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• To describe how fMRI measures brain activity and characterize this method's spatial resolution and temporal resolution.
• To describe how ERPs measure brain activity and characterize this method's spatial resolution and temporal resolution.
• To list one problem with patient lesion evidence.
• To describe how TMS works and characterize this method's spatial resolution and temporal resolution.
• To name two methods that could be combined to measure brain activity with excellent spatial resolution and excellent temporal resolution.
Cognitive neuroscientists employ tools to look inside the brain of participants while they are actively engaged in a mental process. This is no simple feat, and the field of cognitive neuroscience has grown with the advent of techniques that can measure activity in the functioning human brain. These methods vary in popularity, cost, complexity, spatial resolution, and temporal resolution. Each technique has advantages and disadvantages and takes years to master. This chapter briefly describes the most widely used techniques in cognitive neuroscience that will be referred to throughout this book. Section 2.1 briefly reviews the behavioral measures that allow for the interpretation of brain activation results. Section 2.2 discusses techniques with high spatial resolution, such as fMRI, which is the most popular method. fMRI measures the increases in blood flow that occur in active brain regions. This technique has excellent spatial resolution but has poor temporal resolution because the blood flow response is slow. Section 2.3 focuses on techniques with high temporal resolution, such as event-related potentials (ERPs). ERPs measure voltages (i.e., potentials) on the scalp that directly reflect the underlying brain activity. This technique has excellent temporal resolution and limited spatial resolution. In section 2.4, techniques with excellent spatial resolution and excellent temporal resolution are described. These include combined fMRI and ERPs as well as depth electrode recording from patients who have electrodes implanted in their brains for clinical reasons. Section 2.5 considers evidence from patients with brain lesions and cortical deactivation methods such as transcranial magnetic stimulation (TMS). Both of these methods have limited spatial resolution and poor temporal resolution; however, they can assess whether a brain region is necessary for a given cognitive process.
Within the last two decades, the field of cognitive neuroscience has begun to thrive, with technological advances that non-invasively measure human brain activity. This is the first book to provide a comprehensive and up-to-date treatment on the cognitive neuroscience of memory. Topics include cognitive neuroscience techniques and human brain mechanisms underlying long-term memory success, long-term memory failure, working memory, implicit memory, and memory and disease. Cognitive Neuroscience of Memory highlights both spatial and temporal aspects of the functioning human brain during memory. Each chapter is written in an accessible style and includes background information and many figures. In his analysis, Scott D. Slotnick questions popular views, rather than simply assuming they are correct. In this way, science is depicted as open to question, evolving, and exciting.
• To describe the behavioral effects and brain effects that typically occur during implicit memory.
• To identify the brain regions associated with implicit memory.
• To characterize the brain activity frequency bands associated with implicit memory.
• To detail the different neural models of implicit memory.
• To determine whether there is convincing evidence that implicit memory is associated with the hippocampus.
• To describe two different patterns of brain activity that occur during skill learning.
In everyday life, the term memory is used to refer to the conscious experience of a previous event. However, when an event is repeated, there can also be behavioral effects and brain effects that occur outside of conscious experience. Implicit memory refers to a lack of conscious experience or awareness of previously learned information. This includes more efficient or fluent processing of an item when it is repeated (i.e., repetition priming) and skill learning (see Chapter 1). Section 7.1 of this chapter considers the brain regions that have been associated with implicit memory, which include the dorsolateral prefrontal cortex and sensory processing regions (a subset of the regions associated with long-term memory; see Chapters 1 and 3). In section 7.2, the frequency bands of activity associated with implicit memory are discussed, which include gamma activity and alpha activity (a subset of the frequency bands of activity associated with long-term memory; see Chapter 4). Although there is some overlap between implicit memory and long-term memory with regard to the associated regions and frequency bands of activity, there are many notable differences that will be highlighted in this chapter. For instance, in direct opposition to the increases in cortical activity associated with long-term memory, implicit memory is typically associated with decreases in cortical activity. Section 7.3 details theoretical models of neural activity that underlie implicit memory effects and discusses ways in which these models can be distinguished from one another. In the fourth section, 7.4, evidence is considered that has claimed to link the hippocampus to implicit memory, which if true would contradict the evidence that this region is associated with only long-term memory.
• To understand the similarities between phrenology and fMRI.
• To list two advantages of ERPs over fMRI.
• To describe how brain region interaction studies are conducted.
• To characterize how the field of cognitive neuroscience will change in the future.
• To specify whether research on temporal processing in the brain will increase in the future.
Research on human memory is completely dependent on the methods that are employed in the field of cognitive neuroscience, and thus the future of memory research will follow the future of cognitive neuroscience. This final chapter focuses on the cognitive neuroscience techniques that have been employed in the past and the techniques that will be employed in the future. Section 11.1 describes the similarities between fMRI, which identifies brain regions associated with a cognitive process, and phrenology, a pseudoscience from two centuries ago in which each protrusion of the skull was associated with a particular behavioral characteristic. In section 11.2, fMRI is directly compared to ERPs. As fMRI has poor temporal resolution, only ERPs can measure the temporal dynamics of the functioning brain. A cost–benefit analysis favors ERPs, and government agencies are starting to increase funding for research that employs ERPs. Section 11.3 discusses research investigating brain region interactions, which will also receive increased government funding. Brain region interaction research has only recently started to be conducted and involves brain activity frequency analysis or modulating one brain region and measuring how that changes activity in another brain region. Section 11.4 provides an overview of the field of cognitive neuroscience in the future. A distinction is made between human brain mapping, which refers to identifying the brain regions associated with a cognitive process using fMRI, and research that investigates brain region interactions using EEG frequency analysis and combined techniques. It is predicted that human brain mapping research will be assimilated by the field of cognitive psychology and that the field of cognitive neuroscience will consist of human brain region interaction research and will be an area within the field of behavioral neuroscience. The final section, 11.5, shines a spotlight on the dimension of time. To date, temporal processing in the brain has received less attention than spatial localization. However, time is the future of the cognitive neuroscience of memory.
• To describe the cognitive processes and brain regions associated with visual attention.
• To compare the brain regions associated with visual attention to working memory and long-term memory.
• To describe the cognitive processes and brain regions associated with visual imagery.
• To compare the brain regions associated with visual imagery to working memory and long-term memory.
• To list the two primary brain regions associated with language processing and name two ways in which language processing is relevant to memory.
• To identify the two regions that interact to enhance memory for emotional information.
Attention is focused on the contents of all explicit memories. The experience of detailed recollection seems similar to the experience of vivid imagery. This chapter compares the cognitive processes and brain regions associated with memory to the cognitive processes and brain regions associated with attention, imagery, language, and emotion. Section 8.1 reviews the brain regions that have been associated with attention, which include sensory processing regions in addition to dorsolateral prefrontal cortex and parietal cortex control regions. These regions are similar to the regions that have been associated with working memory and long-term memory (except for the additional dependence of long-term memory on the medial temporal lobe; see Chapters 3 and 6). In section 8.2 of this chapter, the brain regions associated with imagery are reviewed, which also include sensory processing regions, the dorsolateral prefrontal cortex, and the parietal cortex. The cognitive processes and brain processes associated with visual imagery are compared to the cognitive processes and brain processes associated with working memory and long-term memory. Section 8.3 details the regions of the brain associated with language processing, which include the left inferior dorsolateral prefrontal cortex and the left posterior lateral temporal cortex. These regions are of relevance to memory studies, which often use words and meaningful objects as stimuli that have language/conceptual representations. The final section, 8.4, considers the brain regions that have been associated with emotion, which include the amygdala (a region just anterior to the hippocampus) and the dorsolateral prefrontal cortex.
• To understand the timing and location of brain activity associated with recollection and familiarity.
• To contrast the evidence on both sides of the scientific debate about activity that has been associated with familiarity.
• To describe what is meant by synchronous activity and how such activity indicates two brain regions interact.
• To list the three frequency bands of brain activity associated with long-term memory.
The large majority of human neuroscience research on long-term memory has focused on identifying the spatial locations of activity associated with this process (see Chapter 3). Although the temporal dimension of brain activity is often ignored, this does not mean that brain activity is static across time. In reality, brain activity changes rapidly across time, and the temporal dynamics of activity must be tracked to understand the brain mechanisms underlying memory. This chapter focuses on the timing of brain activity associated with long-term memory. As discussed previously (see Chapters 1 and 3), recollection refers to retrieval of detailed information, whereas familiarity refers to retrieval of non-detailed information. The chapter begins by introducing event-related potential (ERP) activations (see Chapter 2) that have been associated with familiarity and recollection (section 4.1). Familiarity has been associated with activity in frontal brain regions that occurs within 300 to 500 milliseconds after stimulus onset, while recollection has been associated with activity in parietal brain regions that occurs within 500 to 800 milliseconds after stimulus onset. In section 4.2, a scientific debate that has focused on the ERP activity associated with familiarity is discussed. In section 4.3, it is shown that synchronous activity in two different brain regions (i.e., activation timecourses that increase and decrease together) indicates that these regions interact. Such synchronous activity between regions during long-term memory typically occurs within specific frequency bands of activity including the theta frequency band (4 to 8 cycles per second, i.e., Hertz), the alpha frequency band (8 to 12 Hertz), and the gamma frequency band (greater than 30 Hertz). Theta activity reflects the interaction between the hippocampus and cortical regions during long-term memory, alpha activity reflects inhibition of cortical regions, and gamma activity reflects processing of features in different cortical regions that are combined to create a unified memory.
• To identify the regions of the medial temporal lobe that are associated with item memory, context memory, and binding item information and context information in rats, cats, and monkeys.
• To understand how long-term potentiation links cortical regions to the hippocampus.
• To compare the brain regions that have been associated with memory replay in rats and the brain regions associated with episodic memory in humans.
• To detail the paradigms that have been used to uncover time cells in the hippocampus of rats and monkeys.
• To describe one type of behavioral evidence and one type of brain evidence that indicates mammals have episodic memory.
This book is on the cognitive neuroscience of memory, so why is there a chapter on animal memory? One reason is that the same brain processes associated with memory in animals are often associated with memory in humans. These can be considered core brain processes that mediate memory across species. A second reason is that certain techniques can be used only on animals, such as targeted single-cell recording and brain lesions. The results of such techniques offer a detailed view into the brain mechanisms underlying memory that is not available in humans. This chapter focuses on long-term memory in animals, which relates to the large majority of research conducted with humans. Section 10.1 shows that rats, cats, and monkeys have a medial temporal lobe organization that is the same as humans. The perirhinal cortex is associated with item memory, the parahippocampal cortex is associated with context memory, and the hippocampus is associated with binding item information and context information. In section 10.2, long-term potentiation in the hippocampus is discussed, which is the mechanism by which cortical regions link to the hippocampus. Section 10.3 reviews evidence for memory replay in rats, which refers to reactivation of the same brain regions in the same or the reverse temporal sequence that were activated during a previous event.
• To identify three brain regions most commonly associated with episodic memory.
• To compare the brain regions associated with episodic memory and semantic memory.
• To contrast the two models of long-term memory consolidation.
• To explain what happens during slow wave sleep that promotes long-term memory consolidation.
• To compare the brain regions associated with memory retrieval and memory encoding.
• To describe how behavioral performance and hippocampal activity differ between females and males during long-term memory.
• To explain one way in which the brains of those with superior memory differ from those with normal memory.
This chapter considers the brain regions associated with long-term memory, a type of explicit memory (see Chapter 1). Long-term memory can be broken down into episodic memory and semantic memory. Episodic memory refers to the detailed retrieval of a previous episode, such as when someone remembers a happy moment of his or her life. Semantic memory refers to the retrieval of factual information, such as the definition of a word or the name of the current president. Semantic memories are formed through repeated exposure to information throughout life and lack the details associated with episodic memories. This information is simply known and there is no memory for the previous details of the learning experience. Although episodic memory and semantic memory both refer to conscious forms of retrieval, the degree of detail and subjective experience associated with these types of memory is quite different. It follows that the brain regions associated with episodic memory and semantic memory are also distinct. The first two sections of the chapter (sections 3.1 and 3.2) consider the brain regions associated with episodic memory and semantic memory. Section 3.3 will consider long-term memory consolidation (i.e., the process of creating more permanent memory representations in the brain). In section 3.4, the role of sleep in long-term memory consolidation is examined. Long-term memory consolidation requires the interaction between multiple brain regions in which activity oscillates at specific frequencies. In section 3.5, the brain regions associated with memory encoding will be reviewed.
• To identify the brain regions associated with typical forgetting.
• To understand the experimental paradigms that are used to investigate retrieval-induced forgetting and motivated forgetting.
• To describe the interaction between the dorsolateral prefrontal cortex and the hippocampus during retrieval-induced forgetting and motivated forgetting.
• To compare and contrast the brain regions associated with true memory, false memory for related information, and false memory for unrelated information.
• To determine one way in which flashbulb memories exemplify memory failure.
The previous two chapters focused on the brain mechanisms underlying successful long-term memory. The flip side of memory success is memory failure, and these processes are intimately linked. As will be discussed in more detail within this chapter, understanding memory failure furthers our understanding of memory success. Memory failure can be broadly classified into forgetting and memory distortion. Everyone is experienced with forgetting and, even though we are almost never aware of it, memory distortion. This chapter begins by reviewing the brain regions associated with typical forgetting, which can be attributed to a lack of attention during encoding (section 5.1). In section 5.2, the brain mechanisms underlying retrieval-induced forgetting are considered, which is when retrieval of one item (e.g., the word ‘banana’) has an inhibitory effect on related items (e.g., the word ‘orange’) and increases the rate of forgetting for these items. The brain regions associated with a related process called motivated forgetting, an increase in the rate of forgetting for items that one intentionally tries to forget, is then considered. In the next two sections of the chapter, 5.3 and 5.4, two types of memory distortion are considered: false memories (i.e., memories for information that did not occur) and flashbulb memories (i.e., seemingly picture-like memories for very surprising and consequential events). It has been argued that long-term memory failure reflects an adaptive memory system that works well (Schacter, 1999; Schacter, Guerin & St. Jacques, 2011).
• To identify the brain regions that are thought to store the contents of working memory.
• To describe how information is coded in early sensory regions during visual working memory.
• To list three shortcomings of the evidence or analysis techniques that have been used to associate working memory and the hippocampus.
• To compare and contrast the brain activity frequency bands associated with working memory and long-term memory.
• To understand what types of changes take place in the brain after extensive training on working memory tasks.
Working memory refers to actively holding information in mind during a relatively short period of time, typically seconds (see Chapter 1). Like most long-term memory paradigms, working memory paradigms consist of a study phase, a delay period, and a test phase. During working memory paradigms, information is actively kept in mind during the delay period. Working memory is an explicit process as its contents dominate conscious experience. Working memory has been associated with activity in the dorsolateral prefrontal cortex, the parietal cortex, and sensory processing regions. Thus, the regions associated with working memory are similar to those associated with long-term memory (see Chapter 3), with the notable absence of medial temporal lobe regions such as the hippocampus. Section 6.1 of this chapter details the brain regions that store the contents of working memory during the delay period. It has long been thought that the contents of working memory are stored in the dorsolateral prefrontal cortex, but more recent evidence indicates that storage also takes place in early sensory cortical regions such as V1. In section 6.2, the evidence is evaluated that claims to link working memory with the hippocampus. In section 6.3, brain activity associated with working memory that oscillates at particular frequencies is considered, which includes alpha activity and gamma activity. This also mirrors the findings of long-term memory (see Chapter 4), except for the lack of working memory theta activity. Finally, in section 6.4, changes in brain activity are highlighted that have been linked to training-related increases in working memory capacity. These findings suggest that extensive training (e.g., multiple times a week for many weeks) on working memory tasks can produce long-term improvements in behavioral performance, change the way the brain functions for a period well beyond the time of training, and perhaps even increase intelligence.
• To describe the changes in brain anatomy and fMRI activity in patients with amnestic mild cognitive impairment.
• To identify the regions of the brain that atrophy in patients with early Alzheimer's disease and learn the proteins that are accumulated in these regions.
• To compare the behavioral performance and fMRI activations of mild traumatic brain injury patients and healthy control participants during working memory tasks.
• To understand how surgery on medial temporal lobe epilepsy patients has revealed associations between the left medial temporal lobe and the right medial temporal lobe and verbal long-term memory and visual long-term memory.
• To specify the location of the hippocampal lesion that causes transient global amnesia.
The previous chapters of this book have focused on the neural basis of memory in healthy adults. This chapter discusses five neurological diseases that affect the brain regions associated with explicit memory. Section 9.1 discusses patients with amnestic mild cognitive impairment. These patients have long-term memory deficits due to atrophy of medial temporal lobe regions including the hippocampus. Within a few years of being diagnosed with amnestic mild cognitive impairment, about half of these individuals are diagnosed with Alzheimer's disease, the topic of section 9.2. Patients with early Alzheimer's disease have more severe impairment of long-term memory and atrophy of the medial temporal lobe and the parietal lobe, two regions that have been associated with long-term memory (see Chapter 3). Alzheimer's disease patients also have abnormally high levels of proteins in the medial temporal lobe and the parietal lobe, which is thought to further disrupt processing in these regions. Section 9.3 focuses on patients with mild traumatic brain injury, who typically perform normally on working memory tasks but have increased fMRI activity within the dorsolateral prefrontal cortex and the parietal cortex, relative to healthy control participants. It is generally believed that such increases in fMRI activity reflect compensation, where these regions are recruited to perform normally on the task.