Historical Perspectives (A century’s research on the neurobiology of learning and memory)

1. Historical Perspectives(A century’s research on the neurobiology of learning and memory) READING: Rosenzweig, M. R. (1998). Historical perspectives on the development of the biology of learning and memory. In J. Martinez & R. Kesner (Eds.), Neurobiology of learning and memory (pp. 1-53). San Diego, CA: Academic Press. On reserve in Health Science library. This item is not in the textbook or courseware.

2. General framework (overview) • Top-down approach: • Describing learning and memory in whole organisms and identifying the associated neural regions and systems. • Multiple memory systems: multiple functionally and anatomically distinct brain systems. • Bottom-up approach: • Describing neuronal and synaptic mechanisms and neural networks that may underlie learning and memory

3. Historical Perspectives Part 1, Pre-1960s period

4. 1. The origin of experimental psychology: • the first research psychologies were Germans • a. Fechner & Weber: psychophysics • b. Wilhelm Wundt, 1879, • opened a formal laboratory of experimental psychology at U of Leipzig, • founder of psychology as a science • c. Hermann Ebbinghaus 1850-1909, performed dozens of experiments • i. Study-test method: memorized lists of nonsense syllables and tested himself • ii. excluded the influence of extraneous factors on the outcome of experiments (he realized that a major contaminating factor could be the prior knowledge that subjects bring to an experiment). • iii. novel and successful attempt to test memory experimentally

5. 2. William James 1890, Principles of Psychology, separate chapters to habit, association and memory, Harvard lab (teaching) a. Habit, memory, and other aspects of behaviour are based on physiological properties of the brain, even though not be able to specify b. Separate chapters on habit and memory, a precursor to nondeclarative and declarative memory c. The idea that conscious experience requires memory

6. 2. William James (continued) d. Memory systems include i. consciousness primary memory, ii. secondary memory, which comprises our permanent record of the past. iii. later came to be called short term and long term memory e. His ideas about memory did not start to influence experimental psychology until the 1960s. This extraordinary delay can be attributed to the direct influence of behaviourism.

7. 3. Edward Thorndike (1874-1949), one of the first psychologists to perform systematic experiments on the learning process a. Extended the research to animals b. Instrumental conditioning (operant conditioning),

8. 4. Pavlov, the turn of 19 century, 1904 Nobel price, a. The concept of conditioning from observations on salivary responses b. the first one to demonstrate that salivation could be evoked by a previously neutral stimulus after this has been paired with an effective stimulus.

9. 5. Tolman and cognitive maps, the challenge to behaviorism a. classic paper 1948 “cognitive maps in rats and men” b. Tolman developed operational definitions for mentalistic processes including purposive behavior and expectancy. c. identify the complex cognitive mechanisms, purpose, expectations, and insights that guided behavior d. stated learning generally involved the acquisition of knowledge about the world

12. Neural substrates: 6. “Search for the engram” engram is the term to describe the memory trace that is presumably present in the brain after something has been learned -- debate over whether specific engrams are localized in discrete areas of the brain or interwoven throughout the entire brain

13. 7. Karl Lashley (1929) tried in vain to provide evidence that specific cortical connections are formed during conditioning procedures. a. He found no evidence that destruction or removal of any specific area of the cerebral cortex could either erase the engram or cause learning to be prevented from occurring.

14. 7. Karl Lashley (1929) continued b. His results suggested that the amount of cortical tissue destroyed was far more important than the location of the damage. c. The results forced him to reject the localizationist position, instead, he came to champion the view that specific memories were diffusely distributed throughout the cerebral cortex and that all parts of the cortex were equally involved in learning.

15. 7. Karl Lashley (1929) continued d. “This series of experiments has yielded a good bit of information about what and where the memory trace is not. It has discovered nothing directly on the real nature of the engram. I sometimes feel, in reviewing the evidence on the localization of the memory trace, that the necessary conclusion is that learning is just not possible.” -- Lashley, 1950

16. Historical Perspectives Part 2, Multiple memory systems (Neuropsychology and cognitive psychology) 1960s -

17. 1. After the promising beginning, this ideas disappeared during first half of the century 2. 1960s, 1970s, converging evidence from cognitive neuropsychology, psychology, neurobiology support the view of Multiple memory systems, efforts to experimentally dissociate memory systems emerged as a central research direction

18. 3. Famous amnesic patient H.M. • “frankly experimental” brain surgery • a complete bilateral resection of medial temporal lobes • including hippocampus and amygdala • for relief of intractable epilepsy. • HM’s extremely poor ability on tests requiring the retention of verbal • information over longer periods of time

19. b. Psychological testing • Probe digit task (Waugh and Norman, 1965). Subject listened to a sequence of 16 digits which was then followed by a probe digit. The subject’s task was to name the digit that occurred after the probe. • Example: 5824972537196435 6 (probe)

21. RESULTS • as the number of items occurring after the probe digit increased, • performance on the probe task became poorer, declining at a steeper rate with five or fewer intervening items. • The change in the gradient was taken as indicating the transition from primary to secondary memory, and its occurrence at the five-item point provided the first empirical demonstration of separate primary and secondary memories and the first estimate of primary memory capacity.

22. Probe digit task test of HM (Wicklegen, 1968) Easier version of the probe digit task, showed normal performance Digit span task measures the ability to repeat a random series of numbers in the correct order immediately after seeing or hearing them HM also showed normal primary memory, 7 plus or minus2

23. HM’s extremely poor ability on tests requiring the retention of verbal information over longer periods of time Conclusion: primary memory exists independently of secondary memory

24. Mirror-tracing task (Brenda Miller, 1965) HM did not show deficit on motor learning

25. Declarative vs. Procedural memory • a. HM’s memory problem could not be attributed solely to difficulties with verbal materials • i. he had difficult reproducing or recognizing pictures and spatial designs • ii. could learn some kinds of information about the verbal materials • e.g. could learn to read works printed mirror-reversed (requiring the ability to deal with abstract rules or procedures) • b. the important distinction is probably not between verbal and motor performance, rather between declarative and procedural memory

26. Declarative memory i. Concept (1) facts and information acquired through learning (2) memory that we are aware of accessing ii. Includes: (1) episodic memory: autobiographical, personal history (2) semantic memory: generalized memory, e.g. knowing the meaning of the words

27. Nondeclarative memory • (procedural memory) • shown by performance rather than by conscious recollection. • (2) includes: • (a) skill learning • (b) priming, a change in the processing of a stimulus as a result of prior exposure • (c) conditioning

28. Cognitive psychologists: Tulving’s contribution on episodic memory and semantic memory, Toronto school, the work of theorist: Craik, Schacter, Tulving

29. Logic underlying the use of the evidence from neuropsychology (rationale for cognitive neuropsychology) K. Craik (1943) writes: ‘In any well-made machine one is ignorant of the working of most of the parts - the better they work, the less we are conscious of them... it is only a fault which draws attention to the existence of a mechanism at all.’

30. The concept of modular system • Cognitive neuropsychology assumes that the various components of cognition, including memory, are spatially distributed within the brain and that, furthermore, each of these mental functions is modular. • ii. A modular system is one in which several components interact to perform a function, but each component, or module, is functionally autonomous - i.e. it can continue to operate if other modules cease to work for any reason. • iii. When the brain is damaged, either by accident or by illness, it is assumed that modules may become dissociated from each other in a meaningful way and thereby reveal something about the underlying organization of the system.

31. one-way dissociation or double dissociation