Neuropsychologia 48 (2010) 491–497
Contents lists available at ScienceDirect
Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia
Dissociating the “retrieval success” regions of the brain:
Effects of retrieval delay
W. Huijbers
∗
, C.M.A. Pennartz, S.M. Daselaar
University of Amsterdam, Swammerdam Institute for Life Sciences, Faculty of Science, The Netherlands
article info
Article history:
Received 24 April 2009
Received in revised form 6 October 2009
Accepted 7 October 2009
Available online 14 October 2009
Keywords:
Episodic memory
fMRI
Evidence accumulation
Posterior midline region
Posterior parietal cortex
abstract
There is abundant evidence that the hippocampal formation critically supports episodic memory retrieval,
the remembering of episodes including contextual details. Yet, a group of other brain regions has also been
consistently implicated in successful episodic retrieval. This retrieval success network (RSN) includes the
posterior midline region, medial prefrontal cortex (mPFC), and posterior parietal cortex (PPC). Despite
these consistent findings, the functional roles of the RSN regions remain poorly understood. Given that
vivid remembering leads to high-confidence retrieval decisions, it is unclear whether activity in these
regions reflects episodic long-term memory, or is merely associated with retrieval confidence. In order
to distinguish between these alternatives, we manipulated study-test delays within the context of a
continuous recognition task during fMRI-scanning. The design was based on previous evidence indicating
that retrieval at short delays is easier leading to high-evidence mnemonic decisions, whereas retrieval at
longer delays is more difficult but also more hippocampus-dependent. Confirming previous findings, we
found that retrieval decisions at short delays were more accurate and faster, and that the hippocampus
showed greater activity at longer delays. Within the other RSN regions, we found three distinct activation
patterns as a function of delay. Similar to the hippocampus, the retrosplenial cortex showed increased
activity as a function of retrieval delay. Dorsal PPC and the precuneus showed decreased activity. Finally,
the posterior cingulate, medial PFC and ventral PPC showed a V-shaped pattern. These findings support
the idea that dorsal PPC and the precuneus are involved in decision-related retrieval processes rather
than successful remembering.
© 2009 Elsevier Ltd. All rights reserved.
1. Introduction
In the past few decades, clinical studies in both animals and
humans have yielded overwhelming evidence that the integrity
of the hippocampal formation is critical to normal episodic mem-
ory retrieval, the remembering of episodes including contextual
details (Eichenbaum, Yonelinas, & Ranganath, 2007; Squire, Stark,
& Clark, 2004). More recently, functional MRI studies have con-
sistently implicated another group of brain regions in successful
episodic retrieval. This retrieval success network (RSN) includes
the medial prefrontal cortex (mPFC), the posterior parietal cor-
tex (PPC), and the posterior midline region (PMR), comprising the
precuneus, the posterior cingulate, and the retrosplenial cortex
(Buckner, Andrews-Hanna, & Schacter, 2008; Wagner, Shannon,
Kahn, & Buckner, 2005). Functional Magnetic Resonance Imag-
ing (fMRI) studies of episodic retrieval found greater activity in
these regions during correct recognition of studied items, than dur-
∗
Corresponding author. Tel.: +31205257728.
E-mail address: w.huijbers@uva.nl (W. Huijbers).
ing misses and correctly rejected non-studied items (e.g., Henson,
Hornberger, & Rugg, 2005). Moreover, these differences increase
with retrieval confidence (e.g., Daselaar, Fleck, & Cabeza, 2006).
Yet, despite these consistent findings, the role of the RSN regions in
episodic retrieval remains poorly understood. A critical issue is that
successful retrieval often leads to high-confidence memory deci-
sions. Thus, it is unclear whether activity in the RSN truly reflects
episodic long-term memory or is merely associated with confident
retrieval decisions.
Regarding the latter option, there is substantial evidence that
the posterior parietal regions of the RSN are involved in processes
supporting perceptual decision-making (Platt & Glimcher, 1999;
Shadlen & Newsome, 1996). For instance in the monkey brain,
PPC neurons have been linked to the accumulation and integration
of evidence from multiple perceptual regions prior to a decision
(Shadlen & Newsome, 2001). A functional MRI study on percep-
tual recognition has recently reported analogous findings in the
human PPC (Ploran et al., 2007). Models of recognition memory
also assume that retrieval decisions are based on the accumulation
of evidence (Ratcliff, 1978; Ratcliff & Starns, 2009). These models
assert that memory judgments are based on a diffusion-like process
by which evidence is gathered for or against a memory decision.
0028-3932/$ – see front matter © 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2009.10.006