Background of the study of Learning and memory.
A. Quantification of behavior as it applies to the study of L& M.
the road to the rigorous study of learning and memory was paved by two important people: 1) in the late 1800's/ early 1900's objective measures of mental activity were introduced by Ebbinghaus and 2) the emergence of experimental animal psychology (Darwin) helped pave the road for study of learning and memory processes. Others that have contributed substantially to the development of quantitative methods for studying learning and memory include: **Sechenov, Ivan/Pavlov, Ivan, two Russian physiologists- they developed paradigms where they could restrict an animal or humans behavior, control stimuli and limit responses. Using this simplified paradigm they were able to quantify the behavior and make predictions about what temporal and spatial requirements of stimuli presentation were necessary for the learning process to occur. Their techniques are referred to as classical conditioning techniques and involve the association of an unconditioned stimulus with a conditioned stimulus.
**Thorndike, Edward Lee/B.F Skinner - ibid. instrumental conditioning, operant conditioning. Association of a particular response with a particular consequence.

What are some of the traditional questions and answers that have been provided in the past.
All of the questions Dudai askes on pgs 8-10 fall into these two catagories .
B. Where is memory stored? Physiological-Does not tell us about how memory is stored but can tell us something about what areas or the brain are responsible for various cognitive functions. Thus, from an analysis of the nature of a patients mental disorder we can pinpoint the area that is responsible and begin some sort of remedial treatment.
The earliest record of this question having been addressed comes from the 4th century BC where Aristotle held that the heart was the seat of mental function. While this view held for some time mostly on the reputation of Aristoltle himself it was fundumentally wrong and was challanged first by Alexandrian anatomists who suggested that mental function might be localized in the brain and then by the physician Galen (2nd century AD). He came to this conclusion throught the discection and observation of animals. Galen seems to be the first person to demonstrate that nerves originate in the brain and that motor and sensory function are abolished by brain injuries. **While Galen was correct about the general location he was very mistaken about the specifics. He believed that nutrients from food were taken to the liver and converted to 'animal spirits'. These spirits were then transported throughout the body along the veneous system. The arteries on the otherhand transported 'vital spirits' formed in the left ventricle. These vital spirits were converted to 'animal spirits' in the rete mirabile which was located at the base of the brain and stored in the ventricles. These animal spirits were stored in the ventricles and then transported throughout the body through nerves ( which were assumed to be hollow) when needed.
Medieval scholars attributed even more emphasis to the role of the ventricles. St Augustus writes that the ventricles were, not only the place where 'animal spirits' were stored but also the seat of all mental function. Furthermore, each ventricle was assigned a particular function. The lateral ventricle was called the 1st cell and it **stored sensory imput and therefore was in charge of perception. The third ventricle was called the 2nd cell and it reflected on info recieved from the 1st cell and therefore was in charge of thinking and reason. The fourth ventricle was called the 3rd cell and served the function of memory. In Dudai it is suggested that the idea of memories being stored in the ventricles was first proposed by Costa ben Luca 9th century and Albert Magnus 13th century. While it may be true that they made the prediction, as you've heard these midevil scholars were simply rehashing an old idea. Renaissance anatomoists discovered the errors in the accepted dogma of the time. The demonstrated that there was no rete mirabile in primates (Vesalius). They also showed that the ventricles were simply filled with liquid and were too comple to serve the suggested function (Leonardo Da Vince - cast of cow ventricles).
In the early 1600s Descartes suggested that human memory was subserved by physical traces in the brain that were produced by activity-dependent facilitation of connections in these traces (pg 11 Dudai). He did not however suggest where exactly these traces resided and he was still incorporating the idea of fluid in the ventricles being involved in the process. IN the 17th century William Harvey's demonstration of how blood flows and is separate from other fluds helped dispell the belief that ventricular fluidds were made of converted vital spirits. Furthermore, advances in understanding that nerves functioned by electrical means helped to clear up the role of nerves in controlling movement through hydrolic means (carrying animal spirits to the muscle) - Luigi Galvani, Allesandro Volta.
Support for the storage of memories in the gray matter came from reports by various surgeons in the late 1800's. Theodule Ribot in France (1882), Sergei Korsakoff in Russia (1887) and J. Hughlings Jackson (1888) were some of those who reported that damage to the hippocampal region was associated with profound deficits in the ability to learn.
At the same time much information on the microscopic aspects of the nervous system were being published. Golgi demonstrated the the nervous system was like a net. From his histological studies he suggested that the brain was made up of a large continuous web (retularis school). Using slightly modified techniques Ramon y Cajal provided evidence to suggest that the brain was not one big continuous mass of threadlike tenbdrils but that it was made up of smaller discrete cells just like the rest of the body. These discrete cells were connected to each other by dense contacts. Latter Sir Charles Sherrington would name these contacts 'synapses' **. In the mid 1900's Sir henry Dale and Otto Loewi would show that transmission across synapses is chemical.
Sept 8
Also in the mid 1900's Penfield's (1948 ) work with epileptic patients demonstrated that stimulation of certain areas of the brain could evoke specific memories. These recollections were produced by stimulation of areas near those which the neurosurgeons had shown to be important for learning.
**Karl Spencer Lashley (1950 ) added to the debate by showing that rats who learned a complex maze were unimpaired in their ability to renegotiate the maze even after large areas of the brain were removed. Thus he argued against Penfield's claims of memory localization at least of complex learning (not simple learning which could be disrupted by localized lesions in the sensory cortices. He argued forsomething called equipotentiality and mass action. Both concepts developed to explain the fact that the size of the cortical lesion was more disruptive than the position of the lesion.

More recent work, that we will review, pinpoints various structures as being important. And shows that the importance of that structure is very specific to the task and question that is being asked. Furthermore, we will reconcile the claims of equipotentiality and localization reported by Lashley and Penfield.

C. How do we form memories? Cognitive
Descarte
Sechenov/Pavlov (1862)- specific principles temporal and spatial relationships between stimulus and responses are necessary before learned associations can occur. Some of the most prominent early ideas come from philosophers. eg Hobbes - associationist
1900 French psychologist Alfred Binet exposed children to various objects and then tested their memories in the presence or absence of misleading questions. Binet found that misleading questions produced systematic distortions in children's recollections. Thus, suggesting that retrieved memory is not necessarily an imprint of the input but either 1 )distorted during the storage process, 2) distorted during the retrieval process or 3) both. In any case the retrieved memory is not an exact replica of reality. Suggesting that some reconstruction is occurring.
Ramon Y Cajal (1900) demonstrated that the brain was made of discrete entities he called neurons that seemed to touch each other. Based on this finding he suggested that changes in those areas of contact would be responsible for the storage of information i.e. memories.
Thorndike/Skinner - (1900) ibid
Janet (1928) suggested that new experiences are encoded with respect to, and incorporated into, preexisting knowledge structures accordingly, individual experience may be very difficult to retrieve later as separate, distinct entities (Janet - French psychologist)
Bartlett (1932) stated, based on changes in the retelling of a story, that memories are imaginative reconstruction of past events that are heavily influenced by the rememberer's prexisting knowledge structures of schemas. This view was overlooked in favor of the stimulus-response paradigms and the hypothesis that those paradigms produced (see pavlov, thorndike)
Hebb (1949)- recapitulated Descartes + Ramon Y Cajal's ideas by saying that a trace is layed down through repeated activation of that trace but he extended this by saying that those traces are formed of distributed assemblies of interconnected neurons thus incorporating the ideas of Lashley.

Seriel processing of information was simulated in computers but found lacking because these programs were unable to even complete a rather simple task that is done by humans without even thinking. However, the recent development of techniques for implimenting parallel distributed processes on computers have suggested that this may be an important feature of how memories are stored. This finding lends credence to the Karl Lashleys ideas of equipotentiality and mass action.
Also,Bartlett's original idea of constructivism or memory reconstruction are being revisited due to recent evidence and theories that focus on parallel distributed networks.
Memories are formed in the context of all the information that is available both externally and internally. When memories are formed in isolation we have a deficiet referred to as :source amnesia". Again suggesting that memory is encoded with respect to prior knowledge and other events. However, sometimes distortions or what is recalled can be very insightful with respect to how information is stored. Specific errors may be similar in sound or in context to the word that was to be remembered. This suggests that when testing what was learned we must be careful to analyze even mistakes or else we are in danger of missing important information about the way memories are stored. This is the whole point behind the memory distortion book.
In reading the texts and listening to lectures you need to keep in mind that this course, although limited in scope to learning and memory could just as easily be called animal behavior or intelligence.

In order to forewarn you. I would like to point our a few areas and questions that I will not be addressing.
Social or collective memory
Forgetting except in passing
Attention or focusing

Summary:
Definition of learning
Levels of analysis

  1. Top-down/Bottom up approaches
    1. How is memory formed at a systems level? Schacter
    2. How is memory formed at a molecular level?Dudai
  2. 2 overriding questions and answers provided by earlier work: timelines for historical figures
    1. Where is memory stored?
    2. How do we form memories?
In the following weeks we will look at data that addresses the two overriding questions mentioned above in an attempt to come to some conclusion about the learning process.