Neuropsychologia 80 (2016) 3546

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Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Default network activation during episodic and semantic memory

retrieval: A selective meta-analytic comparison
Hongkeun Kim
Department of Rehabilitation Psychology, Daegu University, Gyeongsangbuk-Do 38453, South Korea

art ic l e i nf o a b s t r a c t

Article history: It remains unclear whether and to what extent the default network subregions involved in episodic
Received 3 August 2015 memory (EM) and semantic memory (SM) processes overlap or are separated from one another. This
Received in revised form study addresses this issue through a controlled meta-analysis of functional neuroimaging studies in-
7 November 2015
volving healthy participants. Various EM and SM task paradigms differ widely in the extent of default
Accepted 9 November 2015
Available online 10 November 2015
network involvement. Therefore, the issue at hand cannot be properly addressed without some control
for this factor. In this regard, this study employs a two-stage analysis: a preliminary meta-analysis to
Keywords: select EM and SM task paradigms that recruit relatively extensive default network regions and a main
Fmri analysis to compare the selected task paradigms. Based on a within-EM comparison, the default network
Default mode network
contributed more to recollection/familiarity effects than to old/new effects, and based on a within-SM
Episodic memory
comparison, it contributed more to word/pseudoword effects than to semantic/phonological effects.
Semantic memory
Meta-analysis According to a direct comparison of recollection/familiarity and word/pseudoword effects, each involving
a range of default network regions, there were more overlaps than separations in default network
subregions involved in these two effects. More specically, overlaps included the bilateral posterior
cingulate/retrosplenial cortex, left inferior parietal lobule, and left anteromedial prefrontal regions,
whereas separations included only the hippocampal formation and the parahippocampal cortex region,
which was unique to recollection/familiarity effects. These results indicate that EM and SM retrieval
processes involving strong memory signals recruit extensive and largely overlapping default network
regions and differ mainly in distinct contributions of hippocampus and parahippocampal regions to EM
retrieval.
& 2015 Elsevier Ltd. All rights reserved.

1. Introduction reportedly specic to EM or SM retrieval.

1.1. The purpose of the study
The distinction between episodic memory (EM) and semantic
memory (SM) has sparked a massive number of experimental and
lesion studies (Tulving, 1972). Here a critical issue concerns whe-
ther and to what extent EM and SM processes involve similar or
differential neural substrates. Although this issue has been in-
vestigated by many functional neuroimaging studies (e.g., Bur-
ianova and Grady, 2007; Dzel et al., 1999; Maguire and Mum-
mery, 1999; Nyberg et al., 2003; Rajah andMcIntosh, 2005; Sha-
pira-Lichter et al., 2013; Wiggs et al., 1998), it remains con-
troversial because of highly mixed results across studies. Previous
studies have indicated the common involvement of various frontal,
temporal, parietal, occipital, and subcortical subregions and de-
monstrated the existence of similar diversity for regions
E-mail address: hongkn1@gmail.com
Such mixed results likely reect the fact that different types of
EM tasks and related activation contrasts recruit different sets of
brain regions and that this also holds for different types of SM
tasks and associated contrasts. For example, in both EM and SM,
some tasks involve a relatively high demand for controlled pro-
cessing, whereas others, a low demand. If EM and SM retrieval
tasks selected for a comparison entail a high demand for con-
trolled processing, common regions for the two tasks may include
fronto-parietal control regions such as the lateral prefrontal cortex
(PFC). However, if both tasks involve a relatively low demand for
controlled processing, then the PFC and other regions are less
likely to show some common activity. In addition, the PFC and
other regions may emerge as specic to either EM or SM retrieval
if the demand for controlled processing is high in one task but low
in the other. Although many studies have addressed these and
other potentially biasing factors, a fully adequate control scheme
has not always been practical because of the complexity of each
task's cognitive processes. Nonetheless, a meaningful and

http://dx.doi.org/10.1016/j.neuropsychologia.2015.11.006
0028-3932/& 2015 Elsevier Ltd. All rights reserved.
36 H. Kim / Neuropsychologia 80 (2016) 3546

theoretically important comparison between EM and SM pro-
cesses requires the careful selection of task paradigms and the
discrimination of commonalities/differences from intrinsic pro-
cesses and those from extrinsic factors.
Recent studies have indicated that both EM and SM retrieval
processes are related to a large-scale, intrinsic network often re-
ferred to as the default network (Raichle et al., 2001), as reviewed
in more detail in the next section. However, it remains largely
unclear whether and to what extent the default network sub-
regions involved in EM and SM processes overlap or are separated
from one another. To address this issue, this study provides a se-
lective and controlled meta-analysis of relevant functional neu-
roimaging studies involving healthy participants.
1.2. The default network and memory retrieval
Recent advances in the eld can provide a useful framework for
a comparison between EM and SM processes and suggest that
both involve the default network. Major components of the default
network include the anteromedial PFC extending laterally to the
dorsal and ventral PFC, the posterior cingulate cortex extending
ventrally to the retrosplenial cortex, the inferior parietal lobule,
the lateral temporal cortex, and the medial temporal lobe, in-
cluding the hippocampal formation (see pink regions in Fig. 1).
Functional roles of the default network are not fully understood,
but there is strong evidence that it plays a critical role in any in-
ternally oriented mental activity such as recall, future thinking,
social cognition, stimulus-independent thought, and other spon-
taneous cognition (AndrewsHanna et al., 2014).
Evidence that EM retrieval is associated with the default net-
work comes from studies using the autobiographical memory
paradigm (Buckner and Carroll, 2007; Kim, 2012). Until recently,
however, similar evidence has been lacking in studies using la-
boratory-based EM tasks. For example, one of the most proto-
typical task paradigms in using neuroimaging in the analysis of
laboratory-based EM retrieval is to compare the correct recogni-
tion of studied items (hit) with the correct non-recognition of
novel items (correct rejection). This old/new (i.e., old 4new) effect
shows only a moderate overlap with default network regions, as
demonstrated in recent meta-analyses (Kim, 2013; McDermott
et al., 2009; Spaniol et al., 2009). Recognition memory may be
based on either vivid recall of an item and associated contexts
(recollection) or a feeling of oldness in the absence of any con-
textual information (familiarity). The contrast of recollection with
familiarity has been widely used to probe the EM retrieval process
involving autonoetic consciousness and other related properties
(Gardiner, 2001). A meta-analysis of this recollection/familiarity
contrast indicated a more consistent relationship with default
network regions (Kim, 2010), providing clearer evidence linking
laboratory-based EM retrieval to the default network. However, no
meta-analysis has directly compared recollection/familiarity and
old/new effects in the extent of default network involvement.
More generally, many laboratory-based EM task types activate at
least some default network regions, but they also show relatively
large differences in the scope of default network involvement. In
this regard, there is a need to determine the extent to which
various EM retrieval paradigms differ in default network involve-
ment and why.
Binder et al.'s (2009) meta-analysis has recently played a key
role in disseminating the notion that SM retrieval is associated
with the default network. Binder et al. restricted their meta-ana-
lysis to studies using verbal stimuli, arguing that much of human
knowledge is represented symbolically through language and in-
dicating their desire to maintain a clear focus on the processing of
concepts rather than percepts. Their analysis considered about 30
types of SM task paradigms simultaneously, many of which were
used in only one or two studies. The most common paradigm in
their analysis, referred to here as the word/pseudoword paradigm,
involves the presentation of words and pseudowords in target and
baseline conditions, respectively, and lexical decisions. The second
most common paradigm is the semantic/phonological paradigm,
which involves semantic and phonological processing tasks for
words in target and baseline conditions, respectively. Their meta-
analysis involving these and other paradigms indicated a clear
overlap with default network regions. However, some caution is
necessary because a consistent relationship with default network
regions may be the property of not all but only some paradigms
included in the meta-analysis. For example, some studies that

dPFC IPS PCC/RSC

pmPFC
midPFC IPL dPCU

vPFC amPFC
aINS pMTG MTL
LTC

Default network Fronto-parietal control network

Fig. 1. Estimates of the default-mode network (shown in pink) and the fronto-parietal cognitive control network (green). Estimates were produced using data from Yeo et al.
(2011), which can be downloaded from the Surface Management System Database (SumsDB; http://sumsdb.wustl.edu/sums). Data were from a clustering analysis of resting-
state functional MRI signals involving 1175 uniformly spaced cortical regions and 1000 subjects. aINS, anterior insula; amPFC, anteromedial prefrontal cortex; dPCU, dorsal
precuneus; dPFC, dorsal prefrontal cortex; IPL, inferior parietal lobule; IPS, intraparietal sulcus; LTC, lateral temporal cortex; midPFC, middle prefrontal cortex; MTL, medial
temporal lobe; PCC, posterior cingulate cortex; pmPFC, posteromedial prefrontal cortex; pMTG, posterior middle temporal gyrus; RSC, retrosplenial cortex; vPFC, ventral
prefrontal cortex. (For interpretation of the references to color in this gure legend, the reader is referred to the web version of this article.)
 H. Kim / Neuropsychologia 80 (2016) 3546
contrasted a semantic processing task with a phonological pro-
cessing task have indicated only modest default network in-
volvement (e.g., Emmorey et al., 2013; Price et al., 1997). Therefore,
as in EM retrieval, there is a need to determine the extent to which
various SM retrieval paradigms differ in default network involve-
ment and why.
Conscious memory retrieval, both episodic and semantic, is
intrinsically associated with attention to internal representations.
Therefore, the involvement of the default network in EM and SM
retrieval is consistent with the notion that the default network
supports internally oriented mentation. Here a related issue of
great importance concerns whether and to what extent these two
processes involve similar or differential default network sub-
regions. Although several studies have compared neural substrates
of these processes, these questions remain largely underexplored
because most studies have been designed without a clear con-
ceptualization of the default network and its role in EM and SM
retrieval.
EM and SM processes differ in many aspects, including their
phenomenological experience, self-references versus cognitive
references, and simultaneous retrieval and integration of what,
where, and when information unique to EM. However, both
EM and SM represent consciously retrieved long-term cognitive
memory (i.e., declarative memory). In addition, their operations
are highly intertwined with each other (Greenberg and Verfaellie,
2010; Irish and Piguet, 2013; Klein, 2013), suggesting some broadly
shared neural substrates. In encoding, one's new SM forms from
repeated episodic experiences, and the semantic processing of
some to-be-remembered material facilitates the storage of new
EM. In retrieval, EM retrieval necessarily includes semantic re-
presentations in that recalling I visited a museum yesterday
necessitates the retrieval of the concepts visit, museum, and
yesterday (Binder et al., 2009), and SM retrieval can be aug-
mented by calling on relevant episodic information (Ryan et al.,
2008). Many memory types are collectively called personal se-
mantics (Renoult et al., 2012) and have some features of both EM
and SM, indicating that the two types of memory have less clear
boundaries than initially conceived. For example, memory for re-
peated personal events (e.g., making morning coffee in the kitch-
en), or repisodic memory (Neisser, 1981), does not fully qualify
as episodic but frequently includes specic spatial and contextual
information. Therefore, a general prediction is that the default
network subregions involved in the two memory processes show
no complete overlap or separation from one another but a mixture
of overlaps and separations.
1.3. A controlled meta-analytic comparison
Many within-study comparisons of EM and SM process studies
have held items constant in the two conditions and required
subjects to process the same items either episodically (e.g., old/
new judgments) or semantically (e.g., living/nonliving judgments)
(Barba et al., 1998; Burianova and Grady, 2007; Dzel et al., 1999;
Hantke et al., 2013; St.-Laurent et al., 2011; Wiggs et al., 1998).
Although ndings vary widely across studies, EM and/or SM tasks
have generally been found to involve default network regions to
some limited extent, and therefore default network regions rarely
gure prominently in either common or specic effects. Although
these ndings may seem puzzling because of the presumed pivotal
role of the default network in mnemonic processing, they em-
phasize the notion that not all conventional memory tasks and
associated activation contrasts consistently tap cognitive processes
supported by the default network. Here the proposed hypothesis is
that not all but only task paradigms that involve both strong
memory signals in target conditions and weak (or no) memory
signals in reference conditions entail a high level of default
37
network involvement. According to this hypothesis, EM and SM
task paradigms involving only modest default network contribu-
tions may be associated with relatively weak memory signals in
target conditions and/or relatively strong memory signals in re-
ference conditions. Some evidence providing support for this hy-
pothesis is presented later. For now, such EM and SM task para-
digms are less useful for addressing the question at hand in this
study (i.e., whether and to what extent the default network sub-
regions involved in EM and SM processes overlap or are separated
from one another).
Various EM and SM task paradigms differ widely in the extent
of default network involvement. Therefore, the issue at hand
cannot be properly addressed without some control for this factor.
A better comparison would require that both EM and the SM task
paradigms selected for a comparison involve default network re-
gions to a relatively large extent. If this requirement is met in only
one task, then the comparison is likely to be biased (i.e., yield
spurious differences), and if it is not met in either condition, then
the comparison cannot identify the motivating factor. To address
this issue, the present meta-analysis was conducted in two stages.
The rst (preliminary) stage involved two within-memory com-
parisons: a comparison of two EM task paradigms (recollection/
familiarity and old/new) to determine which one would engage
the default network in a more consistent manner and a compar-
ison of two SM task paradigms (word/pseudoword and semantic/
phonological) for a similar objective. Although the two task
paradigms in each subject area are not exhaustive of all reported
in the literature, they do represent those that have been most
widely used and thus are considered highly relevant in that area.
The second (main) stage involved a comparison between the EM
task paradigm showing the more consistent involvement of the
default network in the rst-stage comparison and the SM task
paradigm showing the same. Fortunately, as discussed in the Re-
sults section, EM and SM task paradigms were relatively equal in
the extent of default network involvement, rendering their com-
parison balanced in that regard.
2. Materials and methods
2.1. Data collection
The meta-analysis focused on the following four task paradigms: recollection/
familiarity, old/new, word/pseudoword, and semantic/phonological. Functional
neuroimaging studies involving these task paradigms were identied through
multiple searches of the Pubmed and Google Scholar databases. The search terms
included fMRI, PET, episodic memory, recollection, recognition memory,
semantic memory, lexical decision, pseudowords, and semantic processing.
Reference lists of recent meta-analyses (Binder et al., 2009; Kim, 2013, 2015) were
also considered. Then search results were ltered to include only those studies that
(1) included healthy participants, (2) performed whole-brain analyses, and (3) re-
ported activation foci in the standard reference space (i.e., Talairach or Montreal
Neurological Institute [MNI] coordinates). Ultimately, the meta-analysis involved a
total of 105 studies (articles) collectively involving 1792 participants (see Sup-
porting information References for a list of studies included in the meta-analysis).
The imaging modality was functional magnetic resonance imaging in 99 studies
and positron emission tomography in the rest. The sensory modality for the sti-
mulus presentation was visual in 99 studies and auditory in the rest.
2.2. Task paradigms
Table 1 lists the numbers of experiments, peak foci, and participants for each of
the four task paradigms (see Supporting information Tables S1S4 for a detailed
description of studies included in each category). Although most studies have in-
cluded only one relevant experiment of the same task paradigm, some reported
more. In the case of the latter, if multiple experiments involved entirely separate
sets of trials, all were included. Otherwise, only one was selected for inclusion to
prevent these studies from overly inuencing results. The experiment with the
largest number of associated peak coordinates was selected because this approach
was considered to likely enhance the sensitivity of the meta-analysis.
Recollection/familiarity contrasts involved a comparison of recollection
38 H. Kim / Neuropsychologia 80 (2016) 3546
Table 1
A summary of four different task paradigms in the meta-analysis.
Memory type Task paradigms Experiments Foci Participants
EM retrieval Recollection 4familiarity 27 400 529
Old 4new 42 571 746
SM retrieval Word4pseudoword 20 211 338
Semantic 4phonological 17 120 179
EM, episodic memory; SM, semantic memory.
responses with familiarity responses in a recognition memory test. There were 27
experiments of this type. In 17 of these experiments, the stimulus type of retrieval
cue was verbal (words), and in 10, it was pictorial (e.g., object pictures and face
photos). The response method featured a standard remember/know procedure
(Gardiner, 2001) in 13 experiments, condence ratings in 7, and a combination of
remember/know and condence ratings in 7. Condence-based experiments were
included because, (1) although there is no one-to-one direct correspondence, on
average, high- and low-condence recognition align more with remember and
know recognition, respectively (Dunn, 2004) and (2) studies have shown that
experiments based on condence ratings yield activation patterns largely com-
parable to those based on the remember/know procedure (Daselaar et al., 2006;
Kim and Cabeza, 2009).
Old/new contrasts involved a comparison of hit responses with correct rejec-
tion responses in a recognition memory test. There were 42 experiments of this
type. In 27 of these experiments, the stimulus type of retrieval cue was verbal, and
in 15, it was pictorial.
Word/pseudoword contrasts involved the presentation of words and pseudo-
words in target and baseline conditions, respectively, and lexical decision tasks.
There were 20 experiments of this type. A word/pseudoword experiment was in-
cluded only when it appropriately controlled for orthographic and phonological
processes. Therefore, the pseudoword condition always involved the presentation
of orthographically legal and pronounceable letter strings. Although the suitability
of a lexical decision task in this study may be questioned because it required a
relatively shallow level of semantic processing, it was included because (1) the
task has been widely used in previous studies and (2) there is behavioral evidence
that the task involves access to the word meaning instead of or in addition to
presemantic word codes (Balota et al., 2004).
In this study, the semantic/phonological contrasts involved semantic and
phonological processing tasks for words in target and baseline conditions, re-
spectively. The semantic processing tasks involved a two-choice response to
questions such as Does the word represent a living or nonliving object? and
Does the word represent a concrete entity or not? and the phonological pro-
cessing tasks had questions such as Does the word have two syllables or not? and
Does the word have the target phoneme?
2.3. Data analysis
Two types of meta-analyses were conducted. The rst type was a single-study
meta-analysis which separately evaluated each of the four task paradigms. The
second type compared pairs of task paradigms. More specically, the evaluation of
differential effects involved a subtraction meta-analysis, and the evaluation of
common effects involved a conjunction meta-analysis. This type of pairwise ana-
lysis proceeded in two stages. The rst stage involved a comparison of two EM task
paradigms (recollection/familiarity and old/new) and a comparison of two SM task
paradigms (word/pseudoword and semantic/phonological). The second stage in-
volved a comparison of the EM task paradigm that indicated stronger default
network engagement in the rst-stage comparison and the SM type contrast that
indicated the same. The rationale for this two-step approach is explained in the
Introduction section and thus is not detailed here.
2.4. Meta-analysis techniques
All meta-analyses were conducted using the activation likelihood estimation
(ALE) algorithm implemented in GingerALE 2.3.3 (http://www.brainmap.org). The
objective of the ALE algorithm is to determine brain regions showing above-chance
activation convergence across a set of independent studies (Turkeltaub et al., 2012).
To apply the algorithm to the present data set, the standard reference space was
determined for each study, and all activation foci reported in the MNI space were
converted into Talairach coordinates by using icbm2tal (Lancaster et al., 2007). Each
activation focus was then modeled as the center of the 3D Gaussian probability
distribution reecting the spatial uncertainty of each focus. The size of the modeled
Gaussian was calculated based on an extended ALE algorithm accounting for be-
tween-subject variability and between-template/institution variability introduced
by different spatial normalization methods. The algorithm included a step assign-
ing a tighter, taller Gaussian to larger samples to reect the notion of spatial
uncertainly being less (for further details, see Eickhoff et al. (2009)). The modeled
probability distributions across all reported foci were combined within a given
study and then across all studies, which ultimately produced voxelwise ALE values
that estimated the activation likelihood of each voxel across studies. The statistical
signicance of ALE scores was determined in reference to the analytically derived
null distribution of the ALE statistic. A cluster-level familywise error-corrected
threshold was set at p o .05, and a cluster-forming threshold was set at the voxel
level at p o.005.
To apply the ALE algorithm to a subtraction analysis of two ALE maps (Eickhoff
et al., 2011), all studies contributing to either ALE map were pooled, the combined
group was randomly split into two sets of studies having the same size as the
original two groups, and the difference in ALE scores between the two sets of
studies was calculated. This process was repeated 10,000 times to estimate the null
distribution of a chance difference between the two ALE maps. This estimated
chance distribution was used to determine the statistical signicance of observed
differences in ALE scores between the two ALE maps. The threshold was set to a
voxel level at p o .005 and inclusively masked by respective main effects with a
cluster-level familywise error-corrected threshold at p o .05. A conjunction analysis
of the two ALE maps was conducted based on the intersection between the two
maps, each with a cluster-level familywise error-corrected threshold set at p o.05.
The possibility of false-positive ndings in subtraction and conjunction analyses
was further constrained by applying a spatial extent threshold exceeding 500 mm3.
Results of the meta-analysis were visualized through the projection of thre-
sholded ALE maps onto an inated population-average landmark surface (PALS) by
using CARET (Van Essen, 2005). Yeo et al. (2011) recently parcellated the cerebral
cortex region into seven intrinsic networks based on a clustering analysis of rest-
ing-state BOLD signals involving 1175 uniformly spaced cortex regions and 1000
subjects (see Fig. 1). Estimated boundaries of the default network in this study were
projected onto the PALS as exible guidelines for evaluating whether convergence
clusters were located within or outside the default network. When appropriate,
estimated boundaries of the fronto-parietal control network were also projected
onto the PALS (see green regions in Fig. 1). For the visualization of results for the
hippocampal formation, ALE maps were overlaid onto the International Consortium
for Brain Mapping template by using Mango (http://ric.uthscsa.edu/mango).
3. Results
3.1. EM retrieval
Table 2 and Fig. 2 show the results of separate ALE meta-ana-
lyses of recollection/familiarity and old/new effects and a direct
comparison of the two effects. Both effects were predominantly
lateralized to the left. First, recollection/familiarity effects over-
lapped mainly and extensively with default network regions, in-
cluding the bilateral posterior cingulate/retrosplenial cortex, the
bilateral inferior parietal lobule, the bilateral anteromedial PFC,
the bilateral medial temporal lobe including the hippocampal
formation, the left ventral PFC, and the bilateral lateral temporal
cortex, in approximate order of decreasing spatial extent (see
Fig. 2A).
Second, old/new effects showed a more moderate overlap with
default network regions, including the bilateral inferior parietal
lobule, the bilateral posterior cingulate cortex, the left lateral
temporal cortex, and the left parahippocampal cortex, in approx-
imate order of decreasing spatial extent (Fig. 2B). However, they
overlapped extensively with the fronto-parietal control network,
including the bilateral middle PFC, the bilateral intraparietal sul-
cus, the bilateral anterior insula, the bilateral dorsal precuneus,
and the left posteromedial PFC, in approximate order of decreasing
spatial extent.
Finally, the regions that were associated more with recollec-
tion/familiarity effects than with old/new effects involved mainly
default network regions, including the bilateral medial temporal
lobe, the bilateral anteromedial PFC, the bilateral inferior parietal
lobule, and the left retrosplenial cortex (yellow regions in Fig. 2C).
By contrast, reverse effects the regions that were associated
more with old/new effects than with recollection/familiarity ef-
fects involved mainly fronto-parietal control regions, including
the left middle PFC, the bilateral intraparietal sulcus, the bilateral
dorsal precuneus, the left anterior insula, and the left poster-
omedial PFC (cyan regions). The regions common to both task
 H. Kim / Neuropsychologia 80 (2016) 3546 39

Table 2
Results of separate ALE meta-analyses of recollection/familiarity and old/new effects and a direct comparison of two effects.

Network Volume (mm3) ALE Z Talairach H Region Brodmann area

x y z

Recollection 4familiarity
DN 8200 .032 8 56 18 B Posterior cingulate cortex, RSC 23, 30, 31
6360 .033 40 70 28 L Inferior parietal lobule 39
5768 .026 6 50 2 B Anteromedial PFC 9, 10, 24, 32
4192 .027 26 20 6 R Hippocampus, PHC 35, 36
3840 .031 28 26 12 L Hippocampus, PHC 35, 36
1816 .019 28 34 4 L Ventral PFC 47
872 .014 46 66 8 R Inferior parietal lobule 39
672 .018 60 28 12 L Lateral temporal cortex 21
552 .016 54 4 14 R Lateral temporal cortex 21

Other 1536 .030 18 2 10 L Lentiform nucleus

1008 .024 4 8 2 R Caudate nucleus
744 .022 52 60 4 L Posterior temporal cortex 37

Old 4new
DN/FPCN 13,600 .059 34 66 40 L Inferior parietal lobule, intraparietal sulcus 7, 19, 39, 40
9664 .056 8 70 28 B Posterior cingulate cortex, dorsal precuneus 7, 31
5472 .038 32 68 36 R Inferior parietal lobule, intraparietal sulcus 7, 19, 39

DN 5504 .046 4 38 32 B Posterior cingulate cortex 23, 31

1328 .028 60 40 8 L Lateral temporal cortex 21
832 .021 16 30 10 L PHC 35

FPCN 14,320 .040 36 46 10 L Middle PFC 6, 9, 10, 45, 46

4032 .035 6 26 42 L Posteromedial PFC 8, 32
3288 .049 30 18 0 L Anterior insula 13
1696 .030 30 20 4 R Anterior insula 13
768 .021 28 52 12 R Middle PFC 10

Other 8824 .049 10 8 6 B Caudate nucleus, lentiform nucleus

640 .022 14 74 10 L Lingual gyrus 18

Recollection/familiarity 4 old/new
DN 3328 3.89 26 20 10 R Hippocampus, PHC 35, 36
2912 3.89 29 30 14 L Hippocampus, PHC 35, 36
1920 3.89 1 56 5 B Anteromedial PFC 10, 32
1512 3.89 47 73 15 L Inferior parietal lobule 39
896 3.89 7 55 14 L RSC 23
624 3.43 46 68 6 R Inferior parietal lobule 39

Other 1200 3.89 20 9 12 L Lentiform nucleus

736 3.89 50 62 1 L Posterior temporal cortex 37

Old/new 4recollection/familiarity
FPCN 5680 3.89 2 72 38 B Dorsal precuneus 7
5128 3.89 33 63 44 L Intraparietal sulcus 7
3992 3.72 40 53 9 L Middle PFC 10, 46
3296 3.89 26 73 42 R Intraparietal sulcus 7, 19
2336 3.72 40 10 34 L Middle PFC 9
1664 3.72 36 15 8 L Anterior insula 13
1240 3.24 3 25 38 L Posteromedial PFC 32

Other 2064 3.89 9 0 11 L Caudate nucleus

Recollection/familiarity old/new
DN 2976 .027 8 46 30 B Posterior cingulate cortex 23, 31
2184 .031 40 70 30 L Inferior parietal lobule 39

ALE, activation likelihood estimation; B, bilateral; DN, default network; FPCN, fronto-parietal control network; H, hemisphere; L, left; PFC, prefrontal cortex; PHC, para-
hippocampal cortex; R, right; RSC, retrosplenial cortex.

paradigms involved only default network regions, including the
bilateral posterior cingulate cortex (more to the left) and the left
inferior parietal lobule (Fig. 2D).
3.2. SM retrieval
Table 3 and Fig. 3 show the results of separate ALE meta-ana-
lyses of word/pseudoword and semantic/phonological effects and
a direct comparison of the two effects. Both effects were
predominantly lateralized to the left. First, word/pseudoword ef-
fects overlapped exclusively and extensively with default network
regions, including the bilateral inferior parietal lobule, the bilateral
posterior cingulate/retrosplenial cortex, the left dorsal PFC, the
bilateral anteromedial PFC, and the left lateral temporal cortex, in
approximate order of decreasing spatial extent (see Fig. 3A).
Second, semantic/phonological effects overlapped mainly with
default network regions, but the scope of this overlap was modest
relative to that of word/pseudoword effects, including the left
40 H. Kim / Neuropsychologia 80 (2016) 3546

Recollection > Familiarity Subtraction

y=-20 y=-25 y=-20 y=-25

ALE .012 .033 Rec/Fam > Old/New Old/New > Rec/Fam

Old > New Conjunction

ALE .014 .059 Rec/Fam Old/New

Fig. 2. Above-threshold regions in an ALE meta-analysis of (A) recollection 4familiarity effects and (B) old4 new effects, (C) a subtraction analysis of two ALE results, and
(D) a conjunction analysis of two results. Red and green lines indicate estimated boundaries of the default network and the front-parietal control network, respectively. For a
detailed explanation of these lines, see Fig. 1. (For interpretation of the references to color in this gure legend, the reader is referred to the web version of this article.)

inferior parietal lobule, the left anteromedial PFC, the left ventral
PFC, the left dorsal PFC, the left lateral temporal cortex, and the left
retrosplenial cortex, in approximate order of decreasing spatial
extent (Fig. 3B).
Finally, the regions that were associated more with word/
pseudoword effects than semantic/phonological effects involved
exclusively default network regions, including the bilateral pos-
terior cingulate cortex and the bilateral inferior parietal lobule
(Fig. 3C). No regions were associated with the reverse effects. The
regions common to both task paradigms involved only default
network regions, including the left inferior parietal lobule and the
left retrosplenial cortex (Fig. 3D).
3.3. EM versus SM retrieval
The results reported earlier indicate greater contributions of
the default network to recollection/familiarity effects than to old/
new effects and to word/pseudoword effects than to semantic/
phonological effects. Therefore, the issue of whether and to what
extent EM and SM processes involve similar or differential default
network subregions was addressed by a direct comparison of re-
collection/familiarity and word/pseudoword effects. Given that
both effects involved a relatively extensive range of default net-
work regions but few others (see Figs. 2A and 3A), an additional
advantage of the comparison was the clear focus on the default
network. The two effects were also similar in the volume of above-
threshold default network regions approximately 32,272 mm3
for recollection/familiarity and 36,664 mm3 for word/pseudoword
(computed from relevant data in Tables 2 and 3) ensuring a re-
latively balanced comparison in terms of the extent of default
network involvement.
Table 4 and Fig. 4 show the results of this comparison. First,
those regions that were associated more with recollection/famil-
iarity effects than with word/pseudoword effects involved only the
bilateral hippocampal formation and (to a lesser extent) the ad-
jacent parahippocampal cortex region (see Fig. 4A). Second, no
regions were associated with reverse effects. Third, the regions
common to both task paradigms were exclusively within the de-
fault network, including the bilateral posterior cingulate cortex/
retrosplenial cortex (more to the left), the left inferior parietal
 H. Kim / Neuropsychologia 80 (2016) 3546 41

Table 3
Results of separate ALE meta-analyses of word/pseudoword and semantic/phonological effects and a direct comparison of two effects.

Network Volume (mm3) ALE Z Talairach H Region Brodmann area

x y z

Word4pseudoword
DN 11,248 .030 38 70 36 L Inferior parietal lobule 19, 39, 40
9048 .024 6 54 26 B Posterior cingulate cortex, RSC 23, 31
5360 .019 24 16 52 L Dorsal PFC 6, 8
5104 .019 48 66 16 R Inferior parietal lobule 39, 40
3560 .022 2 46 4 B Anteromedial PFC 9, 10, 32
2344 .025 60 48 8 L Lateral temporal cortex 21

Semantic 4phonological
DN 4520 .021 42 66 14 L Inferior parietal lobule 39
2112 .013 38 38 10 L Ventral PFC 47
2096 .013 8 50 38 L Anteromedial PFC, dorsal PFC 8, 9
1640 .017 10 48 6 L Anteromedial PFC 32
816 .013 54 24 8 L Lateral temporal cortex 21
736 .012 50 22 0 L Ventral PFC 45
728 .013 24 18 38 L Dorsal PFC 8
696 .010 56 44 0 L Lateral temporal cortex 21
696 .011 6 56 12 L RSC 23

Other 1480 .013 42 22 18 L Fusiform gyrus 20

672 .012 8 10 18 L Caudate nucleus

Word/pseudoword 4semantic/phonological
DN 4264 3.89 4 36 37 B Posterior cingulate cortex 23, 31
2328 3.89 38 51 19 R Inferior parietal lobule 39, 40
2152 3.54 37 73 41 L Inferior parietal lobule 39
1272 3.35 54 54 23 L Inferior parietal lobule 40

Semantic/phonological 4word/pseudoword
(no above-threshold clusters)

Word/pseudoword semantic/phonological
DN 2568 .018 42 66 16 L Inferior parietal lobule 39
576 .011 6 56 12 L RSC 23

For abbreviations, see Table 2.

lobule, and the left anteromedial PFC (Fig. 4B). In sum, both effects
involved a relatively extensive range of default network regions,
and these regions overlapped to a large extent. However, the re-
cruitment of the hippocampus and parahippocampal region was
unique to recollection/familiarity effects.
4. Discussion
4.1. EM retrieval and the default network
Recollection/familiarity effects overlapped extensively with the
default network, whereas old/new effects, modestly. According to
a direct comparison of the two effects, many default network re-
gions were associated more with recollection/familiarity effects.
Robust contributions of the default network to recollection/fa-
miliarity effects provide direct evidence that recollection involves
the default network more consistently than familiarity. This evi-
dence in turn suggests that the lower involvement of the default
network in old/new effects is tied to the inclusion of familiarity-
based recognition in the target condition. As mentioned earlier,
experiments based on condence ratings yielded activation pat-
terns virtually identical to those based on the remember/know
procedure (Daselaar et al., 2006; Kim and Cabeza, 2009). In this
regard, familiarity recognition typically involves low recognition
condence and (by inference) weak memory signals. This hy-
pothesis resonates with the view that the remember-know pro-
cedure, as it is ordinarily used, may measure memory strength
more than it distinguishes between recollection and familiarity
(Dunn, 2004; Wixted and Mickes, 2010). Along these lines, some
EM (and SM) task paradigms involving only modest default net-
work contributions, including old/new, may involve relatively
weak memory signals in target conditions. The absence of any
hippocampal region associated with old/new effects may also re-
ect some masking by the stronger encoding-related activity of
the region for novel items than for old ones (Stark and Okado,
2003). Old/new effects overlapped more with fronto-parietal
control regions, providing support for the hypothesis that famil-
iarity and associated low recognition condence entail a strong
demand for controlled retrieval processes (Henson et al., 1999;
Kim and Cabeza, 2009; Yonelinas et al., 2005).
This study's results and the ndings of previous meta-analyses
(Kim, 2013; Spaniol et al., 2009) indicate that old/new effects are
only modestly associated with the default network. This merits
further comments in that it seemingly goes against the hypothesis
that EM retrieval processes are critically related to the default
network. First, as mentioned earlier and in conjunction with the
fact that old/new effects include familiarity-based recognition in
target conditions, memory signals during familiarity-based re-
cognition may not be strong enough to activate the default net-
work. Second, low-condence memory judgements generally de-
mand more controlled processing resources than high-condence
ones (Henson et al., 2000), and studies have shown that the task-
induced deactivation of the default network increases as the task
demand increases (McKiernan et al., 2003, 2006). Along this line,
familiarity-based recognition may induce the default network's
relatively strong deactivation. Finally, new responses may be
based on a recall-to-reject process in which mismatching
42 H. Kim / Neuropsychologia 80 (2016) 3546

Word > Pseudoword Subtraction

Word/Pseudo > Sem/Pho

ALE .010 .030
Sem/Pho > Word/Pseudo (none)

Semantic > Phonological Conjunction

ALE .008 .021 Word/Pseudo Sem/Pho

Fig. 3. Above-threshold regions in an ALE meta-analysis of (A) word4pseudoword effects and (B) semantic 4 phonological effects, (C) a subtraction analysis of two ALE
results, and (D) a conjunction analysis of two results. Red lines indicate estimated boundaries of the default network (see Fig. 1). (For interpretation of the references to color
in this gure legend, the reader is referred to the web version of this article.)

information that is retrieved from memory is used to reject test
foils that are similar to studied items (Rotello et al., 2000, p. 67).
In this regard, reference conditions of old/new effects may involve,
at least in some conditions, memory signals that activate the de-
fault network to some extent.
McDermott et al.'s (2009) meta-analysis compared activated
regions in studies of laboratory-based EM retrieval versus auto-
biographical memory retrieval and reported very few overlapping
regions, suggesting that the two paradigms may typically probe
very different retrieval-related processes. However, studies in-
cluded in the laboratory-based analysis used mainly old/new task
paradigms. That is, if a meta-analysis compares recollection/fa-
miliarity to autobiographical tasks, then results may indicate more
overlaps involving default network regions. In conjunction with
the fact that old/new effects, not recollection/familiarity effects,
involve familiarity-based recognition in target conditions, this line
of reasoning indicates that laboratory-based recollection, not fa-
miliarity, shares relatively extensive processes with

Table 4
Results of a direct comparison of recollection/familiarity and word/pseudoword effects.

Network Volume (mm3) ALE Z Talairach H Region Brodmann area

x y z

Recollection/familiarity 4 word/pseudoword
DN 2616 3.89 22 19 11 R Hippocampus, PHC 35
744 3.72 27 21 7 L Hippocampus

Word/pseudoword4 recollection/familiarity
(No above-threshold clusters)

Recollection/familiarity word/pseudoword
DN 4984 .025 42 68 24 L Inferior parietal lobule 39
4904 .023 10 50 28 B Posterior cingulate cortex, RSC 23, 31
2576 .020 2 48 4 L Anteromedial PFC 10, 32

For abbreviations, see Table 2.

 H. Kim / Neuropsychologia 80 (2016) 3546
Subtraction
y=-20 y=-25
Rec/Fam > Word/Pseudo
Word/Pseudo > Rec/Fam (none)
Conjunction
Rec/Fam Word/Pseudo
Fig. 4. Above-threshold regions in (A) a subtraction analysis of
recollection4familiarity effects (EM retrieval) and word 4pseudoword effects (SM
retrieval) and (B) a conjunction analysis of two effects. Red lines indicate estimated
boundaries of the default network (see Fig. 1). (For interpretation of the references
to color in this gure legend, the reader is referred to the web version of this
article.)
autobiographical retrieval. This is perhaps not surprising in that,
although the distinction between recollection and familiarity is
seldom made explicit in studies of autobiographical retrieval, most
cases of retrieval within the paradigm likely involve recollection.
4.2. SM retrieval and the default network
Both word/pseudoword and semantic/phonological effects in-
volved mainly default network regions, but this involvement was
much more extensive for word/pseudoword effects. According to a
direct comparison of the two effects, many default network re-
gions were associated more with word/pseudoword effects. Sub-
stantial contributions of the default network to word/pseudoword
effects likely reect access to meaning in the word, not pseudo-
word, condition. Behavioral studies have provided strong evidence
that the mere presentation of words typically activates semantic
processes even when subjects are not required to do so (Lesch and
Pollatsek, 1993; Lucas, 2000; Shaffer and LaBerge, 1979). In this
regard, the lower involvement of the default network in semantic/
phonological effects may reect relatively automatic semantic
processing in the phonological condition and the associated partial
subtraction of semantic activation (Price et al., 1996). Consistent
with this hypothesis, previous studies contrasting a semantic task
involving words with a phonological task involving pseudowords
have indicated more consistent default network involvement
(Binder et al., 2008; Cappa et al., 1998; Dmonet et al., 1992). If
relatively easy phonological tasks are used, then the modest in-
volvement of the default network in semantic/phonological effects
may additionally reect the lesser task-induced deactivation of the
default network in reference conditions (McKiernan et al., 2003,
43
2006). If this line of reasoning is correct, then some SM (and EM)
task paradigms involving only modest default network contribu-
tions, including semantic/phonological, may involve relatively
strong memory signals in reference conditions. Many SM tasks
emphasize the retrieval of overlearned information such as the
lexicality and meaning of common words and (by inference)
strong memory signals. In such SM tasks, conclusions about which
task paradigms do or do not engage the default network to a large
extent may be dependent mainly on the strength of memory sig-
nals in the reference task.
Seemingly disparate cognitive functions engage the default
network, including future thinking, remembering, conceiving the
perspective of others (theory of mind), and navigation (Spreng
et al., 2009). A well-known theory (Buckner and Carroll, 2007)
proposes that mentally projecting oneself into an alternative si-
tuation may be a key process common to these functions. A dif-
ferent view (Hassabis and Maguire, 2007) suggests that mental
scene construction is a more fundamental common process. Yet
another view (Schacter et al., 2007) holds that these functions
commonly involve the extraction and recombination of relevant
episodic details from stored memories. On the one hand, the
lexicality decision (or word recognition) is not explicitly linked to
self-projection, mental scene construction, or constructive episo-
dic simulation, which suggests that any such process are sufcient
but not required for default network activation. On the other hand,
most (if not all) instantiations of these processes necessarily in-
clude some semantic representation. Therefore, as suggested in
Binder et al. (2009), the default network may be more funda-
mentally portrayed as a semantic system.
4.3. EM versus SM retrieval and the default network
This study's results as well as previous ndings indicate that
various episodic and semantic task paradigms differ widely in the
extent of default network involvement. Therefore, a theoretically
more meaningful comparison of default network subregions in-
volved in EM versus SM processes requires this factor to be con-
trolled for to some extent. Along this line, the present study
compares recollection/familiarity effects (EM retrieval) and word/
pseudoword effects (SM retrieval), each involving a range of de-
fault network regions and relatively few other regions. According
to the results, the two effects had more overlaps than separations
in default network subregions involved in the two effects. More
specically, overlaps included the bilateral posterior cingulate/
retrosplenial cortex, the left inferior parietal lobule, and the left
anteromedial PFC region, whereas separations included only the
hippocampal formation and (to a lesser extent) the adjacent
parahippocampal cortex region, which was unique to recollection/
familiarity effects. This implies that EM and SM retrieval processes
involving strong memory signals recruit extensive and largely
overlapping default network regions and differ mainly in distinct
contributions of hippocampus and parahippocampal regions to EM
retrieval.
Noteworthy is the broad overlap of recollection/familiarity and
word/pseudoword effects in the default network. First, the overlap
goes beyond the unequivocal but banal observation that all EM
retrieval processes include SM representations because language
requirements are held constant in target and reference conditions
of recollection/familiarity contrasts. Second, it highlights the
within-component anatomical specicity of both effects. That is,
both involved the inferior parietal lobule component more pos-
teriorly (i.e., the angular gyrus), the posterior cingulate/retro-
splenial cortex component more ventrally, and the anteromedial
PFC component more ventrally. Third, the broad overlap involving
the default network resonates with the recently proposed semantic
scaffolding hypothesis, which suggests that semantic knowledge
44 H. Kim / Neuropsychologia 80 (2016) 3546
provides a framework or scaffolding that facilitates most (if not all)
forms of episodic remembering and future thinking (Greenberg
and Verfaellie, 2010; Irish and Piguet, 2013). Finally, the results
raise the question of what type of common process is recruited
during episodic recollection and word recognition. Here a plau-
sible hypothesis is that strong memory signals, both episodic and
semantic, may typically capture a subject's attention away from
the visual stimulus (i.e., perceptual decoupling) and drive toward
internal information. This hypothesis is consistent with the evi-
dence that the default network supports internally oriented
mentation (AndrewsHanna et al., 2014) and may provide some
testable predictions.
As discussed in the Introduction section, previous within-study
comparisons of laboratory-based EM and SM tasks have typically
indicated limited overlaps involving the default network. How-
ever, such studies have employed EM and/or SM task paradigms
involving relatively weak contributions of the default network,
including old/new and semantic/phonological contrasts. A smaller
set of previous studies has employed the autobiographical mem-
ory paradigm. For example, Maguire and Frith (2003) compared a
condition involving true/false judgments about autobiographical
event statements (e.g., You were the Christmas star in the school
nativity play) and a condition involving true/false judgments
about autobiographical fact statements (e.g., Winkey and Frawley
were friends at school). Such studies (e.g., Addis et al., 2004;
Holland et al., 2011; Levine et al., 2004; Maguire and Frith, 2003;
Zysset et al., 2002) have generally indicated that autobiographical
EM and SM tasks both involve an extensive range of default net-
work regions and that these default network regions show some
broad overlaps. Here the interpretive ambiguity concerns whether
and to what extent such broad overlaps reect common self-re-
ferential components or more memory-specic commonalities.
The results of the present study demonstrate that at least some
commonalities are due to memory-specic processes by showing
that a similar overlap can be observed even when self-referential
processing is not a common component. The study also extends
the literature on autobiographical memory by showing that epi-
sodic and semantic processing tasks at even very low levels can
yield similar overlaps and separations.
The bilateral hippocampal formation and the adjacent para-
hippocampal region were largely specic to recollection/famil-
iarity effects, providing support for the hypothesis that these re-
gions play a more pivotal role in EM retrieval than in SM retrieval
(Nadel et al., 2000; Tulving and Markowitsch, 1998; Vargha-Kha-
dem et al., 1997). Previous within-study comparisons of auto-
biographical EM retrieval versus autobiographical SM retrieval
have frequently indicated greater contributions of the hippo-
campal region to EM retrieval (Holland et al., 2011; Levine et al.,
2004; Maguire and Frith, 2003). Studies of patients with hippo-
campal damage have generally indicated some severe decit in
clinical measures of EM processes but largely intact SM processes
based on standardized tests (e.g., Parkin and Leng, 1993). There-
fore, the hippocampus may be the key neural structure under-
pinning some distinct characteristics of EM relative to SM, such as
the retrieval of detailed contextual information (Yonelinas, 2013),
scene construction (Maguire and Mullally, 2013), and a sense of
reliving or autonoetic consciousness (Klein, 2013; Tulving and
Markowitsch, 1998). Although word/pseudoword effects involved
no hippocampal or other medial temporal lobe activation, pre-
vious studies have shown that more complex SM tasks such as
autobiographical fact retrieval, the retrieval of public facts, and
semantic uency can recruit some hippocampal regions (Maguire
and Frith, 2003; Sheldon and Moscovitch, 2012). Therefore, the
hippocampal contribution to EM versus SM may be a relative
difference instead of a sharp dichotomy. Hippocampal contribu-
tions to SM tasks may reect the fact that some SM retrieval
involves elements of contextual information, scene construction,
and even a sense of reliving (Rubin and Umanath, 2015) as well as
an episodic retrieval strategy during SM tasks.
A competing theory to the hypothesis that the hippocampus
plays a more important role in EM retrieval than SM retrieval is
the consolidation model of memory (Squire et al., 2004), which
posits that both EM and SM are relatively equally dependent on
the hippocampal region and that it plays a time-limited role in
both types of memory. The hippocampal region supports the
maintenance and retrieval of both EM and SM but only until
memory consolidation is complete. Proponents of this view have
obtained supporting evidence from both neuroimaging and lesions
studies, including evidence that hippocampal activity during SM
retrieval is related to the age of memory (Manns et al., 2003;
Smith and Squire, 2009). The consolidation model of memory can
account for greater hippocampal involvement in recollection/fa-
miliarity than word/pseudoword effects because laboratory-based
recollection involves newly acquired memories, whereas word
recognition, remotely acquired memories. In this regard, it remains
unclear whether and to what extent differential hippocampal in-
volvement reects memory type or age or both. One related con-
cern is that if memory consolidation necessarily (or typically) in-
volves a transformation of episodic memory to semantic memory
(Winocur and Moscovitch, 2011), the question of whether the
hippocampus supports episodic memory or newly acquired
memory becomes a denitional, not empirical, issue.
According to the results, no default network (and other) region
was associated more with word/pseudoword effects than with
recollection/familiarity effects. It is difcult to argue strongly from
negative evidence, and the results may reect the specic task
paradigms included in the study. However, previous within-study
comparisons of autobiographical EM retrieval and auto-
biographical SM retrieval have rarely indicated any default net-
work region being activated more by the SM condition than by the
EM condition. In this regard, the present study as well as previous
studies indicates dissociation involving EM but not that involving
SM. A single case of dissociation involving EM provides support for
Tulvings (1989) hierarchical view of memory organization, which
suggests that the episodic system depends on but goes beyond
the capabilities of the semantic system. The semantic system, on
the other hand, is capable of operating in the absence, or in-
dependently, of the episodic system (p. 362). As suggested in
Barba et al. (1998), this hierarchical organization requires that
episodic memory involves all regions contributing to semantic
memory and some additional regions not contributing to semantic
memory.
4.4. Methodological considerations and conclusions
A possible criticism of the present selective comparison is
that the broader inclusion of task paradigms in each domain may
yield more generalizable and possibly more useful results. If a
study's aim is only the numerical integration of results across a
large number of studies with relatively few theoretical concerns,
then a more inclusive approach may be more suitable. However,
given that the present study also pursues a theoretical clarication
of previous ndings, there were some reasons behind this study's
choice of a more selective approach. First, a more inclusive com-
parison may yield less interpretable results because different task
paradigms within each domain are often associated with strikingly
heterogeneous effects (e.g., old/new vs. recollection/familiarity). To
illustrate this point, if a comparison involves old/new effects (EM)
and word/pseudoword effects (SM), then results may probably
(but not very plausibly) indicate that EM processes involve mainly
fronto-parietal control network regions, whereas SM processes,
mainly default network regions. Such nontrivial confounding and
 H. Kim / Neuropsychologia 80 (2016) 3546
interpretive ambiguity may not necessarily be mitigated by in-
cluding more paradigms in the analysis because each domain is
now represented by multiple and often markedly dissimilar
effects.
Second, the ultimate aim of this study is to determine whether
and the extent to which the EM and SM processes involve similar
or differential default network subregions. This issue may be
better addressed using EM and SM task paradigms involving re-
latively extensive default network involvement, such as recollec-
tion/familiarity and word/pseudoword, than using those involving
only modest default network involvement, such as old/new and
semantic/phonological. In addition, in most cases, an ideal mea-
sure of both episodic and semantic memory processes involves
high memory signals in target conditions and low or no memory
signals in baseline conditions. As discussed earlier, both old/new
and semantic/phonological task paradigms are likely to fail this
criterion to a nontrivial extent, and their inclusion is likely to make
results less interpretable in exchange for supercially higher but
not necessarily more meaningful generalizability. Taken together,
these considerations suggest that a selective comparison involving
recollection/familiarity and word/pseudoword task paradigms but
not the other two task paradigms may provide better control for
biasing factors and more interpretable and useful results.
Of course, it is true that this study's results are generalizable
only to the two selected task paradigms and thus not necessarily
to other EM and SM task paradigms. Therefore, any generalization
based on this study's results should be made with caution. The
foregoing discussion provides some support for the hypothesis
that recollection/familiarity and word/pseudoword task paradigms
but not old/new and semantic/phonological ones involve both
strong memory signals in target conditions and weak memory
signals in baseline conditions. If this argument is correct at least to
some extent, then this study's results may be generalized not only
to the two selected task paradigms but also to other EM and SM
task paradigms involving strong memory signals in target condi-
tions and weak memory signals in reference conditions. In this
regard, it is worth noting again that previous within-study com-
parisons of autobiographical EM and SM retrieval processes have
generally indicated that they both involve an extensive range of
default network regions and that these default network regions
show extensive overlaps (e.g., Addis et al., 2004; Holland et al.,
2011; Levine et al., 2004; Maguire and Frith, 2003; Zysset et al.,
2002). These results provide evidence that what can be inferred
from this study's results is not strictly limited to the two selected
paradigms but also to some EM and SM task paradigms involving
strong memory signals.
In conclusion, the present comparison of recollection/famil-
iarity and word/pseudoword effects provides evidence that EM
and SM retrieval processes involving strong memory signals re-
cruit extensive and largely overlapping default network regions
and differ mainly in distinct contributions of hippocampus and
parahippocampal regions to EM retrieval. The broad overlap pro-
vides support for the notion that SM retrieval is a logical pre-
requisite for EM retrieval and also for a more formal but associated
view that semantic knowledge provides scaffolding for EM re-
trieval (Greenberg and Verfaellie, 2010; Irish and Piguet, 2013).
The greater association of the hippocampal formation with re-
collection/familiarity effects than with word/pseudoword effects is
consistent with its presumed pivotal role in EM, not SM, retrieval,
although it remains unclear whether it may alternatively (or ad-
ditionally) reect a greater role of the hippocampus in the retrieval
of newly acquired memories than in the retrieval of remote and
consolidated memories. More broadly, the study provides evi-
dence that neural correlates of EM and SM processes show more
overlaps than separations. This evidence is more compatible with
the view that a single unied memory system supports all
45
declarative memory retrieval processes than with the view that
two spatially segregated and independent memory systems sup-
port EM and SM processes. However, it should be noted that by
focusing exclusively on default network regions and thus on strong
memory signals, the present study ignores potential EM and SM
differences involving other brain regions that may support more
controlled and other retrieval-related processes. In this regard, an
important area of future research is to compare EM and SM re-
trieval processes in terms of not only retrieval success but also
other components such as modes, effort, and orientation (Rugg
and Wilding, 2000).
Acknowledgments
This research was supported by the Daegu University Research
Grant in 2014.
Appendix A. Supplementary material
Supplementary data associated with this article can be found in
the online version at http://dx.doi.org/10.1016/j.neuropsychologia.
2015.11.006.
References
Addis, D.R., Moscovitch, M., Crawley, A.P., McAndrews, M.P., 2004. Recollective
qualities modulate hippocampal activation during autobiographical memory
retrieval. Hippocampus 14, 752762.
AndrewsHanna, J.R., Smallwood, J., Spreng, R.N., 2014. The default network and
selfgenerated thought: component processes, dynamic control, and clinical
relevance. Ann. N Y. Acad. Sci. 1316, 2952.
Balota, D.A., Cortese, M.J., Sergent-Marshall, S.D., Spieler, D.H., Yap, M., 2004. Visual
word recognition of single-syllable words. J. Exp. Psychol. Gen. 133, 283316.
Barba, G.D., Parlato, V., Jobert, A., Samson, Y., Pappata, S., 1998. Cortical networks
implicated in semantic and episodic memory: common or unique? Cortex 34,
547561.
Binder, J.R., Desai, R.H., Graves, W.W., Conant, L.L., 2009. Where is the semantic
system? A critical review and meta-analysis of 120 functional neuroimaging
studies. Cereb. Cortex 19, 27672796.
Binder, J.R., Swanson, S.J., Hammeke, T.A., Sabsevitz, D.S., 2008. A comparison of ve
fMRI protocols for mapping speech comprehension systems. Epilepsia 49,
19801997.
Buckner, R., Carroll, D., 2007. Self-projection and the brain. Trends Cogn. Sci. 11,
4957.
Burianova, H., Grady, C.L., 2007. Common and unique neural activations in auto-
biographical episodic and semantic retrieval. J. Cogn. Neurosci. 19, 15201534.
Cappa, S., Perani, D., Schnur, T., Tettamanti, M., Fazio, F., 1998. The effects of se-
mantic category and knowledge type on lexical-semantic access: a PET study.
Neuroimage 8, 350359.
Daselaar, S.M., Fleck, M.S., Cabeza, R., 2006. Triple dissociation in the medial tem-
poral lobes: recollection, familiarity, and novelty. J. Neurophysiol. 96,
19021911.
Dmonet, J.F., Chollet, F., Ramsay, S., Cardebat, D., Nespoulous, J.L., Wise, R., Rascol,
A., Frackowiak, R., 1992. The anatomy of phonological and semantic processing
in normal subjects. Brain 115, 17531768.
Dzel, E., Cabeza, R., Picton, T.W., Yonelinas, A.P., Scheich, H., Heinze, H.-J., Tulving,
E., 1999. Task-related and item-related brain processes of memory retrieval.
Proc. Natl. Acad. Sci. USA 96, 17941799.
Dunn, J., 2004. Remember-know: a matter of condence. Psychol. Rev. 111,
524542.
Eickhoff, S.B., Bzdok, D., Laird, A.R., Roski, C., Caspers, S., Zilles, K., Fox, P.T., 2011. Co-
activation patterns distinguish cortical modules, their connectivity and func-
tional differentiation. Neuroimage 57, 938949.
Eickhoff, S.B., Laird, A.R., Grefkes, C., Wang, L.E., Zilles, K., Fox, P.T., 2009. Coordinate-
based activation likelihood estimation meta-analysis of neuroimaging data: a
random-effects approach based on empirical estimates of spatial uncertainty.
Hum. Brain Mapp. 30, 29072926.
Emmorey, K., Weisberg, J., McCullough, S., Petrich, J.A., 2013. Mapping the reading
circuitry for skilled deaf readers: an fMRI study of semantic and phonological
processing. Brain Lang. 126, 169180.
Gardiner, J.M., 2001. Episodic memory and autonoetic consciousness: a rst-person
approach. Philos. Trans. R. Soc. Lond. B: Biol. Sci. 356, 13511361.
Greenberg, D.L., Verfaellie, M., 2010. Interdependence of episodic and semantic
memory: evidence from neuropsychology. J. Int. Neuropsychol. Soc. 16,
46 H. Kim / Neuropsychologia 80 (2016) 3546
748753.
Hantke, N., Nielson, K.A., Woodard, J.L., Breting, L.M.G., Butts, A., Seidenberg, M.,
Carson Smith, J., Durgerian, S., Lancaster, M., Matthews, M., 2013. Comparison
of semantic and episodic memory BOLD fMRI activation in predicting cognitive
decline in older adults. J. Int. Neuropsychol. Soc. 19, 1121.
Hassabis, D., Maguire, E., 2007. Deconstructing episodic memory with construction.
Trends Cogn. Sci. 11, 299306.
Henson, R., Rugg, M., Shallice, T., Dolan, R., 2000. Condence in recognition memory
for words: dissociating right prefrontal roles in episodic retrieval. J. Cogn.
Neurosci. 12, 913923.
Henson, R.N.A., Rugg, M.D., Shallice, T., Josephs, O., Dolan, R.J., 1999. Recollection
and familiarity in recognition memory: an event-related functional magnetic
resonance imaging study. J. Neurosci. 19, 39623972.
Holland, A.C., Addis, D.R., Kensinger, E.A., 2011. The neural correlates of specic
versus general autobiographical memory construction and elaboration. Neu-
ropsychologia 49, 31643177.
Irish, M., Piguet, O., 2013. The pivotal role of semantic memory in remembering the
past and imagining the future. Front. Behav. Neurosci. 7, 27.
Kim, H., 2010. Dissociating the roles of the default-mode, dorsal, and ventral net-
works in episodic memory retrieval. Neuroimage 50, 16481657.
Kim, H., 2012. A dual-subsystem model of the brains default network: Self-refer-
ential processing, memory retrieval processes, and autobiographical memory
retrieval. Neuroimage 61, 966977.
Kim, H., 2013. Differential neural activity in the recognition of old versus new
events: an activation likelihood estimation meta-analysis. Hum. Brain Mapp.
34, 814836.
Kim, H., 2015. Encoding and retrieval along the long axis of the hippocampus and
their relationships with dorsal attention and default mode networks: the
HERNET model. Hippocampus 25, 500510.
Kim, H., Cabeza, R., 2009. Common and specic brain regions in high- versus low-
condence recognition memory. Brain Res. 1282, 103113.
Klein, S.B., 2013. Making the case that episodic recollection is attributable to op-
erations occurring at retrieval rather than to content stored in a dedicated
subsystem of long-term memory. Front. Behav. Neurosci. 7 (3), 114.
Lancaster, J., Tordesillas-Gutierrez, D., Martinez, M., Salinas, F., Evans, A., Zilles, K.,
Mazziotta, J., Fox, P., 2007. Bias between MNI and Talairach coordinates ana-
lyzed using the ICBM-152 brain template. Hum. Brain Mapp. 28, 11941205.
Lesch, M.F., Pollatsek, A., 1993. Automatic access of semantic information by pho-
nological codes in visual word recognition. J. Exp. Psychol. Learn. Mem. Cogn.
19, 285294.
Levine, B., Turner, G.R., Tisserand, D., Hevenor, S.J., Graham, S.J., McIntosh, A.R.,
2004. The functional neuroanatomy of episodic and semantic autobiographical
remembering: a prospective functional MRI study. J. Cogn. Neurosci. 16,
16331646.
Lucas, M., 2000. Semantic priming without association: a meta-analytic review.
Psychon. Bull. Rev. 7, 618630.
Maguire, E.A., Frith, C.D., 2003. Aging affects the engagement of the hippocampus
during autobiographical memory retrieval. Brain 126, 15111523.
Maguire, E.A., Mullally, S.L., 2013. The hippocampus: a manifesto for change. J. Exp.
Psychol. Gen. 142, 11801189.
Maguire, E.A., Mummery, C.J., 1999. Differential modulation of a common memory
retrieval network revealed by positron emission tomography. Hippocampus 9,
5461.
Manns, J.R., Hopkins, R.O., Squire, L.R., 2003. Semantic memory and the human
hippocampus. Neuron 38, 127133.
McDermott, K.B., Szpunar, K.K., Christ, S.E., 2009. Laboratory-based and auto-
biographical retrieval tasks differ substantially in their neural substrates.
Neuropsychologia 47, 22902298.
McKiernan, K., DAngelo, B., Kaufman, J., Binder, J., 2006. Interrupting the stream of
consciousness: an fMRI investigation. Neuroimage 29, 11851191.
McKiernan, K., Kaufman, J., Kucera-Thompson, J., Binder, J., 2003. A parametric
manipulation of factors affecting task-induced deactivation in functional neu-
roimaging. J. Cogn. Neurosci. 15, 394408.
Nadel, L., Samsonovich, A., Ryan, L., Moscovitch, M., 2000. Multiple trace theory of
human memory: computational, neuroimaging, and neuropsychological re-
sults. Hippocampus 10, 352368.
Neisser, U., 1981. John Deans memory: a case study. Cognition 9, 122.
Nyberg, L., Marklund, P., Persson, J., Cabeza, R., Forkstam, C., Petersson, K.M., Ingvar,
M., 2003. Common prefrontal activations during working memory, episodic
memory and semantic memory. Neuropsychologia 41, 371377.
Parkin, A.J., Leng, N.R.C., 1993. Neuropsychology of the Amnestic Syndrome.
Lawrence Erlbaum Associates, Hove, UK.
Price, C.J., Moore, C., Humphreys, G., Wise, R., 1997. Segregating semantic from
phonological processes during reading. J. Cogn. Neurosci. 9, 727733.
Price, C.J., Wise, R., Frackowiak, R., 1996. Demonstrating the implicit processing of
visually presented words and pseudowords. Cereb. Cortex 6, 6270.
Raichle, M., MacLeod, A., Snyder, A., Powers, W., Gusnard, D., Shulman, G., 2001. A
default mode of brain function. Proc. Natl. Acad. Sci. USA 98, 676682.
Rajah, M.N., McIntosh, A.R., 2005. Overlap in the functional neural systems involved
in semantic and episodic memory retrieval. J. Cogn. Neurosci. 17, 470482.
Renoult, L., Davidson, P.S., Palombo, D.J., Moscovitch, M., Levine, B., 2012. Personal
semantics: at the crossroads of semantic and episodic memory. Trends Cogn.
Sci. 16, 550558.
Rotello, C.M., Macmillan, N.A., van Tassel, G., 2000. Recall-to-reject in recognition:
evidence from ROC curves. J. Mem. Lang. 43, 6788.
Rubin, D.C., Umanath, S., 2015. Event memory: a theory of memory for laboratory,
autobiographical, and ctional events. Psychol. Rev. 122, 123.
Rugg, M., Wilding, E., 2000. Retrieval processing and episodic memory. Trends
Cogn. Sci. 4, 108115.
Ryan, L., Cox, C., Hayes, S.M., Nadel, L., 2008. Hippocampal activation during epi-
sodic and semantic memory retrieval: comparing category production and
category cued recall. Neuropsychologia 46, 21092121.
Schacter, D.L., Addis, D.R., Buckner, R.L., 2007. Remembering the past to imagine the
future: the prospective brain. Nat. Rev. Neurosci. 8, 657661.
Shaffer, W.O., LaBerge, D., 1979. Automatic semantic processing of unattended
words. J. Verbal Learn. Verbal Behav. 18, 413426.
Shapira-Lichter, I., Oren, N., Jacob, Y., Gruberger, M., Hendler, T., 2013. Portraying
the unique contribution of the default mode network to internally driven
mnemonic processes. Proc. Natl. Acad. Sci. USA 110, 49504955.
Sheldon, S., Moscovitch, M., 2012. The nature and time-course of medial temporal
lobe contributions to semantic retrieval: an fMRI study on verbal uency.
Hippocampus 22, 14511466.
Smith, C.N., Squire, L.R., 2009. Medial temporal lobe activity during retrieval of
semantic memory is related to the age of the memory. J. Neurosci. 29, 930938.
Spaniol, J., Davidson, P., Kim, A., Han, H., Moscovitch, M., Grady, C., 2009. Event-
related fMRI studies of episodic encoding and retrieval: meta-analyses using
activation likelihood estimation. Neuropsychologia 47, 17651779.
Spreng, R., Mar, R., Kim, A., 2009. The common neural basis of autobiographical
memory, prospection, navigation, theory of mind, and the default mode: a
quantitative meta-analysis. J. Cogn. Neurosci. 21, 489510.
Squire, L.R., Stark, C., Clark, R., 2004. The medial temporal lobe. Annu. Rev. Neurosci.
27, 279306.
Stark, C., Okado, Y., 2003. Making memories without trying: medial temporal lobe
activity associated with incidental memory formation during recognition. J.
Neurosci. 23, 67486753.
St-Laurent, M., Abdi, H., Burianov, H., Grady, C.L., 2011. Inuence of aging on the
neural correlates of autobiographical episodic and semantic memory retrieval.
J. Cogn. Neurosci. 23, 41504163.
Tulving, E., 1972. Episodic and semantic memory. In: Tulving, E., Donaldson, W.
(Eds.), Organization of Memory. Academic Press, Oxford, pp. 381402.
Tulving, E., 1989. Remembering and knowing the past. Am. Sci. 77, 361367.
Tulving, E., Markowitsch, H.J., 1998. Episodic and declarative memory: role of the
hippocampus. Hippocampus 8, 198204.
Turkeltaub, P.E., Eickhoff, S.B., Laird, A.R., Fox, M., Wiener, M., Fox, P., 2012. Mini-
mizing withinexperiment and withingroup effects in activation likelihood
estimation metaanalyses. Hum. Brain Mapp. 33, 113.
Van Essen, D., 2005. A population-average landmark- and surface-based (PALS)
atlas of human cerebral cortex. Neuroimage 28, 635662.
Vargha-Khadem, F., Gadian, D.G., Watkins, K.E., Connelly, A., Van Paesschen, W.,
Mishkin, M., 1997. Differential effects of early hippocampal pathology on epi-
sodic and semantic memory. Science 277, 376380.
Wiggs, C.L., Weisberg, J., Martin, A., 1998. Neural correlates of semantic and epi-
sodic memory retrieval. Neuropsychologia 37, 103118.
Winocur, G., Moscovitch, M., 2011. Memory transformation and systems con-
solidation. J. Int. Neuropsychol. Soc. 17, 766780.
Wixted, J.T., Mickes, L., 2010. A continuous dual-process model of remember/know
judgments. Psychol. Rev. 117, 10251054.
Yeo, B.T.T., Krienen, F.M., Sepulcre, J., Sabuncu, M.R., Lashkari, D., Hollinshead, M.,
Roffman, J.L., Smoller, J.W., Zllei, L., Polimeni, J.R., 2011. The organization of the
human cerebral cortex estimated by intrinsic functional connectivity. J. Neu-
rophysiol. 106, 11251165.
Yonelinas, A.P., 2013. The hippocampus supports high-resolution binding in the
service of perception, working memory and long-term memory. Behav. Brain
Res. 254, 3444.
Yonelinas, A.P., Otten, L.J., Shaw, K.N., Rugg, M.D., 2005. Separating the brain regions
involved in recollection and familiarity in recognition memory. J. Neurosci. 25,
30023008.
Zysset, S., Huber, O., Ferstl, E., Von Cramon, D., 2002. The anterior frontomedian
cortex and evaluative judgment: an fMRI study. Neuroimage 15, 983991.