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Cognitive Neuroscience LEARNING AND MEMORY 509 Cognitive PART Neuroscience © 2010 Sinauer Associates, Inc. This material cannot be copied, VI reproduced, manufactured, or disseminated in any form without express written permission from the publisher. CHAPTER 17 Learning and Memory 511 CHAPTER 18 Attention and Higher Cognition 549 CHAPTER 19 Language and Hemispheric Asymmetry 583 © 2010 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured, or disseminated in any form without express written permission from the publisher. © 2010 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured, or disseminated in any form without express written permission from the publisher. Previous page Simultanenous registration maps of EEG and functional MRI In this technique the excellent spatial resolution of fMRI and the excellent temporal resolution of EEG are combined to provide an improved method for tracking brain activation associated with complex cognitive processing. (Image by Jan C. De Munck V, VU University Medical Center, Amsterdam.) © 2010 Sinauer Associates, Inc. This material cannot be copied, LEARNING AND MEMORY 511 reproduced, manufactured, or disseminated in any form without express written permission from the publisher. CHAPTER Learning and Memory 17 Trapped in the Eternal Now Henry Molaison, known to the world as “patient H.M.” in a classic series of re- search articles, was probably the most famous research subject in the history of brain science. Henry started to suffer seizures during adolescence, and by his late 20s, in 1953, Henry’s epilepsy was out of control. Because tests showed that his sei- zures began in both temporal lobes, a neurosurgeon removed most of the anterior temporal lobe, including much of the amygdala and hippocampus, on both sides. After Henry recovered from the operation, his seizures were milder, and they could be controlled by medication. But this relief came at a terrible, unforeseen price: Henry had lost the ability to form new memories (Scoville and Milner, 1957). For more than 50 years after the surgery, until his death in 2008, Henry could retain Functional Perspectives on any new fact only briefl y; as soon as he was distracted, the newly acquired infor- Memory 511 mation vanished. Long after the surgery, he didn’t know his age or the current date, and he didn’t know that his parents (with whom he lived well into adult- There Are Several Kinds of Memory and Learning 512 hood) had died years previously. His IQ remained a little above average (Corkin et al., 1997) because most IQ tests monitor problem solving that doesn’t require Memory Has Temporal Stages: Short, remembering new facts for more than a few minutes. Henry knew that something Intermediate, and Long 517 was wrong with him, because he had no memories from the years since his sur- gery, or even memories from earlier the same day. Successive Processes Capture, Store, and Retrieve Information in the Every day is alone in itself, whatever enjoyment I’ve had, and whatever sorrow I’ve Brain 519 had. … Right now, I’m wondering, have I done or said anything amiss? You see, at this moment everything looks clear to me, but what happened just before? That’s Different Brain Regions Process what worries me. It’s like waking from a dream. I just don’t remember. (B. Milner, Different Aspects of Memory 523 1970, p. 37) Neural Mechanisms of Henry’s inability to form new memories meant that he couldn’t construct a lasting Memory 529 relationship with anybody new. No matter what experiences he might share with Memory Storage Requires Neuronal someone he met, Henry would have to start the acquaintance anew the following Remodeling 529 day, because he would have no recollection of ever meeting the person before. What happened to Henry, and what does his experience teach us about learning Invertebrate Nervous Systems Show and memory? Plasticity 533 Synaptic Plasticity Can Be Measured in ll the distinctively human aspects of our behavior are learned: the languages Simple Hippocampal Circuits 534 we speak, how we dress, the foods we eat and how we eat them, our skills Aand the ways we reach our goals. So much of our own individuality depends Some Simple Learning Relies on on learning, the process of acquiring new information; and memory, the ability to Circuits in the Mammalian store and retrieve that information. Cerebellum 540 In the Adult Brain, Newly Born Functional Perspectives on Memory Neurons May Aid Learning 542 Conditions that impair memory are particularly frightening; they can make it im- Learning and Memory Change as possible to take part in normal social life and can rob us of our identity. We can We Age 543 discover a great deal about learning and memory by examining how they fail. © 2010 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured, or disseminated in any form without express written permission from the publisher. 512 CHAPTER 17 learning The process of acquiring new and Clinical case studies show that memory can fail in very different ways, indicating relatively enduring information, be havior pat- that there are different forms of learning and memory, and that multiple brain re- terns, or abilities, characterized by modifi ca- gions are involved. The clinical research has provided guidance for further studies tions of behavior as a result of practice, study, employing animal models and brain-imaging technology; together, these diverse or experience. approaches are generating a comprehensive picture of the brain’s mechanisms of memory 1. The ability to retain infor ma- learning and memory. tion, based on the mental process of learning or encoding, retention across some interval of time, and retrieval or reactivation of the There Are Several Kinds of Memory and Learning memory. 2. The specifi c information that is stored in the brain. The terms learning and memory are so often paired that it sometimes seems as if one patient H.M. A patient who, because of necessarily implies the other. We cannot be sure that learning has occurred unless damage to medial temporal lobe structures, a memory can be elicited later. Many kinds of brain damage, caused by disease was unable to encode new declarative memo- or accident, impair learning and memory, and specifi c cases continue to provide ries. Upon his death we learned his name was powerful lessons. We’ll start by looking at some of these cases before progressing Henry Molaison. to newer brain-imaging techniques that provide novel information about brain amnesia Severe impairment of memory. regions involved in learning and memory. Study of different types of memory retrograde amnesia Diffi culty in retriev- impairment has revealed different classes of learning and memory, which we will ing memories formed before the onset of discuss later in the chapter. First, let’s look at a few cases of memory impairment amnesia. that have posed puzzles and have generated a great deal of controversy. anterograde amnesia The inability to form new memories beginning with the onset of a For patient H.M., the present vanished into oblivion disorder. Patient H.M.—Henry Molaison, whom we met at the start of the chapter—suffered from amnesia (Greek for “forgetfulness”), a severe impairment of memory. In Hen- ry’s case, most old memories remained intact, but he had diffi culty recollecting any events after his surgery. Loss of memories formed prior to an event (such as surgery or trauma) is called retrograde amnesia (from the Latin retro-, “backward,” and gradi, “to go”), and is not uncommon. What was striking about Henry was a far more unusual symptom: his apparent inability to retain new material for more than a brief period. The inability to form new memories after an event is called anterograde amnesia (the Latin antero- means “forward”). Over the very short term, Henry’s memory was normal. If given a series of six or seven digits, he could immediately repeat the list back without error. But if he was given a list of words to a b 1 cm study and then tested on them after other tasks had intervened, he could not repeat the list or even recall that there was a list. So Henry’s case provided clear neuroanatomical evidence that short- term memory differs from long-term memory—a distinction long Normal H.M. recognized by biological psychologists on behavioral grounds (a) (W. James, 1890). We will discuss long- and short-term memory in more depth later in this chapter. Henry’s surgery removed the amygdala, most of the hip- pocampus, and some surrounding cortex from both temporal H lobes (Figure 17.1). The memory defi cit seemed to be caused by loss of the hippocampus, because other surgical patients who had EC received the same type of damage to the amygdala, but less dam- age to the hippocampus, did not exhibit memory impairment. (b) But to the puzzlement of researchers, it soon became evident that experimental bilateral hippocampal lesions in laboratory animals 17.1 Brain Tissue Removed from Henry Molaison (Patient H H.M.) (a) MRI scans of a normal subject (left) and Henry (right) show that Henry’s hippocampus (H) and entorhinal cortex (EC) were extensively PH damaged, bilaterally. (b) Henry had a small amount of posterior hip- pocampus left (see drawing for orientation), but the parahippocampal Cer cortex (PH) was totally gone, and the cerebellum (Cer) was dramatically shrunken. (From Corkin et al., 1997; courtesy of Suzanne Corkin.) © 2010 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured, or disseminated in any form without express written permission from the publisher. LEARNING AND MEMORY 513 seemed not to produce widespread memory defi cits (Isaacson, (a) The mirror-tracing task 1972). What might account for this discrepancy between humans and lab animals? An interesting early fi nding suggested that Henry’s memory defi cit might involve verbal function.
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