalism - knowledge gained by reasoning logic and intuition. –In both instances, knowledge is independent of experience

7. Origins of the study of L&M Aristotle (350 BC) - 4 laws of memory 1. Similarity 2. Contrast 3. Contiguity 4. Frequency John Locke (1690) Empiricism (Associationism) - knowledge is gained by experience as provided to the mind by the senses. -these views advanced by others - esp David Hume in the 1700s -Enquiry concerning human understanding, 1748

8. Origins of the study of memory Ebbinghaus (1885) Used introspection to study forgetting in himself list of 12-16 consonant-vowel-consonant (CVC) nonsense syllabus (e.g., KEG, MIW) memorize the list by repeating until recalled, then record # of trials and wait 20 min - 31 d Relearn the list = SAVINGS Most forgetting in first 20 min, very little between 20 min and 31 d

9. Origins of the study of memory

10. Ebbinghaus also discovered serial position effect (primacy & recency effects) Glanzer, M. & Cunitz, A.R. (1966). Two storage mechanisms in free recall. Journal of Verbal Learning and Verbal Behavior, 5, 351-60

11. Serial position effect is ubiquitous

12. Origins of the study of memory William James (1890) argued there must be at least two types of memory Primary memory -information currently contained in consciousness Secondary memory -stored information which can be brought into consciousness First of many suggestions that there were multiple memory systems.

13. Origins of the study of memory The study of memory developed into information processing theory in the 1960s *Multi-store Memory System *Sensory Memory *Short Term Memory *Long Term Memory *Information Processing in Memory *Encoding *Storage *Retrieval

14. Origins of the study of learning Behaviorists tended to be somewhat hostile and engaged in many turf battles In the domain of clinical psychology they were battling it out with the psychoanalytic approach. Very much against introspection, tended to become extreme in this domain, and rejected cognition as well. In the domain of animal behavior, they were battling it out with the ethologists. Ethologists believed everything was innate and nothing was learned, hence behaviorists believed nothing was innate and everything learned.

15. Behaviorist Tradition Behaviorism focuses on objectively observable behavior and discounts unobservable mental activities. Behaviorists mostly studied animal and human learning focusing on observable behavior and ways to change behavior. Their studies of learning came to be known as learning theory and their studies of how to change behavior is known as behavior modification Largely relationships between external stimuli and events and their ability to influence behavior Radical behaviorism allows no intervening processes (e.g., BF Skinner) while other forms are more lenient.

16. Behaviorism John B Watson, the father of American Behaviorism – Began a scientific movement for Psychology, against several mystical squishy fields, esp. Freudian Psychoanalysis

17. Watson & Raynor's famous experiment with "Little Albert"

18. Watson & Rayner's (1920) famous experiment with "Little Albert"

19. Watson & Rayner's famous experiment with "Little Albert"

20. Behaviorism of the time was very anti- genetics, in part because of the evil mental testing movement Give me a dozen healthy infants, well- formed, and my own specified world to bring them up in and I'll guarantee to take any one at random and train him to become any type of specialist I might select--doctor, lawyer, artist, merchant- chief, and yes, even beggarman and thief, regardless of his talents, penchants, tendencies, abilities, vocations, and race of his ancestors - John Watson Concept of "Tabula rasa" - organisms are born as blank slates John Watson and Rosalie Rayner

21. Tabula rasa v. intelligence Notice our education system is still scarred by the nature-nurture debate “Social scientists” who think everything is nurture Mental testers who think everything is nature As a result, very little study of genetics of learning until 1992

22. Russian Reflexology Ivan Pavlov CSCR URUSCS CS-US pairing

23. Pavlov’s Little Albert CR

24. Learning and Performance Distinction between learning/memory and performance an important problem in animal and human studies Learning can only be inferred by performance: in humans, a response or a change in behavior in rats, a change in behavior Performance may not occur: Latent learning [Tolman & Honzik (1930)] Test anxiety Require extensive controls

25. Operant/Intrumental Conditioning Law of Effect (1911) “Of several responses made to the same situation, those which are accompanied by or closely followed by satisfaction to the animal will, all other things being equal, be more firmly connected with the situation, so that, when it recurs, they will be more likely to recur… discomfort… weakened. The greater the satisfaction or discomfort, the greater the strengthening or weakening of the bond.” Edward Thorndike

26. Operant/Intrumental Conditioning Response is paired with an outcome (appetitive or aversive), R-S* Response increases -Positive Reinforcement-app -Negative Reinforcement-avs Response decreases -Punishment -avs -Time-out/Diff Reinf Other -app

27. Behaviorism split into two camps Learning theory - theories relating environmental events and behavior could have a limited number of inferred processes & phenomena e.g., association, generalization, etc. Radical behaviorism - could only make laws between behavior and environmental events (only things that are observable)- typified by Skinner

28. Learning theory today Learning theory was more successful but ultimately self-destructed by dozens of wrong assumptions, most of which were excessively simplistic equipotentiality of stimuli single associative value indepedence of path invariance of path

29. Learning theory today Survives primarily as a subdiscipline of behavioral neuroscience Strong integration of principles and rigor into other fields of study

30. Animal Cognition Edward Tolman Cognitive Psychology emphasizes internal mental representations and operations between them, both it and computer science are referred to as information theory. Memory and knowledge are cognitive terms. Tolman emphasized flexibility of animal knowledge, in contrast to the very mechanistic views of most behaviorists Also rejected the idea of Tabula rasa

31. Tolman's path integration experiment

32. Tolman & Tryon: Selective breeding of maze bright and maze dull rats

34. Influence of Rearing Environment Cooper & Zubeck (1958) compare with Rosenzweig & Tryon (1950) Enrichment first done by D.O. Hebb (1949)

35. Behavioral Neuroscience & Neuropsychology Karl Lashley, father of Behavioral Neuroscience Laws of equipotentiality and mass action

36. Neuropsychology Studies the breakdown of function after brain damage in humans First original case of Lebourgne by Paul Broca (1861), Broca’s language production area (Wernicke-1894 - language reception) Strong push toward localization of function Early push was from Gall’s Organology (Spurzheim’s Phrenology) Read the critique by S.I. Franz

37. Neuropsychology Theodore Ribot (1882) -Reviewed cases of retrograde amnesia associated with brain damage --in most cases memory acquired remotely before the insult was preserved compared to that acquired recently --Ribot’s law of regression - loss of memory is inversely related to the time elapsed between the event to be remembered and the injury. Ribot concluded memories need a certain amount of time to become organized and fixed.

38. Neuropsychology Alois Alzheimer (1906) Reported an institutionalized female patient with progessive dementia. After being shown objects and recognizing them, she immediately forgot them and circumstances under which she learned them. Both anterograde and retrograde amnesia were present, and obeyed Ribot’s law as well. Sergei Korsokoff (1887) Memory impairment in chronic Alcoholics which obeyed Ribot’s law.

39. Neuropsychology Georg Muller & Alfons Pilzecker (1900) - large number of experiments in normal subjects -same as Ebbinghaus, but nonsense syllables in pairs -Recall second item when probed with first from pairs -Spontaneous recall of pairs from the same list (perseveration) which had a time-gradient of a few minutes -- speculated reflected transient brain activity -Put distractor lists between training and recall, get retroactive interference -- this follows a time gradient so that distractors are only effective for a few minutes --Concluded brain activity perseverates after new learning and activity serves to consolidate memory.

40. Neuropsychology Several interesting case studies since then, but the most influential was not until the 1950s. Phenomenon of consolidation not studied a whole lot by animal learning people until recently Phases of memory are a recurring theme which we will struggle to understand throughout the class.

41. Where are we today? Further than any other area of neuroscience - not very far Strong stratification of the study of learning and memory hinder vertical integration Try to build an interdisciplinary framework from molecules to cognition No single engram identified at the molecular- genetic level