High‐resolution fMRI is an essential technique for evaluating theories of MTL function that had previously been tested only in animals. The combination of hr‐
fMRI and multivariate pattern‐information analysis techniques, in particular, has substantially advanced our understanding of how memory is represented in MTL subregions, allowing for greater parity and convergence with animal studies. High‐
resolution fMRI techniques also have increasing translational relevance. Many of the paradigms described in this review are being applied to clinical populations with neurological and psychiatric disorders associated with memory impairment (e.g., Gaisler‐Salomon et al., 2009; Schobel et al., 2009; Suthana et al., 2009; Yassa et al., 2010), providing further insight into the relationship between pathological changes to MTL subregions and disease processes that affect memory. New technical develop- ments are likely to advance the field further, affording greater leverage to characterize the critical MTL computations and representations that underlie episodic memory.
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The Wiley Handbook on the Cognitive Neuroscience of Memory, First Edition.
Edited by Donna Rose Addis, Morgan Barense, and Audrey Duarte.
© 2015 John Wiley & Sons, Ltd. Published 2015 by John Wiley & Sons, Ltd.
Memory Retrieval and the Functional Organization
of Frontal Cortex
Erika Nyhus and David Badre
7
Introduction
To be adaptive, the long‐term memory system must support retrieval of previously stored knowledge that has high utility given our current task and goals. In this fram- ing, the problem of memory retrieval concerns balancing the recovery of useful information on the one hand against the inherent costs associated with retrieval itself (Anderson and Milson, 1989). Central to striking this balance is cognitive control function (sometimes called executive function), or the ability to leverage abstract goals and contextual representations in order to adaptively influence retrieval and memory‐based performance.
The prefrontal cortex (PFC) is necessary for cognitive control function, including during the cognitive control of memory. Whereas damage to medial temporal lobe (MTL) structures produces amnesia that catastrophically impairs the encoding of new information and retrieval of recently encoded information (Scoville and Milner, 1957), damage to the PFC results in more subtle memory deficits (Moscovitch, 1992;
Stuss and Alexander, 2005). For example, PFC patients are impaired in contexts that require retrieval of specific information (i.e., source memory tasks; Janowsky et al., 1989; Swick, Senkfor, and Van Petten, 2006), reliance on retrieval strategies (Moscovitch and Melo, 1997), overcoming interference (Moscovitch, 1982; Squire, 1982; Winocur, Kinsbourne, and Moscovitch, 1981), ordering information at retrieval (Shimamura, Janowsky, and Squire, 1990), or retrieval with limited cue support (e.g., free recall; Janowsky, Shimamura, and Squire, 1989; Jetter et al., 1986; Stuss et al.
1994). Neuroimaging studies have similarly implicated PFC in relation to specific manipulations of cognitive control at retrieval (Badre and Wagner, 2007; Fletcher and Henson, 2001; Rugg and Wilding, 2000). Thus, PFC is a crucial component of the system that supports cognitive control of memory.
In general, the mechanism of cognitive control can be described in terms of a pro- cess of guided activation (Miller and Cohen, 2001), wherein contextual or goal information is maintained in working memory and thereby has the opportunity to provide a top‐down influence on processing elsewhere. Nevertheless, a central debate
in the study of cognitive control concerns whether cognitive control is a unitary process or involves a diverse set of functionally distinguishable control processes (e.g., Cooper, 2010).
Mirroring the debate about the componentiality of cognitive control, the PFC is likely not a functionally homogeneous structure supporting a unitary executive, but may contain distinct subsystems that support different forms of cognitive control.
Neuroimaging studies have provided the primary evidence in favor of functional dissociations in PFC (Badre and Wagner, 2007; Simons and Spiers, 2003; Spaniol et al., 2009). Though debate still exists as to whether there are undifferentiated “multiple‐demand zones” within the PFC – such as within the mid‐dorsolateral PFC (Duncan, 2010) – it seems now widely accepted that functional distinctions likely exist, such as between ventral and dorsal lateral PFC (Petrides, 2002; Simons and Spiers, 2003), rostral versus caudal PFC (Buckner, 2003; Race, Shanker, and Wagner, 2008), left versus right lateral PFC (Nolde, Johnson, and Raye, 1998;
Tulving et al., 1994), and lateral PFC versus medial and subcortical systems (Kuhl et al., 2008; Scimeca and Badre, 2012). Considerable controversy remains, how- ever, regarding the validity of these distinctions and how to map them onto both individual experimental tasks and real‐world behavior.
The cognitive control of memory is also likely componential, emerging from a set of interacting component processes. For example, cognitive control has the oppor- tunity to influence retrieval performance in several ways, ranging from processes that structure inputs to the memory system (e.g., cue elaboration) to output control that monitors the outcome of retrieval and selects which representations are permitted to influence decision and action (Benjamin, 2007; see also Chapter 5). These and other distinct memory control processes could likewise be supported by different brain sys- tems. Though research has only begun to refine understanding of these mechanisms and their neural correlates, several distinctions have been proposed regarding differential control processing in the PFC (e.g., Badre and Wagner, 2007; Nyberg, Cabeza, and Tulving, 1996; Simons and Spiers, 2003; Spaniol et al., 2009; Tulving et al., 1994).
Here we will focus on an example of functional specification within PFC related to cognitive control of memory retrieval. We will discuss the hypothetical specialization within ventrolateral PFC (vlPFC) between controlled retrieval and post‐retrieval selection/monitoring operations. We will conclude by broadening the discussion of this distinction to consider the participation of these PFC subregions within distinct larger‐scale functional networks.