Daniel L. Schacter

The brain’s default network: Anatomy, function, and relevance to disease

Thirty years of brain imaging research has converged to define the brains default network—a novel and only recently appreciated brain system that participates in internal modes of cognition. Here we synthesize past observations to provide strong evidence that the default network is a specific, anatomically defined brain system preferentially active when individuals are not focused on the external environment. Analysis of connectional anatomy in the monkey supports the presence of an interconnected brain system. Providing insight into function, the default network is active when individuals are engaged in internally focused tasks including autobiographical memory retrieval, envisioning the future, and conceiving the perspectives of others. Probing the functional anatomy of the network in detail reveals that it is best understood as multiple interacting subsystems. The medial temporal lobe subsystem provides information from prior experiences in the form of memories and associations that are the building blocks of mental simulation. The medial prefrontal subsystem facilitates the flexible use of this information during the construction of self‐relevant mental simulations. These two subsystems converge on important nodes of integration including the posterior cingulate cortex. The implications of these functional and anatomical observations are discussed in relation to possible adaptive roles of the default network for using past experiences to plan for the future, navigate social interactions, and maximize the utility of moments when we are not otherwise engaged by the external world. We conclude by discussing the relevance of the default network for understanding mental disorders including autism, schizophrenia, and Alzheimers disease.

Implicit Memory: History and Current Status

Memory for a recent event can be expressed explicitly, as conscious recollection, or implicitly, as a facilitation of test performance without conscious recollection. A growing number of recent studies have been concerned with implicit memory and its relation to explicit memory. This article presents an historical survey of observations concerning implicit memory, reviews the findings of contemporary experimental research, and delineates the strengths and weaknesses of alternative theoretical accounts of implicit memory. It is argued that dissociations between implicit and explicit memory have been documented across numerous tasks and subject populations, represent an important challenge for research and theory, and should be viewed in the context of other dissociations between implicit and explicit expressions of knowledge that have been documented in recent cognitive and neuropsychological research. Psychological studies of memory have traditionally relied on tests such as free recall, cued recall, and recognition. A prominent feature of these tests is that they make explicit reference to, and require conscious recollection of, a specific learning episode. During the past several years, however, increasing attention has been paid to experimental situations in which information that was encoded during a particular episode is subsequently expressed without conscious or deliberate recollection. Instead of being asked to try to remember recently presented information, subjects are simply required to perform a task, such as completing a graphemic fragment of a word, indicating

Implicit and explicit memory for new associations in normal and amnesic subjects

Two experiments examined whether repetition priming effects on a word completion task are influenced by new associations between unrelated word pairs that were established during a single study trial. On the word completion task, subjects were presented with the initial three letters of the response words from the study list pairs and they completed these fragments with the first words that came to mind. The fragments were shown either with the paired words from the study list (same context) or with other words (different context). Both experiments showed a larger priming effect in the same-context condition than in the different-context condition, but only with a study task that required elaborative processing of the word pairs. This effect was observed with college students and amnesic patients, suggesting that word completion performance is mediated by implicit memory for new associations that is independent of explicit recollection.

Remembering the past and imagining the future: Common and distinct neural substrates during event construction and elaboration

People can consciously re-experience past events and pre-experience possible future events. This fMRI study examined the neural regions mediating the construction and elaboration of past and future events. Participants were cued with a noun for 20s and instructed to construct a past or future event within a specified time period (week, year, 5-20 years). Once participants had the event in mind, they made a button press and for the remainder of the 20s elaborated on the event. Importantly, all events generated were episodic and did not differ on a number of phenomenological qualities (detail, emotionality, personal significance, field/observer perspective). Conjunction analyses indicated the left hippocampus was commonly engaged by past and future event construction, along with posterior visuospatial regions, but considerable neural differentiation was also observed during the construction phase. Future events recruited regions involved in prospective thinking and generation processes, specifically right frontopolar cortex and left ventrolateral prefrontal cortex, respectively. Furthermore, future event construction uniquely engaged the right hippocampus, possibly as a response to the novelty of these events. In contrast to the construction phase, elaboration was characterized by remarkable overlap in regions comprising the autobiographical memory retrieval network, attributable to the common processes engaged during elaboration, including self-referential processing, contextual and episodic imagery. This striking neural overlap is consistent with findings that amnesic patients exhibit deficits in both past and future thinking, and confirms that the episodic system contributes importantly to imagining the future.

Remembering the past to imagine the future: The prospective brain.

A rapidly growing number of recent studies show that imagining the future depends on much of the same neural machinery that is needed for remembering the past. These findings have led to the concept of the prospective brain; an idea that a crucial function of the brain is to use stored information to imagine, simulate and predict possible future events. We suggest that processes such as memory can be productively re-conceptualized in light of this idea.

The cognitive neuroscience of constructive memory: remembering the past and imagining the future

Episodic memory is widely conceived as a fundamentally constructive, rather than reproductive, process that is prone to various kinds of errors and illusions. With a view towards examining the functions served by a constructive episodic memory system, we consider recent neuropsychological and neuroimaging studies indicating that some types of memory distortions reflect the operation of adaptive processes. An important function of a constructive episodic memory is to allow individuals to simulate or imagine future episodes, happenings and scenarios. Since the future is not an exact repetition of the past, simulation of future episodes requires a system that can draw on the past in a manner that flexibly extracts and recombines elements of previous experiences. Consistent with this constructive episodic simulation hypothesis, we consider cognitive, neuropsychological and neuroimaging evidence showing that there is considerable overlap in the psychological and neural processes involved in remembering the past and imagining the future.

The Evolution of Multiple Memory Systems

The existence of multiple memory systems has been proposed in a number of areas, including cognitive psychology, neuropsychology, and the study of animal learning and memory. We examine whether the existence of such multiple systems seems likely on evolutionary grounds. Multiple systems adapted to serve seemingly similar functions, which differ in important ways, are a common evolutionary outcome. The evolution of multiple memory systems requires memory systems to be specialized to such a degree that the functional problems each system handles cannot be handled by another system. We define this condition as functional incompatibility and show that it occurs for a number of the distinctions that have been proposed between memory systems. The distinction between memory for song and memory for spatial locations in birds, and between incremental habit formation and memory for unique episodes in humans and other primates provide examples. Not all memory systems are highly specialized in function, however, and the conditions under which memory systems could evolve to serve a wide range of functions are also discussed. Memory is a function that permits animals and people to acquire, retain, and retrieve many different kinds of information. It allows them to take advantage of previous experience to help solve the multitude of problems with which their environment confronts them, such as how to recognize the familiar, predict events, return to particular places, and assess the consequences of behavior Recently the question has arisen as to whether the

Priming and the brain

This work was supported by National Institute on Aging AG08441 and the Human Frontiers Science Foundation. We thank Alex Martin and Eric Kandel for comments and discussion and Carolyn Brenner for help with preparation of the manuscript.

Medial Temporal Lobe Activations in fMRI and PET Studies of Episodic Encoding and Retrieval

Early neuroimaging studies often failed to obtain evidence of medial temporal lobe (MTL) activation during episodic encoding or retrieval, but a growing number of studies using functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) have provided such evidence. We review data from fMRI studies that converge on the conclusion that posterior MTL is associated with episodic encoding; too few fMRI studies of retrieval have reported MTL activations to allow firm conclusions about their exact locations. We then turn to a recent meta‐analysis of PET studies (Lepage et al., Hippocampus 1998;8:313–322) that appears to contradict the fMRI encoding data. Based on their analysis of the rostrocaudal distribution of activations reported during episodic encoding or retrieval, Lepage et al. (1998) concluded that anterior MTL is strongly associated with episodic encoding, whereas posterior MTL is strongly associated with episodic retrieval. After considering the evidence reviewed by Lepage et al. (1998) along with additional studies, we conclude that PET studies of encoding reveal both anterior and posterior MTL activations. These observations indicate that the contradiction between fMRI and PET studies of encoding was more apparent than real. However, PET studies have reported anterior MTL encoding activations more frequently than have fMRI studies. We consider possible sources of these differences. Hippocampus 1999;9:7–24.

Functional-Anatomic Correlates of Object Priming in Humans Revealed by Rapid Presentation Event-Related fMRI

Human functional-anatomic correlates of object repetition were explored in a cohort of 20 subjects using fMRI. Subjects performed an object classification task where the target objects were either novel or repeated. Objects appeared rapidly, one every 2 s, in a randomly intermixed task design similar to traditional behavioral, event-related potential (ERP), and single-unit physiological studies. Recently developed event-related fMRI methods were used to analyze the data. Clear effects of repetition were observed. Brain areas in midlevels of the processing hierarchy, including extrastriate visual cortex extending into inferotemporal cortex and left dorsal prefrontal cortex, showed reductions in the amount of activation after repetition. By contrast, early visual areas and output motor areas were activated equally by both novel and repeated objects and did not show effects of repetition, suggesting that the observed correlates of repetition were anatomically selective. We discuss these findings in relation to previous positron emission tomography (PET) and fMRI studies of item repetition and single-unit physiological studies; we also address the broad impact that rapid event-related fMRI is likely to have on functional neuroimaging.