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Psychomusicology: Music, Mind, and Brain

© 2018 American Psychological Association 2018, Vol. 28, No. 2, 82–100


0275-3987/18/$12.00 http://dx.doi.org/10.1037/pmu0000210

Listening for Memories: Attentional Focus Dissociates Functional Brain


Networks Engaged by Memory-Evoking Music
Benjamin Kubit and Petr Janata
University of California, Davis

How we experience music depends largely on how we orient our attention. For example, we can orient
attention toward aspects of the music or toward thoughts that are evoked while listening. Often, those
thoughts are memories that are associated with the music. To better understand how task set and the
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

associated attentional focus recruit brain networks during attentive music listening, we performed two
This document is copyrighted by the American Psychological Association or one of its allied publishers.

experiments: (a) a single-factor behavioral experiment manipulating Attentional Focus and (b) a balanced
2 ⫻ 2 factorial design functional MRI experiment crossing Attentional Focus with Memory Status.
Before listening to 30-s song excerpts, half of which were known to be memory evoking in the functional
MRI experiment, participants were instructed to either focus on the retrieval of memories evoked by the
music, or to determine how many instruments were playing. Attentional Focus accounted for the majority
of variability in brain activity and modulated both functional connectivity and community structure of
brain regions during music listening. Consistent with previous research on music-evoked autobiograph-
ical memories (MEAMs), we found that the medial temporal lobe may not be necessary for MEAMs to
be experienced. However, focusing attention toward MEAMs increased medial temporal lobe activity
and behavioral ratings of memory reliving. Focusing on MEAMs also increased connectivity between the
left hippocampus and default-mode network regions supporting autobiographical recall, in addition to
sensorimotor regions that music familiarity is known to modulate. The results substantiate the notion that
how we experience music and associated memories depends on how we attend to the music. We provide
initial evidence suggesting that music-evoked remembering integrates a sensorimotor representation of
the music with episodic representations for the related autobiographical content.

Keywords: attention, autobiographical memory, default-mode network, hippocampus

Supplemental materials: http://dx.doi.org/10.1037/pmu0000210.supp

A 2014 survey based in the United States estimated 93% of the a study. Accordingly, music serves as a powerful tool with which
population report listening to music and do so for an average of 25 to study the interaction of brain networks in response to dynamic
hr a week (Nielsen, 2014). Such a high rate of musical engagement and naturally occurring stimuli that are variously bound to auto-
in the population helps to explain why reports of music-evoked biographical memory content. The present study uses music to
memories of people, places, and events are common phenomena in investigate the interaction between memory and attention systems
both young and old populations (Cuddy, Sikka, & Vanstone, 2015; using a task that reflects common methods of engaging with music,
Janata, Tomic, & Rakowski, 2007; Sloboda & O’Neill, 2001). regardless of whether or not an individual has musical training.
Unlike in the case of traditional memory studies, the memory cue How we experience music depends, in part, on how we direct
(i.e., a piece of music) and associated episodes are bound outside attentional resources. For example, when listening to a familiar
of the laboratory, that is, during normal life, typically years before song we may be reminded of childhood friends and direct our
attention toward memories of them and associated places and
events. On another occasion, while listening to a live performance,
we may focus attention on a guitar solo or on the accompanying
Benjamin Kubit and Petr Janata, Department of Psychology and Center
instrumentation. Thus, how the brain engages with music depends
for Mind and Brain, University of California, Davis.
We are grateful to Fred Barrett for his advice on the functional MRI not only on a listener’s familiarity with it or the memories that
analyses, Melanie Rothfuss for her assistance in data collection, and might be associated with it, but also on the attentional focus of the
Victoria Chen, Jessica Yeung, Neda Sefidpour, Alex Greene, and Chriselle listener (Janata, Tillmann, & Bharucha, 2002). We performed a
Johanna Vinson for their help in quantifying the number of instruments and functional MRI (fMRI) study to understand how networks of brain
percent vocals present in each musical excerpt. This project was made areas reconfigure to support particular experiential states, from
possible by support, in part, from the Child Family Fund for the Center for
among the multitude of possible states enabled by music. The
Mind and Brain.
Correspondence concerning this article should be addressed to Petr
design manipulated the locus of attention that listeners were asked
Janata, Department of Psychology and Center for Mind and Brain, Uni- to adopt while listening to a large number of popular music
versity of California, Davis, 267 Cousteau Place, Davis, CA 95618. E-mail: excerpts that either were or were not memory evoking on a
pjanata@ucdavis.edu per-person basis.

82
ATTENTION AND MEMORY-EVOKING MUSIC 83

To maximize differences between experiential states, we created served when other hippocampus-dependent memories are impaired
a stark contrast in attentional focus. Despite using music that we (Baird & Samson, 2009; Cuddy et al., 2015), the results from
knew ahead of time was able to evoke autobiographical memories Janata (2009) suggests that MTL involvement is not necessary
in a participant, we predicted that focusing attention on the com- for experiencing MEAMs. However, the content specificity of
plement of instruments used in any given excerpt would recruit the MEAMs of patients with MTL damage remains unknown. The
fronto-parietal control network (FPCN). By contrast, we expected MTL may play a critical role in the explicit search for and
that focusing attention on the memories evoked by the music elaboration of MEAMs. These predictions are in accordance with
would preferentially drive activity within default-mode network multiple trace theory, which views the hippocampus as necessary
(DMN) regions known to be active during autobiographical mem- for the recollection of specific, vivid episodes (Nadel & Mosco-
ory tasks. Critically, our design allowed us to test the role of the vitch, 1997; Winocur, Moscovitch, & Bontempi, 2010). Research-
hippocampus in music-evoked autobiographical remembering spe- ers have postulated that directing attention toward memory content
cifically, while providing more general insight into the nature of may lead to more hippocampally mediated integration across brain
hippocampal involvement in the recollection of memories associ- areas involved in forming the event models that support rich
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

ated with personally relevant complex natural stimuli. episodic recollection. However, the question of how attentional
This document is copyrighted by the American Psychological Association or one of its allied publishers.

focus interacts with memory specificity when it comes to hip-


pocampal recruitment for remembering experiences has not been
Music-Evoked Autobiographical Memories
directly addressed (Ranganath & Ritchey, 2012).
Music provides a means of accessing bits of lifetime period,
general event, and event-specific information that constitute auto-
Memory for Music
biographical memories (Conway & Pleydell-Pearce, 2000; Janata
et al., 2007). Music-evoked autobiographical memories (MEAMs) Separate from the episodic and autobiographical memories that
are characterized by similar phenomenological dimensions of re- may be associated with a piece of music is the memory for a piece
living and engagement of brain networks as are memories evoked of music itself. Memory for music is often assessed using mental
by picture cues and interview techniques. Autobiographical mem- imagery tasks, and is typically associated with activity in premotor
ories evoked by music elicit vivid mental imagery and emotions regions of the frontal lobe and in the basal ganglia (Halpern &
(Belfi, Karlan, & Tranel, 2016; Janata et al., 2007), as do nonmu- Zatorre, 1999; Herholz, Halpern, & Zatorre, 2012; Janata & Graf-
sical memories (Sutin & Robins, 2007). Neuroimaging studies of ton, 2003; Leaver, Van Lare, Zielinski, Halpern, & Rauschecker,
MEAMs have demonstrated increases in blood-oxygen-level- 2009). Separately from music, these brain regions support implicit
dependent signal in DMN regions, including lateral parietal, tem- sequence learning (Censor, Dayan, & Cohen, 2014; Sun, Miller,
poral, medial prefrontal, and posterior cingulate cortices (Ford, Rao, & D’Esposito, 2007; Willingham, 1998) and are crucial for
Addis, & Giovanello, 2011; Janata, 2009). The DMN also partic- auditory imagery in general, such as in speech perception and
ipates in the search for and elaboration of autobiographical mem- production (Lima, Krishnan, & Scott, 2016).
ories unrelated to music (Cabeza & St Jacques, 2007; Svoboda, Recent research has demonstrated that representations of long-
McKinnon, & Levine, 2006). known familiar songs can be distinguished from newly encoun-
Our use of tasks for explicitly orienting attention either toward tered (unfamiliar) music in the presupplementary motor area (pre-
or away from memories while listening to memory-evoking music SMA) and adjacent cingulate regions (Jacobsen et al., 2015),
was intended to address an issue regarding the role of the hip- suggesting these regions maintain long-term representations
pocampus in autobiographical memory retrieval. Medial temporal unique to specific pieces of music. Much like procedural memo-
lobe (MTL) regions, in particular the hippocampus, play a prom- ries, memory for music is often preserved despite extensive deg-
inent role in the retrieval of autobiographical memories (Daselaar radation of hippocampal and cortical regions involved in episodic
et al., 2008). However, the role of the hippocampus during music- and semantic memory retrieval (Belfi & Tranel, 2014; Huijgen et
evoked remembering has been unclear. Janata (2009) did not find al., 2015). Activity in the pre-SMA was found in previous MEAM
activity in MTL regions during a music listening task in which imaging studies (Ford et al., 2011), and has been shown to increase
⬃30% of the presented songs evoked memories, yet in which there in accordance with individual ratings of familiarity with music, but
was no explicit instruction to retrieve autobiographical memories. not the salience of evoked memories (Janata, 2009). Both neuro-
In other words, an excerpt could be unfamiliar, familiar but not logical and neuroimaging evidence suggest that involvement of
memory evoking, or familiar and memory evoking. The memories sensorimotor regions, specifically the pre-SMA, in MEAMs may
and remembering experiences were therefore typical of spontane- be independent of attentional focus and rely on more automatic
ous autobiographical memories (Rubin & Berntsen, 2009) rather procedural processes typical of sequence learning.
than memories retrieved obligatorily in response to a retrieval cue.
By contrast, Ford et al. (2011) used a task that required memory
Attention and Music
search and categorization for each presented music item and found
increased MTL activity as the event specificity of an autobiograph- Although the degree to which music and memories engage the
ical memory increased. DMN seems to depend on the activation of self-relevant associa-
We postulate that deliberate memory retrieval when listening to tions, explicit instructions to orient attention within an auditory
music involves the hippocampus (Ford et al., 2011), whereas scene, such as to detect a specific timbre or tone, result in activa-
passive or spontaneous memory retrieval engages mostly non- tion of parietal and frontal regions involved in the control of
MTL regions (Janata, 2009). In agreement with literature that attention (Janata et al., 2002). Focusing attention toward the struc-
suggests that memory for music, as well as for MEAMs, is pre- ture of a song engages the FPCN, which supports sustained and
84 KUBIT AND JANATA

adaptive attentional resources during auditory and visuospatial Methods for stimulus presentation in both experiments followed
tasks unrelated to music (Dosenbach, Fair, Cohen, Schlaggar, & those used in Janata (2009).
Petersen, 2008). The FPCN supports top-down attentional control
and involves coordination between regions of the lateral prefrontal Procedure
cortex, the anterior insula, dorsal anterior and middle cingulate
cortices, dorsal parietal regions, and the precuneus. Activity in Figure 1B depicts the structure of a single trial in either exper-
dorsal parietal and prefrontal areas within the FPCN is negatively iment. Each excerpt was preceded by a cue to either attend-music
correlated with activity in the DMN during resting states (Fox et or attend-memory. A participant’s goal during attend-music trials
al., 2005; Raichle & Snyder, 2007) and during a variety of atten- was to focus attention on the structural features of the song, thus
tion, memory retrieval, and social tasks (Jack et al., 2013; Spreng, directing attention away from any memories associated with the
Stevens, Chamberlain, Gilmore, & Schacter, 2010). As such, the song. To ensure that participants focused attention on the music
networks have been hypothesized to represent two distinct modes during these trials, immediately following each trial participants
of attentional focus: analytic externally focused attention that rated the proportion of time that they spent singing along covertly
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is emotionally detached and emotionally engaged internally fo- in their minds, the proportion of the excerpt containing vocals (1:
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cused attention supported by the FPCN and DMN, respectively 0 –20%, 2: 20 – 40%, 3: 40 – 60%, 4: 60 – 80%, and 5: 80 –100%),
(Andrews-Hanna, 2012; Buckner, Andrews-Hanna, & Schacter, and the number of instrument groups present in the excerpt (1: one
2008; Kim, 2012). It remains unclear how the FPCN and DMN or two, 2: three, 3: four, 4: five, and 5: six or more). Participants
interact during spontaneous autobiographical memory retrieval, chose from eight possible instrument groups: percussion, bass,
and whether directing attention away from MEAMs influences brass, guitar, strings, woodwind, keyboards, and vocals. Instru-
activity in the DMN. ment groups were based on timbral similarity to accommodate the
wide range of genres sampled and to reduce the difficulty of the
task for nonmusicians.
The Present Study The goal during attend-memory trials was to focus attention on
Previous research demonstrating greater activity and communi- memories associated with a song and to search for autobiograph-
cation within the DMN has comprised a rich phenomenology of ical associations if none spontaneously presented themselves. To
the emotional, social, and autobiographical dimensions of music- encourage the focusing of attention toward the memories, follow-
related experiences (Barrett & Janata, 2016; Ford, Rubin, & Gio- ing each attend-memory trial participants had to rate the number of
vanello, 2016; Janata et al., 2007; Wilkins, Hodges, Laurienti, memories experienced (1: one, 2: two, 3: three, 4: four, 5: five or
Steen, & Burdette, 2014), but it has not explicitly examined how more), the proportion of time spent thinking about specific events,
the allocation of attention configures the neural networks support- and the proportion of time spent thinking about specific individ-
ing those experiences. Similarly, research demonstrating FPCN uals (1: 0 –20%, 2: 20 – 40%, 3: 40 – 60%, 4: 60 – 80%, and 5:
involvement in tasks directing attention toward musical features 80 –100%). Following every excerpt in both attend-music and
such as timbre and tonality typically ignore the listener’s internal attend-memory trials, participants were cued to rate the strength of
experience and use musical stimuli that purposefully minimize autobiographical association (salience), vividness of evoked men-
personal relevance (Janata et al., 2002). tal imagery, and familiarity with the excerpt on 5-point scales (1:
Because pieces of music unfold over time, they can serve as a not at all, 2: weakly, 3: moderately, 4: strongly, and 5: extremely).
sustained and potent cue for autobiographical memories that may
arise spontaneously even if attention is directed elsewhere. Behavioral Data Analysis
Whereas previous studies examining the interactions between at-
tention and memory focused on encoding periods (Chun & Turk- Linear mixed models were used to examine the effects of
Browne, 2007) or directed attention to different aspects of the Attentional Focus and Memory Status on poststimulus responses
retrieved memory itself (Denkova, Dolcos, & Dolcos, 2015), mu- by separately predicting salience, vividness, and familiarity as an
sic provides us with the opportunity to investigate memory re- interaction between Attentional Focus and Memory Status, while
trieval during both internally and externally focused attention. The allowing mean response to vary across participants. Allowing for
goal of Experiment 1 was to investigate the influence of attentional individual differences in poststimulus responses significantly im-
focus on the phenomenology of MEAM recollection. The goal of
Experiment 2 was to investigate how evoked autobiographical
memories and the focus of attention interact across brain networks
engaged by music. In both experiments, we manipulated the locus
of attention while participants listened to familiar and novel 30-s
excerpts of music.

General Method

Stimuli
Musical stimuli were 30-s preview excerpts from the iTunes
Music store of songs that were in the Billboard top-100 charts Figure 1. (A) Diagrams of the experimental structure and (B) the time-
during the time participants were between the ages of 7–19 years. course of a single trial.
ATTENTION AND MEMORY-EVOKING MUSIC 85

proved model fit (␹2 ⫽ 14.8, df ⫽ 2). Approximately 8.2% of the read and agreed to a consent form followed by a series of test
total variance in salience ratings could be attributed to differences sounds to ensure that their speaker volume was set at a comfortable
between participants. Participants were modeled as a random ef- level. Next, participants answered questions from the Music En-
fect for every linear mixed model. gagement Style Scale (unpublished) as well as the Brief Affective
All mixed model analyses were performed using the MIXED Neuroscience Personality Scales (Barrett, Robins, & Janata, 2013).
procedure in SAS software (Version 9.4). Results from Type III Neither scale was used in analyses related to the present study.
tests for fixed effects, Tukey’s procedure for pairwise compari- Participants performed 15 attend-music and 15 attend-memories
sons, and least-squares estimated means are reported for each trials (Figure 1B). We did not know ahead of time which songs
model. Kenward–Roger corrections (Kenward & Roger, 1997) would be memory evoking for any given participant and therefore
were applied to adjust for small sample size. Effect sizes are could not implement, a priori, the balanced 2 ⫻ 2 factorial design
reported as partial eta squared (␩p2) for main effects and Cohen’s used in Experiment 2. However, our stimulus selection methods
dav for t tests (Lakens, 2013). resulted in a collection of songs that evoked autobiographical
memories in 92% of participants. Participants who did not report
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Experiment 1 experiencing at least four memory-evoking excerpts during both


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Attentional Focus conditions were excluded from all analyses. For


We used a single-factor design to examine the influence of the remaining participants, songs rated as at least “weakly autobi-
attentional focus on behavioral ratings made after listening to ographical” (2 out of 5) were categorized as memory evoking for
memory-evoking music. We hypothesized that a listener’s focus of all analyses. The post hoc categorization of Attentional Focus trials
attention would influence the reported salience and vividness of as memory evoking or nonevoking allowed us to build linear mixed
evoked autobiographical memories. models using Attentional Focus and Memory Status to predict
behavioral ratings.
Materials and Method
Results
Participants. Participants were 48 University of California,
Davis, undergraduate student volunteers who participated for par- For clarity and brevity, we only report results from mixed
tial course credit. No participants reported neurological or hearing models examining the influence of Attentional Focus and Memory
impairments. Sixteen participants were excluded from analyses Status on the three questions answered at the end of every trial
because an insufficient number of excerpts evoked autobiograph- (i.e., autobiographical salience of evoked memories, vividness of
ical memories during the experiment, resulting in a final sample of evoked memories, and familiarity with the music; Figure 2A). The
17 women and 15 men (age: range ⫽ 18 – 44 years, M ⫽ 20.7 ⫾ mixed model predicting autobiographical salience revealed sig-
3.9 years). nificant fixed effects of Memory Status, F(1, 961.87) ⫽ 1664.48,
Procedure. The study was completed online using Ensemble, p ⬍ .001, ␩p2 ⫽ 0.634. Pairwise comparisons demonstrated higher
a web-based survey delivery and experiment management system salience ratings during memory-evoking compared with non–
(http://atonal.ucdavis.edu/ensemble/; Tomic & Janata, 2007) in memory-evoking songs, t(961.9) ⫽ 40.80, p ⬍ .001, dav ⫽ 0.779.
coordination with a MATLAB engine (The Mathworks, Natick, The mixed model predicting vividness showed a significant
MA) that selected stimuli and trial types. Participants completed fixed effect of Memory Status, F(1, 954.42) ⫽ 687.52.89, p ⬍
the paradigm online at a quiet location of their choosing using a .001, ␩p2 ⫽ 0.419, as well as an interaction between Memory Status
laptop or desktop computer. Before the study began, participants and Attentional Focus, F(1, 947.56) ⫽ 8.37, p ⫽ .004, ␩p2 ⫽ 0.009.

Figure 2. Influence of Attentional Focus (attend-memory/attend-music) and Memory Status (memory-evoking/


nonevoking) on postmusic behavioral ratings of autobiographical salience, vividness, and familiarity. Solid and
stippled fills correspond to memory-evoking and nonevoking stimuli, respectively. Error bars represent 95%
confidence interval. All spanning lines indicate differences that are significant at p ⬍ .001 except where noted.
Significance value markers: ⴱ p ⬍ .05. ⴱⴱ p ⬍ .01.
86 KUBIT AND JANATA

Pairwise comparisons showed the interaction was due to increased was directed toward the perceptual analysis of the music. We
rated vividness during attend-memory trials compared with attend- hypothesized that activity within the FPCN would increase if
music trials when songs were memory evoking, t(941.3) ⫽ 2.49, participants attended to structural aspects of music, and would
p ⫽ .010, dav ⫽ 0.294, whereas there was no significant difference increase in core DMN regions and the MTL if participants focused
in vividness ratings between attention conditions for songs that attention on searching for and elaborating autobiographical mem-
were nonevoking, t(942.5) ⫽ 1.71, p ⫽ .090, dav ⫽ 0.026. ories associated with the music, regardless of whether or not the
The mixed model predicting familiarity revealed a significant music was originally memory evoking. We also hypothesized that
fixed effect of Memory Status, F(1, 963.59) ⫽ 577.85, p ⬍ .001, the connectivity between DMN and MTL regions would depend
␩p2 ⫽ 0.375, and a significant effect of Attentional Focus, F(1, on the attentional focus while listening to memory-evoking music.
936.78) ⫽ 7.28, p ⬍ .007, ␩p2 ⫽ 0.008. Pairwise comparisons show
greater familiarity ratings for memory-evoking songs compared
with nonevoking songs, t(963.6) ⫽ 24.04, p ⬍ .001, dav ⫽ 0.498, Materials and Method
as well as for songs during attend-music compared with attend- Participants. Eleven UC Davis undergraduate and graduate
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

memories trials, t(936.8) ⫽ 2.70, p ⫽ .007, dav ⫽ 0.304. See Table


students (six females, 18 –30 years; mean age: 22 years) partici-
This document is copyrighted by the American Psychological Association or one of its allied publishers.

S1 in the online supplemental materials for all comparisons.


pated in the experiment after providing informed consent. Four of
the participants reported having at least 1 year of formal musical
Discussion training and none reported neurological or hearing impairments.
These participants had been invited to take part in the functional
Overall, Attentional Focus and Memory Status influenced the
neuroimaging experiment because each had experienced at least 16
music listening experience. As expected, based on the experimen-
MEAMs in a preliminary MEAM sampling study. None of the
tal design, memory-evoking trials were reported as more autobio-
participants in the fMRI study had participated in Experiment 1.
graphically salient, vivid, and familiar compared with non–
All data collection procedures were approved by the University of
memory-evoking trials. Songs heard during attend-music trials,
California at Davis Institutional Review Board.
during which participants had to direct their attention to structural
We recognize that our sample may be considered small by
aspects of the music, were rated as more familiar than songs heard
present day expectations that arose in response to reports that some
during attend-memories trials. This suggests that focusing on the
types of fMRI analysis practices were prone to false positives
tonal and rhythmic information that makes a song unique leads
to greater feelings of familiarity with the music, regardless of (Type I error; Eklund, Nichols, & Knutsson, 2016). Below, we (a)
whether a song is memory evoking. In addition, participants re- adopt a nonparametric analysis strategy that helps guard against
ported increased vividness for memory-evoking songs during Type I error and performs well with small samples, and (b) discuss
attend-memories trials compared with attend-music trials. The issues of power for this specific study.
interaction between Attentional Focus and Memory Status indi- Stimuli. During the preliminary MEAM sampling study, par-
cates that explicitly orienting attention toward the elaboration of ticipants heard a random sample of 40 musical excerpts. Unlike in
associated memories during music listening results in a richer Experiment 1, the preliminary sampling study did not explicitly
recollective experience, whereas focusing on the structural aspects instruct participants to search for memories; rather, participants
of the music decreases the vividness with which evoked memories passively listened to music and reported any spontaneously evoked
are experienced. autobiographical memories. On the basis of the per-participant
The results from Experiment 1 demonstrate that the locus of results from the preliminary MEAM sampling study, we were able
attention during music listening changes both the experiencing of to present music that we knew was capable of evoking MEAMs in
the music and of associated autobiographical memories. These each participant. The preliminary MEAM sampling study was
findings motivated further investigation into the attentional pro- completed on average ⬃2 months before the fMRI session. Sixteen
cesses that underlie differences in how people experience memory- additional excerpts of 30-s duration were chosen randomly for
evoking music. We designed and implemented Experiment 2 to each participant, using the aforementioned criteria, for presenta-
examine brain network connectivity and modularity as a function tion in the fMRI session.
of attention while participants listened to memory-evoking music. Procedure. The experiment was controlled by Presentation
software (Neurobehavioral Systems, Inc., Albany, CA) in coordi-
nation with a MATLAB engine (The Mathworks, Natick, MA) that
Experiment 2 selected stimuli and submitted responses via Ensemble. For each
Using a 2 ⫻ 2 factorial design, we examined the influence of participant, all information pertaining to scanner timing pulses,
attentional focus and autobiographical salience during music lis- stimulus event codes, trial-type, and responses to questions were
tening on behavioral ratings and the connectivity of the FPCN, recorded to a text file that was used subsequently to construct the
DMN, and the MTL. We utilized univariate models of the fMRI design matrices used in the statistical analyses.
data to identify regions of interest (ROIs) that were then used in The experiment followed a balanced 2 ⫻ 2 factorial design
network-level connectivity and community modularity analyses. crossing Attentional Focus (attend-music or attend-memory) with
The fMRI study described in Experiment 2 was motivated by the Memory Status (memory evoking or nonevoking). Four trials of
results of the initial behavioral experiment. On the basis of the results each of the four possible trial types were randomly ordered within
of Experiment 1, we expected participants to report more vivid each run. Thirty-two song excerpts, each 30 s in duration, were
evoked memories when attention was directed toward the search presented across two fMRI scanning runs. Each run contained 16
for memories, compared with those experienced while attention songs and lasted ⬃18 min (Figure 1A). Data were collected for
ATTENTION AND MEMORY-EVOKING MUSIC 87

three resting-state runs during which participants lay passively playing, poststimulus question and response epochs, and Atten-
with eyes closed in the scanner for 5 minutes. tional Focus trial cue presentation. An additional six motion co-
Immediately after each trial, participants used a five-button variates (X, Y, Z, pitch, yaw, roll) as well as run and session
response device using the fingers of their left hand to respond to constants were also included in the model. A high-pass filter of
each of the six poststimulus questions. Participants’ eyes were 120 s was applied and resting-state runs served as the implicit
closed during scanning runs. Music excerpts, trial-type cues, and baseline for model estimation.
poststimulus question cues were all presented auditorily via insert Participant-level music regressors were collapsed into the
earphones (Model S14: Sensimetrics Corporation, Malden, Mas- four potential trial types: attend-music and memory-evoking excerpts,
sachusetts). Sound isolation headphones were worn over the ear- attend-music and nonevoking excerpts, attend-memory and memory-
phones as extra padding to prevent scanner noise from interfering evoking excerpts, attend-memory and nonevoking excerpts. A trial
with music listening. Before the scanning runs, the volume of the was considered memory evoking if it was rated at least “weakly
music was adjusted so that is was as loud as the participant could autobiographical” (2 out of 5) on the Autobiographical Salience
comfortably stand but without noticeable distortion of the audio scale in the scanner. Nonevoking trials thus had an autobiograph-
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quality. ical salience rating of “not at all autobiographical” (1 out of 5).


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To improve performance in the MRI scanner, participants re- Classification of music previously identified as memory evoking
viewed question-response mappings with a visual aid during the before the scanner session was consistent with subsequent classi-
20-min transit to the scanning facility. Before entering the scanner, fication based on responses made in the scanner. From the non-
participants practiced the number of instrument groups question evoking category, participants endorsed on average three addi-
until they answered correctly for each of the six practice songs. tional music excerpts as memory evoking that were not presented
To further reduce variability among participants that might be during the preliminary study. These stimuli were thus reassigned to
associated with encountering a strange and potentially anxiety- the memory-evoking category for the analysis. On average partic-
inducing environment, we had them complete an additional eight ipants had fewer trials for nonevoking attend-memory trials com-
practice trials in a mock MRI scanner. pared with all other conditions. The minimum number of trials for
fMRI acquisition parameters. MRI data were collected on a a condition across all participants was five. A full factorial analysis
3T Siemens Trio (Siemens Medical Solutions, Erlangen, Ger- of variance was carried out using the first-level general linear
many) at the Imaging Research Center at the UC Davis Medical models. Additional one-sample t tests were evaluated contrasting
Center. Three sets of MR images were acquired for each partici- attend-memory and attend-music trials for both memory-evoking
pant. The first was a high-resolution 3D magnetization prepared and nonevoking trials. A mean high-resolution anatomical image
rapid gradient echo (MPRAGE) structural image (field of view ⫽ volume was created from study participants’ anatomical images
256 ⫻ 256 mm, 192 slices, resolution ⫽ 1 ⫻ 1 ⫻ 1 mm, time and segmented to create a group-level gray matter mask. A mask
repetition [TR] ⫽ 2.5 ms, time echo [TE] ⫽ 4.77 ms, flip angle ⫽ made from the intersection of participants’ EPIs was used along
7 degrees). The second was a whole brain volume of T1-weighted with the gray matter mask to limit regions of the brain included for
axial images (32 slices, resolution ⫽ 0.8 ⫻ 0.8 ⫻ 3.2 mm, TR ⫽ statistical analysis.
617 ms, TE ⫽ 9.6 ms, flip angle ⫽ 70 degrees). The third set was Cluster mass thresholding. Results from the full factorial
a whole brain volume of multiband gradient echoplanar images analysis of variance and all subsequent analyses were corrected for
(EPI) using the same 32 slice positions and orientations as the family-wise error rate at p ⬍ .05 using threshold-free cluster
previously collected T1-weighted images (resolution ⫽ 3.2 mm enhancement (TFCE) with 5,000 permutations (Smith & Nichols,
isotropic, TR ⫽ 1.03 s, TE ⫽ 25 ms, flip angle ⫽ 65 degrees, 2009). The algorithm performs cluster-based inferences, taking
multiband acceleration factor ⫽ 2). Zero-fill interpolation was into account both voxel extent and voxel height. MatlabTFCE
applied to the EPIs in k-space. (Thornton, 2016) and custom code were used to implement the
fMRI preprocessing parameters. The fMRI data were ana- methods described by Smith and Nichols in conjunction with the
lyzed using SPM5 software (http://www.fil.ion.ucl.ac.uk/spm/ sign-flipping permutation approach described in Winkler, Ridg-
software/spm5/) for the following preprocessing steps applied to way, Webster, Smith, and Nichols (2014) to define the null cluster
the series of blood-oxygen-level-dependent EPI-image volumes: mass distribution for each contrast. The observed value of the
realignment of the first volume of the all runs to the first volume TFCE statistic of a given voxel was compared with the corre-
of the first run, realignment of the EPIs to the first volume of each sponding derived null distribution, to determine the probability of
run, coregistration of the reference EPI volume to the T1-weighted observing the value by chance alone. Nonparametric approach
axial images, coregistration of the coplanar volume to the partic- avoids assumptions of parametric methods associated with in-
ipant’s high-resolution structural volume, and spatial normaliza- creased Type I error (Eklund et al., 2016), particularly for exper-
tion of the high-resolution volume to the Montreal Neurological iments with low degrees of freedom (Nichols & Hayasaka, 2003).
Institute 152-scan (MNI152) T1 template with propagation of the Functional connectivity. Significant clusters from the con-
normalization parameters to the coplanar and EPI volumes. The trast of attend-memory and attend-music trials were used as ROIs
normalized images were resliced to consist of 2-mm isotropic in a functional connectivity analysis. Time series from clusters
voxels and smoothed with a 5-mm full-width-half-max (FWHM) were extracted in a two-step process. First, at the level of individ-
isotropic kernel. ual participants, residual time series from a GLM controlling for
fMRI statistical analyses. All analyses were performed in noise covariates, including global mean signal (cf. Murphy & Fox,
MATLAB using a combination of third party toolboxes (SPM5, 2017; Power et al., 2014), and main effects of Attentional Focus
IPEM Toolbox) and custom code. The main model consisted of and Memory Status were extracted for each cluster and trial type
regressors for each 30-s period during which a musical excerpt was during music presentation and rest epochs. Second, principle com-
88 KUBIT AND JANATA

ponents analysis was used to extract the first n-components of each Attentional Focus by Memory Status condition, the left hippocam-
time series. Components of the noise-corrected time series data pal ROI time series extracted from a sphere with a 6-mm radius
were included in a cluster’s reconstructed time series if the vari- centered at the peak of activation, and four regressors modeling the
ance explained by a component was greater than the amount of interaction between (product of) each condition with the ROI time
variance explained by 95% of a null distribution of randomly series. A high-pass filter of 120 s was applied to the design matrix.
permuted time series. For each of the four task conditions and rest The same six motion covariates and run and session constants from
epochs, the Pearson correlation between each clusters’ recon- the main model were also included in the generalized psychophys-
structed time course was calculated within participants and Fish- iological interactions model. One-sample t tests were evaluated
er’s r-to-z transformation was applied to facilitate comparison across participants, contrasting hippocampal functional connectiv-
across participants. Between-participants one-sample t tests were ity estimated in each PPI condition to the functional connectivity
used to threshold average group-level correlation matrices with z during resting-state runs.
values significantly different from zero. The resulting p values for Power. Neuroimaging studies with small sample sizes can find
each of the five correlation matrices were corrected for multiple false positives due to a lack of power (Button et al., 2013). The
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comparisons using a false discovery rate of p ⬍ .05. positive predictive value (PPV) quantifies the probability that an
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To examine network-level properties during task and rest, each observed effect is a true effect. The PPV depends on a study’s
correlation matrix was transformed to a directed adjacency matrix, power, Type I error rate, and the prestudy odds ratio (R). R is the
assigning z significantly greater than zero a value of one and a priori probability that the investigated effect is a true effect.
significantly less than zero a value of negative one. We used the We believe the PPV of our observed effects is maximized in three
Louvain modulatory algorithm implemented in the Brain Connec- ways. First, although the overall number of participants is low, the
tivity Toolbox (Rubinov & Sporns, 2010) to detect communities in number of volumes collected during resting-state scans complies
each adjacency matrix. Because this algorithm has an element of with experiment-based recommendations for maximizing reliabil-
randomness to it, we obtained 10,000 solutions to find the most ity of per-participant estimates (Birn et al., 2013). Second, as
consistent community assignments. The variability associated with previously mentioned, the use of TFCE effectively reduces Type I
these solutions is illustrated in Figure S2 in the online supplemen- error rates. Third, our hypotheses are based on two well-
tal materials. First, five resting-state communities identified by the documented effects in the neuroimaging literature (the DMN and
algorithm were consistently found across iterations to include five FPCN networks), effectively increasing the prestudy odds ratio (R)
nodes: (a) FPCN regions together with the left inferior frontal for each main effect. Specifically, given the results of countless
gyrus (IFG), (b) frontal DMN regions including the left medial studies, the expected effect of attention-demanding tasks is one of
prefrontal cortex (MPFC), (c) posterior DMN regions including reduced activity in the DMN, thus resulting in a high value of R in
the left MTL, (d) right mid-cingulate cortex (MCC), and (e) the our Attend Music versus Attend Memories contrast. With regard to
left pre-SMA. Next, for every task-based matrix, community mem- the effects we report for the hippocampus, we believe that the
bership was assigned relative to the five key clusters found during appropriate scope for our “estimate” of R is the broader set of
rest. Cluster N from task matrix A was assigned to a resting-state studies that drove our hypothesis: that vivid remembering and
community if the algorithm consistently assigned the cluster to the active attempts to retrieve autobiographical content will increase
same community as one of the five resting-state node communi- activity in the hippocampus. Thus, even though no previous study
ties. If a cluster did not consistently belong to any of the resting- has examined the interaction of attention and autobiographical
state communities it was assigned to a unique task-evoked com- remembering in a fully crossed factorial design, the previous
munity. expectation that we would observe greater activity in the hip-
MTL beta-weights. Probabilistic maps from the Anatomy pocampus during elaboration of memories associated with
Toolbox (Eickhoff et al., 2005) were used to define MTL ROIs at memory-evoking music is quite high, given the literature showing
the bilateral hippocampus (hipp), consisting of CA1, CA2, and increases in hippocampal activity during effortful retrieval of
CA3, dentate gyrus (DG), entorhinal cortex (EC), and subiculum memory content and vivid remembering.
complex (SUB). Within these ROIs, beta-estimates for activity in Analyses of behavioral data. To examine the influence of
each of the four conditions specified in the full factorial analysis autobiographical salience on performance during attend-music tri-
were extracted using the MarsBar toolbox (Brett, Anton, Vala- als, five research assistants independently assessed the number of
breque, & Poline, 2002). Linear mixed models treating participant instrument groups present for each excerpt. If the number of
as a random effect were implemented in SAS (Version 9.4) to instrument groups was not unanimously agreed upon across re-
examine the effects of Attentional Focus and Memory Status on search assistants, a further group assessment was carried out until
activity in regions of the MTL. a unanimous decision was reached. These ratings were used as the
Psychophysiological interaction. Results from the main effect correct answers to the “number of instrument groups” question.
of Attentional Focus were masked using the left hippocampal ROI Similar methods were used to score the responses for the percent
identified from the MTL beta-weight analysis. Small volume cor- of the excerpt containing vocals. Research assistants used Audac-
rection with a false discovery rate of ␣ ⫽ .05 and voxel extent ity (http://audacityteam.org/) to label the period of time a human
threshold of five voxels revealed a significant peak of activation voice was present in each excerpt. For both sets of responses,
located at x ⫽ ⫺22, y ⫽ ⫺12, z ⫽ ⫺20. Custom code was used participants’ answers were coded as correct or incorrect for anal-
to construct a generalized psychophysiological interactions model ysis.
that tested for differences in whole brain and left hippocampal A preliminary analysis revealed no difference in original sa-
functional connectivity between each possible trial type and lience ratings between Attentional Focus, F(1, 180) ⫽ 0.31, p ⫽
resting-state baseline. The model consisted of regressors for each .577, ␩p2 ⫽ 0.001. That is, there was no a priori difference in the
ATTENTION AND MEMORY-EVOKING MUSIC 89

average autobiographical salience of songs presented during ratings outside of the scanner under passive listening conditions.
attend-music and attend-memory trials. Salience ratings made during attend-music trials were significantly
lower than the preliminary ratings, t(165.1) ⫽ 2.43, p ⫽ .016,
dav ⫽ 0.183. As expected, a similar model predicting the differ-
Results
ence between prescanner and in-scanner ratings of familiarity with
Behavioral results. Overall, the results of the analyses of the a song did not produce any significant effect of Attentional Focus,
behavioral data indicated that the behavioral manipulations were F(1, 165) ⫽ 0.5, p ⫽ .480, ␩p2 ⫽ 0.003. Because participants’ eyes
effective, particularly in decreasing the typical characteristics of remained closed when listening to music in the present experiment,
autobiographical remembering experiences when attentional focus whereas no explicit instructions were given to participants in the
was oriented toward the structural aspects of autobiographically preliminary study, the difference in imagery vividness ratings was
salient music. In total, we ran three distinct sets of analyses to not tested.
examine (a) the influence of Memory Status on behavioral ratings Effects of Attentional Focus and Memory Status. Figure 2B
specific to each Attentional Focus condition, (b) the influence of shows the results of three separate models, each predicting, based
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Attentional Focus on the per-participant consistency of autobio- on per-trial data, the answers to one of the poststimulus questions
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graphical salience ratings made in the fMRI scanner compared as a function of Attentional Focus and Memory Status. The mixed
with those obtained in the preliminary MEAM sampling study, and model predicting salience revealed significant fixed effects of
(c) the influence of both Memory Status and Attentional Focus on Memory Status, F(1, 351) ⫽ 566.17, p ⬍ .001, ␩p2 ⫽ 0.617,
behavioral ratings made after every trial. Attentional Focus, F(1, 342) ⫽ 5.99, p ⫽ .015, ␩p2 ⫽ 0.017, and an
Effects of Memory Status. Memory Status was used in a interaction between Memory Status and Attentional Focus, F(1,
linear mixed model to predict postsong responses unique to attend- 351) ⫽ 6.55, p ⫽ .011, ␩p2 ⫽ 0.018. Pairwise comparisons dem-
memory or attend-music trials. The mixed models predicting Num- onstrated higher autobiographical salience ratings during attend-
ber of Memories, Percent Specific Event, and Percent Specific memory compared with attend-music trials for memory-evoking
Individuals reported during attend-memory trials all revealed sig- songs, t(347) ⫽ 4.23, p ⬍ .001, dav ⫽ 0.331, whereas no difference
nificant main effects of Memory Status, F(1, 169) ⫽ 99.94, p ⬍ was detected in salience ratings between attention conditions for
.001, ␩p2 ⫽ 0.372; F(1, 175) ⫽ 64.15, p ⬍ .001, ␩p2 ⫽ 0.268; F(1, nonevoking songs, t(248.6) ⫽ 0.12, p ⫽ .999, dav ⫽ 0.011.
170) ⫽ 57.83, p ⬍ .001, ␩p2 ⫽ 0.254. Pairwise comparisons The mixed model predicting vividness showed a significant
showed that participants reported more memories, t(169.4) ⫽ 10, fixed effect of Memory Status, F(1, 350) ⫽ 151.09, p ⬍ .001, ␩p2 ⫽
p ⬍ .001, dav ⫽ 0.560, and more time thinking about specific 0.302, as well as an interaction between Memory Status and
events, t(175.3) ⫽ 8.01, p ⬍ .001, dav ⫽ 0.697, and specific Attentional Focus, F(1, 351) ⫽ 9.52, p ⫽ .002, ␩p2 ⫽ 0.026.
individuals, t(170.4) ⫽ 7.6, p ⬍ .001, dav ⫽ 0.474, associated with Pairwise comparisons showed the interaction was due to higher
the music during memory-evoking music trials compared with vividness ratings during attend-memory trials compared with
nonevoking music trials. attend-music for memory-evoking songs, t(346.8) ⫽ 3.30, p ⫽
The mixed model predicting Percent Sing Along during attend- .006, dav ⫽ 0.255, whereas there was no significant difference
music trials revealed a main effect of Memory Status, F(1, 175) ⫽ in vividness ratings between cue types for nonevoking songs,
84.38, p ⬍ .001, ␩p2 ⫽ 0.325. Participants reported spending more t(348.4) ⫽ 1.47, p ⫽ .459, dav ⫽ 0.038. No main effect of
time silently singing along with the music during memory-evoking Attentional Focus was found, F(1, 342) ⫽ 0.620, p ⫽ .432, ␩p2 ⫽
music trials compared with nonevoking music trials, t(175.4) ⫽ 0.001.
9.19, p ⬍ .001, dav ⫽ 0.705. The mixed model predicting familiarity revealed a significant
Table S2 in the online supplemental materials shows the mar- fixed effect of Memory Status, F(1, 349) ⫽ 198.22, p ⬍ .001, ␩p2 ⫽
ginal predicted probabilities from two mixed effects logistic re- 0.362. Pairwise comparisons showed greater familiarity ratings
gressions using the 5-point Autobiographical Salience and Imag- for memory-evoking songs compared with nonevoking songs,
ery Vividness scales to predict whether a participant correctly t(345.9) ⫽ 10.37, p ⬍ .001, dav ⫽ 0.354. No main effect of
identified the number of instrument groups during attend-music Attentional Focus or interaction was found, F(1, 342) ⫽ 0.68, p ⫽
trials. The mixed models revealed a main effect of Salience, F(1, .409, ␩p2 ⫽ 0.002; F(1, 349) ⫽ 1.550, p ⫽ .214, ␩p2 ⫽ 0.004. See
164) ⫽ 4.03, p ⫽ .046, ␩p2 ⫽ 0.024, and Vividness, F(1, 164) ⫽ Table S3 in the online supplemental materials for all comparisons.
4.68, p ⫽ .032, ␩p2 ⫽ 0.028: The more salient the memory or vivid Overall, behavioral results were as expected and consistent
the mental imagery evoked by a song the greater the probability of across the preliminary MEAM sampling, initial behavioral, and
correctly identifying the number of instrument groups. Similar fMRI experiments. A comparison between ratings made in the
models predicting whether a participant correctly identified the fMRI scanner compared with those made previously outside of
percentage of the song excerpt containing vocals revealed no the scanner without any explicit retrieval instructions suggests that
effects of Salience, F(1, 164) ⫽ 0.35, p ⫽ .557, ␩p2 ⫽ 0.002, or the effect of Attentional Focus on memory-evoking music is char-
Vividness, F(1, 164) ⫽ 0.05, p ⫽ .820, ␩p2 ⫽ 0.000. acterized as a decrease in the reported salience of experienced
Effects of Attentional Focus. Attentional Focus was used in a autobiographical memories when attention is focused on the mu-
linear mixed model to predict the difference between autobio- sic. The effect of Memory Status was demonstrated for questions
graphical salience ratings made during the preliminary study and unique to attend-memories trials (e.g., Percent Specific Event), as
the salience ratings made in the scanner for the same song. The well as those unique to attend-music trials (e.g., Percent Sing
mixed model revealed a significant main effect of Attentional Along). In addition, increases in autobiographical salience ratings
Focus, F(1, 165) ⫽ 5.89, p ⫽ 0.030, ␩p2 ⫽ 0.034. Salience ratings increased participants’ chances of correctly identifying the Num-
made after attend-memory trials were more similar to the previous ber of Instruments in a song (Table S2 in the online supplemental
90 KUBIT AND JANATA

materials). Taken together, these results confirm that autobio- factorial analyses of variance. Figure 3 shows the contrast of
graphical information associated with a song was evoked during attend-memory and attend-music trials. Greater activity during
both attend-music and attend-memories trials. Most importantly, attend-memory trials was found throughout DMN regions includ-
the results from the fMRI experiment replicate our initial finding ing lateral parietal, temporal, medial prefrontal and posterior cin-
that attentional focus modulated the vividness of the remembering gulate cortices. Additional activations were observed bilaterally in
experience even though the experienced familiarity of the song the MTL, left superior frontal gyrus (SFG), dorsolateral prefrontal
remained unchanged (Figure 2A). cortex (DLPFC), amygdala and somatosensory cortex, and in right
Neuroimaging results. The neuroimaging results showed that lateralized primary motor cortex extending into MCC (Table 1;
attentional focus influenced the organization of, and functional
Figure 3, regions shown in warm colors). During attend-music
connectivity between, brain networks, particularly when partici-
trials, activity increases were observed in FPCN regions including
pants listened to memory-evoking music. We conducted separate
bilateral IFG, superior parietal lobule (SPL), and right lateralized
neuroimaging analyses to (a) identify whole-brain ROIs compris-
ing DMN and FPCN regions modulated by Attentional Focus, (b) anterior insula (aI). Increased activation during attend-music trials
was also observed in the right DLPFC and left pre-SMA (Table 2;
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examine connectivity and community modularity of DMN and


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FPCN during music listening as a function of Attentional Focus Figure 3, regions in cool colors).
and Memory Status, and (c) examine MTL activity and connec- Connectivity of Attentional Focus ROIs. Figure S1 in the
tivity as a function of Attentional Focus and Memory Status. online supplemental materials displays the resting-state correlation
Effects of Attentional Focus and Memory Status. The main matrix between all Attentional Focus ROIs. Overall, functional
effect of Attentional Focus accounted for variance in 11,043 vox- connectivity between ROIs during rest blocks replicated task-
els, whereas the main effect of Memory Status was significant in evoked network structure. During rest, DMN ROIs in which ac-
only 746 voxels, and no interaction effect was found in the full tivity increased during attend-memory trials showed positive

Figure 3. Influence of each of the two tasks (attend-memory in reddish colors; attend-music in bluish
colors) on brain activity during the music excerpts. The group-level results depicted here are from a direct
contrast of the two task conditions. The white contour lines enclose the volume in which data were available
for all subjects. ROI labels refer to key nodes identified in the community membership analysis (see Figure
5). IFG ⫽ inferior frontal gyrus; MTL ⫽ medial temporal lobe; pSMAa ⫽ presupplementary motor area a;
MPFC ⫽ medial prefrontal cortex; MCC ⫽ mid-cingulate cortex; M1 ⫽ primary motor cortex; TFCE ⫽
threshold-free cluster enhancement; ROI ⫽ region of interest. See the online article for the color version
of this figure.
ATTENTION AND MEMORY-EVOKING MUSIC 91

Table 1
Summary of Significant Clusters From Attend-Memory Greater Than Attend-Music Trials

Left hemisphere (mm) Right hemisphere (mm)


Brain regions BA x y z Nvox Cmass x y z Nvox Cmass

Frontal
MPFC 10/11 ⫺9 57 7 495 3,009
vMPFC 11 4 49 ⫺16 4 1,023
DLPFC 46 ⫺20 38 32 23 1,100
SFGa 9 ⫺22 36 44 1 941
SFGb 9 ⫺16 37 44 12 967
mSFG 8/9 ⫺10 34 53 48 1,024
OFCb 11 ⫺38 28 ⫺16 9 1,105
Temporal
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aTP 38 ⫺43 16 ⫺24 2 986


⫺58 ⫺4 ⫺24 ⫺20
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MTG 21 4 994 60 1 2 1,068


Parietal
MCC/M1 4/23/24 6 ⫺14 43 184 1,644
Som. C 5 ⫺9 ⫺45 58 50 1,077
PCC/MCC 23/24/31 ⫺8 ⫺49 28 1,157 2,699
PCC 23/31 5 ⫺54 25 186 1,665
AG 39/40 ⫺49 ⫺62 30 377 1,879 47 ⫺61 28 2 958
Limbic
Amygdala 25 ⫺24 ⫺8 ⫺12 19 1,070
MTL 28/34/35/36 ⫺27 ⫺37 ⫺12 495 1,801
pMTL 36 ⫺33 ⫺37 ⫺3 3 945 26 ⫺34 ⫺10 1 955
Note. BA ⫽ brodmann area; Nvox ⫽ #voxels; Cmass ⫽ cluster mass; MPFC ⫽ medial prefrontal cortex; vMPFC ⫽ ventral medial prefrontal cortex;
DLPFC ⫽ dorsolateral prefrontal cortex; SFGa ⫽ superior frontal gyrus a; SFGb ⫽ superior frontal gyrus b; mSFG ⫽ medial superior frontal gyrus;
OFCb ⫽ orbitofrontal cortex b; aTP ⫽ anterior temporal pole; MTG ⫽ middle temporal gyrus; MCC ⫽ mid-cingulate cortex; M1 ⫽ primary motor cortex;
Som. C ⫽ somatosensory cortex; PCC ⫽ posterior cingulate cortex; AG ⫽ angular gyrus; MTL ⫽ medial temporal lobe; pMTL ⫽ posterior MTL.

within-network correlations and negative between-network corre- strated increased connectivity during memory-evoking attend-
lations with frontoparietal ROIs. music trials.
Figure 4 shows the contrasts of ROI time-series correlation Community membership of Attentional Focus ROIs. To bet-
estimates for memory-evoking, attend-memory, and attend-music ter understand how each brain area of interest associated with other
trials. For memory-evoking music trials, attending to memories brain areas under the different task conditions, we summarized the
resulted in a reduction of the negative correlations between right community membership for each ROI during rest and in the different
prefrontal FPCN and medial DMN ROIs, as well as a reduction of task conditions in tabular form (Figure 5A) and in brain space (Figure
positive correlations between the right IFG and left DLPFC within 5B). Five reliable resting-state communities were found. The first
the FPCN community. Attending to the music during memory- community consisted mostly of FPCN regions in which increased
evoking trials led to greater connectivity between pre-SMA and activity was observed during attend-music trials, and included bilat-
several left lateralized ROIs including middle temporal gyrus eral IFG, SPL, and right lateralized DLPFC and aI. The FPCN
(MTG), posterior cingulate cortex (PCC) extending dorsally into community also included left somatosensory cortex and SFG ROIs
MCC, and DLPFC. In addition, left IFG and right SPL demon- during resting-state periods, though these areas showed greater activ-

Table 2
Summary of Significant Clusters From Attend-Music Greater Than Attend-Memory Trials

Left hemisphere (mm) Right hemisphere (mm)


Brain regions BA x y z Nvox Cmass x y z Nvox Cmass

Frontal
DLPFC 46 44 36 22 39 953
pSMAa 6 ⫺4 12 50 1 1,015
pSMAb 6 ⫺6 14 48 1 971
IFG 44/45 ⫺49 11 21 40 946 47 12 24 98 947
Parietal
SPL 7/40 ⫺41 ⫺47 47 381 948 39 ⫺50 48 287 955
Insular cortex
Anterior insula 33 23 4 10 969
Note. BA ⫽ Brodmann Area; Nvox ⫽ #voxels; Cmass ⫽ cluster mass; DLPFC ⫽ dorsolateral prefrontal
cortex; pSMAa ⫽ presupplementary motor area a; pSMAb ⫽ presupplementary motor area b; IFG ⫽ inferior
frontal gyrus; SPL ⫽ superior parietal lobule.
92 KUBIT AND JANATA
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Figure 4. Attention-related differences in the correlations of the activity in the regions of interest (ROIs)
defined by the attend-memory versus attend-music contrast (Figure 3; Tables 1 and 2), when listening to
memory-evoking music. The matrix depicts the significant differences (expressed as Fisher’s z scores and
corrected for multiple comparisons using a false discovery rate criterion of p ⬍ .05) in the correlation
estimates obtained separately for attend-memory and attend-music trials. Correlations that were more
positive during attend-memory trials are shown in red, whereas those that were more positive during
attend-music trials are shown in blue. The boxes demarcated with black boundary lines comprise sets of
ROIs that formed distinct communities during the resting-state periods. ROI labels denoted in black are
those regions in which the simple contrast of evoked activity indicated greater activity during attend-
memory than attend-music trials, and vice versa for those ROIs denoted in gray. Gray dashed lines separate
left from right hemisphere loci within each resting-state community, and ROIs are ordered in a rostro-caudal
direction within each hemisphere. L⫽ left; R ⫽ right; pSMAa ⫽ presupplementary motor area a; pSMAb ⫽
presupplementary motor area b; pMTL ⫽ posterior MTL; MCC ⫽ mid-cingulate cortex; M1 ⫽ primary
motor cortex; SFGa ⫽ superior frontal gyrus a; IFG ⫽ inferior frontal gyrus; SPL ⫽ superior parietal
lobule; DLPFC ⫽ dorsolateral prefrontal cortex; aI ⫽ anterior insula; MTL ⫽ medial temporal lobe; PCC ⫽
posterior cingulate cortex; AG ⫽ angular gyrus; vMPFC ⫽ ventral medial prefrontal cortex; MTG ⫽ middle temporal
gyrus; aTP ⫽ anterior temporal pole; MPFC ⫽ medial prefrontal cortex; OFCa ⫽ orbitofrontal cortex a; OFCb ⫽
orbitofrontal cortex b; SFGb ⫽ superior frontal gyrus b; mSFG ⫽ medial superior frontal gyrus. See the online article
for the color version of this figure.

ity during attend-memory trials. A second resting state community ROIs depended on how attention was oriented when participants
was made of up two ROIs located in the pre-SMA; both demonstrated listened to memory-evoking music. Notably, during attend-memory
greater activity during attend-music trials. trials, the pre-SMA, pMTL, SFG, and somatosensory ROIs joined the
The remaining communities consisted exclusively of ROIs that MCC/M1 community and left-lateralized AG, MTG, and anterior
demonstrated greater activity during attend-memory trials. Specif- temporal pole (aTP) joined the left MTL community. Motor and
ically, the third community consisted of the MPFC and left later- somatosensory ROIs did not form a unique community during attend-
alized angular gyrus (AG), lateral and anterior temporal ROIs, music trials with memory-evoking music. Rather than forming a
orbitofrontal cortex (OFC), DLPFC, and SFG, as well as the right distinct community, the somatosensory and pre-SMA ROIs joined the
posterior MTL. A fourth community consisted of the left MTL and left MPFC community, the MCC/M1 ROI joined the left MTL
posterior DMN regions including PCC, and right lateralized AG, community, and pMTL and SFG ROIs joined the IFG community.
MTG, and ventral MPFC. The final resting-state community con- Community membership did not consistently vary as a function of
sisted of right lateralized primary motor cortex, extending to MCC, Memory Status.
the left amygdala, and posterior MTL (pMTL). Effects of Attentional Focus and Memory Status on MTL
Although community membership during tasks largely adhered to beta-estimates. Given the discrepant results in the previous lit-
the resting-state communities, the community membership of several erature regarding the involvement of the hippocampus and MTL
ATTENTION AND MEMORY-EVOKING MUSIC 93
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Figure 5. Community membership analysis. (A) Region of interest (ROI) community membership as a
function of task conditions. Community membership based on reliable membership with key nodes: left
MPFC (blue), left MTL (cyan), left IFG (yellow), right MCC (green), and left pre-SMA (purple). White
entries represent ROIs that formed a task-specific community (Task Com) not found in the resting state
connectivity. Gray ROI labels indicate greater activity during attend-music than attend-memory trials. (B)
Brain-space plot of ROIs and communities. Colors represent resting state community membership. ROIs
with red boundaries switched membership away from their resting state community during at least one task
condition. ROI labels refer to key nodes identified in the community membership analysis. L⫽ left; R ⫽
right; pSMAa ⫽ presupplementary motor area a; pSMAb ⫽ presupplementary motor area b; pMTL ⫽
posterior MTL; MCC ⫽ mid-cingulate cortex; M1 ⫽ primary motor cortex; SFGa ⫽ superior frontal gyrus
a; IFG ⫽ inferior frontal gyrus; SPL ⫽ superior parietal lobule; DLPFC ⫽ dorsolateral prefrontal cortex;
aI ⫽ anterior insula; MTL ⫽ medial temporal lobe; PCC ⫽ posterior cingulate cortex; AG ⫽ angular gyrus;
vMPFC ⫽ ventral medial prefrontal cortex; MTG ⫽ middle temporal gyrus; aTP ⫽ anterior temporal pole;
MPFC ⫽ medial prefrontal cortex; OFCa ⫽ orbitofrontal cortex a; OFCb ⫽ orbitofrontal cortex b; SFGb ⫽
superior frontal gyrus b; mSFG ⫽ medial superior frontal gyrus. See the online article for the color version
of this figure.

more generally during MEAM tasks, we examined the role of the during attend-memory trials for memory-evoking music. The con-
hippocampus in particular detail. Results for all models of MTL dition resulted in increased connectivity between the left hip-
activity are presented in Figure 6. Both left and right hippocampus pocampus and clusters in the right lateral auditory cortices and
ROIs demonstrated fixed effects of Attentional Focus, L F(1, superior temporal gyrus (STG), MPFC, PCC, and extrastriate
31.1) ⫽ 17.43, p ⬍ .001; R F(1, 31.2) ⫽ 10.08, p ⫽ .003, and cortex compared with resting baseline. Directing attention toward
Memory Status, L F(1, 31.1) ⫽ 5.65, p ⫽ .024; R F(1, 31.1) ⫽ music-evoked memories also led to increases in functional con-
5.68, p ⫽ .024. Pairwise comparisons showed larger beta-estimates nectivity between left hippocampus and bilateral striatum (Table
in the left hippocampus for memory-evoking songs during attend- 3). Directing attention toward the features of the music, and thus
memory compared with attend-music trials, t(30.97) ⫽ 2.73, p ⫽ .048. away from music-evoked memories, did not lead to any significant
The effect was not found in the right hippocampus, t(30.99) ⫽ 2.57, clusters compared with resting baseline.
p ⬍ .067. Both dentate gyrus ROIs and subiculum ROIs demonstrated
a fixed effect of Attentional Focus, L F(1, 31) ⫽ 24.75, p ⬍ .001; R
Discussion
F(1, 31.1) ⫽ 12.44, p ⬍ .001; L F(1, 31.1) ⫽ 22.65, p ⬍ .001; R F(1,
31.1) ⫽ 22.65, p ⬍ .001, respectively. Pairwise comparisons showed Our goal with the present study was to examine the interaction
that beta-estimates in both subiculum ROIs were larger for memory- between different forms of attentional focus and music that dif-
evoking music when attending to memories compared with trials in fered in its autobiographical salience, in terms of brain networks
which the attentional focus was on the music, L: t(30.78) ⫽ 4.04, p ⫽ underpinning the listening experiences. Both behavior and mea-
.002; R: t(30.98) ⫽ 2.89, p ⫽ .033. No fixed effect or interactions sures of brain activity varied as a function of Attentional Focus and
were observed in the left or right entorhinal cortex ROIs. See Tables Memory Status. The effects of manipulating the focus of attention
S6 and S7 in the online supplemental materials for all models and were particularly pronounced for those songs that had previously
comparisons. been identified as memory evoking (autobiographically salient) by
Effects of Attentional Focus and Memory Status on hippocam- the individual being tested.
pal connectivity. Figure 7 shows the contrast that identified The behavioral results indicated that participants successfully
regions of increased connectivity with the left hippocampal ROI adopted either externally or internally focused attentional states in
94 KUBIT AND JANATA

DMN and MTL structures, and focusing attention on music re-


sulted in greater activity within the FPCN (see Figure 3). Our
results are in agreement with research characterizing the structural
(Damoiseaux & Greicius, 2009) and functional division between
the DMN and FPCN networks as representing a division be-
tween externally focused, goal-directed attention supported by
FPCN regions and internally focused attention supported by the
DMN (Andrews-Hanna, 2012; Buckner et al., 2008; Kim,
2012).
Previous research has demonstrated similar antagonistic brain
activity during externally focused attention (Mason et al., 2007;
McKiernan, Kaufman, Kucera-Thompson, & Binder, 2003; Li,
Yan, Bergquist, & Sinha, 2007). McKiernan et al. (2003) demon-
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strated that the more that attentional resources are focused toward
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Figure 6. Medial temporal lobe sensitivity to attentional orientation and a visuospatial task, the greater the deactivation of DMN regions,
the memory-evoking nature of the music. The means of the estimated beta including the anterior and PCC, compared with rest. Performance
values for each task condition are shown separately for different MTL of a novel task also leads to more deactivation in MPFC, PCC, and
subregions within each brain hemisphere. Significance value markers: other DMN regions, compared with the performance of a practiced

p ⬍ .05. ⴱⴱ p ⬍ .01. ⴱⴱⴱ p ⬍ .001. L⫽ left; R ⫽ right; Hipp ⫽ task (Mason et al., 2007). The authors explain the results as
hippocampus; DG ⫽ dentate gyrus; EC ⫽ entorhinal cortex; SUB ⫽ evidence of the increasing external attention demands of a novel
subiculum complex; MTL ⫽ medial temporal lobe; ROIs ⫽ regions of
task inhibiting task-unrelated self-referential processes in the
interest.
DMN. The MPFC and PCC are central hubs of the DMN and
integrate information between DMN subsystems and the FPCN
during the construction and elaboration of autobiographical mem-
response to the prestimulus cues, and responses to the three post- ories (Andrews-Hanna, Reidler, Sepulcre, Poulin, & Buckner,
stimulus questions unique to each cue-type indicated that partici- 2010; St. Jacques, Kragel, & Rubin, 2011). Given that greater
pants successfully performed the tasks as instructed. The coarse MPFC involvement in memory processing is associated with in-
comparison of memory-evoking music with nonevoking music creased self-referential processing (Cabeza et al., 2004) and in-
conformed to expectations based on previous research in which creased PCC activity is associated with both increased sense of
ratings of autobiographical salience, vividness of mental imagery, reliving (Daselaar et al., 2008) and vividness of recalled events
and familiarity ratings were significantly higher for memory- (Gilboa, Winocur, Grady, Hevenor, & Moscovitch, 2004), deacti-
evoking music (Barrett et al., 2010; Janata, 2009; Janata et al., vation of these regions should impact the richness of the recollec-
2007; Figure 2). Of greater importance was the fact that directing tion experience. The present results suggest that focusing attention
attention toward memories significantly increased MEAM salience externally during music listening decreases self-referential activity
and vividness ratings, relative to the task of focusing attention in DMN regions critical for autobiographical recall and corre-
toward the musical structure, but only for memory-evoking music. sponds to a decrease in reported reliving experiences of MEAMs.
Although salience ratings for attend-memories trials were compa- The focus of attention during memory-evoking music trials also
rable to salience ratings for the same memories spontaneously modulated functional connectivity between and within brain net-
recalled months before the scanner study, ratings for memory- works. The decrease in FPCN activity when attending to memories
evoking music reported after attend-music trials were significantly was concurrent with the weakening of negative time series corre-
diminished compared with previous ratings. Participants’ familiar- lations between prefrontal FPCN and DMN communities, as well
ity ratings were not influenced by Attentional Focus, suggesting as a more sparsely connected FPCN (Figure 4). The elimination of
that phenomenological dimensions directly related to the experi- negative connectivity between prefrontal DMN and FPCN regions
ence of reliving autobiographical memories (Sutin & Robins, while attending memories may represent the engagement of spe-
2007) can be modulated by attention. Overall, the behavioral cific prefrontal regions (Daselaar et al., 2008; Svoboda et al.,
results replicated the results from Experiment 1, affirming our a 2006) and the formation of process-specific alliances to aid in
priori interest in examining the influence of Attentional Focus on retrieval (Cabeza & Moscovitch, 2013). On the other hand, the
the brain networks engaged by memory-evoking music. increased interconnectivity within FPCN and negative connectiv-
Attention modulates DMN activity during MEAMs. ity with DMN when attending to music resemble anticorrelated
Attentional Focus accounted for the majority of task-related vari- patterns of resting-state connectivity (Fox et al., 2005).
ability in brain activity and modulated DMN and MTL regions The increased negative correlations between networks when the
previously associated with MEAMs and likability of music task requires actively ignoring memories may also be indicative of
(Janata, 2009; Wilkins et al., 2014). Attending to memories was top-down suppression of memory regions engaged by spontane-
associated with greater hippocampal activity compared with base- ously triggered MEAM content. During a task in which partici-
line, whereas attending to music led to decreases in DMN and pants had to actively ignore a memory for picture pairs, Benoit,
increases in medial and lateral premotor areas (see Tables S4 and Hulbert, Huddleston, and Anderson (2015) found that increased
S5 in the online supplemental materials). The direct contrast be- negative top-down coupling between the DLPFC and hippocam-
tween attention conditions showed the predicted effects clearly. pus was associated with a reduction in involuntarily recalled
Focusing attention on memories resulted in greater activity within memories. Although we did not find negative connectivity be-
ATTENTION AND MEMORY-EVOKING MUSIC 95
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Figure 7. Influence of Attentional Focus on the functional connectivity of the left hippocampus during trials
involving memory-evoking music. The yellow sphere marks the location of hippocampal seed (6-mm radius)
used for the analysis. TFCE ⫽ threshold-free cluster enhancement. See the online article for the color version
of this figure.

tween MTL regions and the DLPFC, the right DLPFC demon- MTL, rather than the MPFC community these ROIs belonged to
strated greater negative correlations with MPFC when attention during the resting state. These results suggest that when attention
was directed away from MEAMs, consistent with the role of the is directed toward autobiographical content, regions implicated in
lateral prefrontal cortex in manipulating (Blumenfeld & Ranga- the elaboration of autobiographical memories (Berryhill, Phuong,
nath, 2006) and suppressing the contents of memory (Benoit & Picasso, Cabeza, & Olson, 2007; Daselaar et al., 2008) and auto-
Anderson, 2012). biographical planning of events (Spreng et al., 2010) demonstrate
The differences between ROIs within known functional brain a concomitant increase in communication with the MTL. When
networks described above were made more succinct by consider- attention was directed away from salient autobiographical associ-
ing the community structure of these brain areas as a function of ations, these lateral posterior DMN ROIs instead shared commu-
the participant’s experiential state. The presence of autobiograph- nity membership with frontal DMN regions. MEAMs were rated
ical associations failed to evoke unique community structures as less salient and imagery as less vivid when attention was
within the DMN (Figure 5A), supporting our observations that directed toward music rather than memories, suggesting that di-
Attentional Focus had a greater influence over brain activity than versity within the DMN and MTL community structure is impor-
did Memory Status. tant for the construction of coherent and salient memories. Lateral
Of particular interest was the way in which Attentional Focus temporal and AG regions provide semantic information about
influenced community structure of the DMN when participants people and places (Svoboda et al., 2006), and posterior midline
listened to music that evoked autobiographical associations. When DMN regions serve to integrate spatial, temporal, and causal
attending to memories, ROIs including the left AG, MTG, and information within an episode into a coherent event (Ranganath &
anterior temporal pole shared community membership with the left Ritchey, 2012). The specialization within DMN regions likely
96 KUBIT AND JANATA

Table 3
Summary of Significant Clusters Correlated With Left Hippocampal Time Series During
Memory-Evoking Attend-Memory Trials

Left hemisphere (mm) Right hemisphere (mm)


Brain regions BA x y z Nvox Cmass x y z Nvox Cmass

Frontal
MPFC 10 6 58 18 10 1,392
MPFC 10 6 60 12 1 1,088
Temporal
STG 22 58 ⫺26 3 9 1,164
pSTGa 22 50 ⫺30 12 1 1,092
pSTGb 22 48 ⫺32 14 1 1,113
Parietal
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RSC 30 ⫺9 ⫺53 7 9 1,124 8 ⫺57 20 182 828


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PCC 23/31 ⫺8 ⫺58 20 61 1,711


31 ⫺12 ⫺62 12 1 1,072
31 ⫺6 ⫺67 16 9 1,094
Occipital
Ext. C 37 ⫺46 ⫺70 ⫺10 1 1,149
Subcortical
Dorsal striatum ⫺17 13 1 548 2,531 20 14 ⫺1 607 3,222
⫺14 ⫺8 18 22 1,246
Ventral striatum 8 8 ⫺6 18 1,229
Note. BA ⫽ Brodmann Area; Nvox ⫽ #voxels; Cmass ⫽ cluster mass; MPFC ⫽ medial prefrontal cortex;
STG ⫽ superior temporal gyrus; pSTGa ⫽ posterior STG a; pSTGb ⫽ posterior STG b; RSC ⫽ retrosplenial
cortex; PCC ⫽ posterior cingulate cortex; Ext. C ⫽ extrastriate cortex.

reflects the granularity of the DMN (Andrews-Hanna et al., 2010) ever, when attending memories the same representations are used
and the integration of multiple sources of information during the as a context in which to search for related autobiographical con-
reconstruction of an autobiographical memory (Conway & tent. Given the extensive evidence that memory for music, for
Pleydell-Pearce, 2000). example, for familiar melodies, is supported by sensorimotor re-
Attention modulates sensorimotor activity during MEAMs. gions including the SMA and pre-SMA (Halpern & Zatorre, 1999;
The dynamic temporal and tonal patterns in a piece of music are Jacobsen et al., 2015; Leaver et al., 2009), we hypothesize that
the basis for a very different type of memory cue compared during memory-evoking attend-memory trials the left MTL and
with the pictures or words that are commonly used in studies of MPFC communities likely represent information unique to evoked
autobiographical memory. Unlike static pictures, music can be autobiographical content, whereas the community including the
thought of as sequences of complex sounds structured in time. pre-SMA regions represents information particular to the music
Consequently, the autobiographical cue a piece of music provides itself. These results demonstrate the attention-induced flexibility of
consists of information that changes as the song unfolds over time. these brain regions and highlight the utility of thinking about
Subcortical and cortical sensorimotor regions that support interval MEAMs in association with a sensorimotor representation for the
timing (Schaefer, 2014) and the consolidation of nonmusical pro- music that is distinct from representations of related autobiograph-
cedural knowledge (Censor et al., 2014; Willingham, 1998) are ical and episodic content.
also implicated in the perception of and memory for music (Lidén The left IFG also plays an important role in processing se-
et al., 2016; Jacobsen et al., 2015; Janata & Grafton, 2003). quences of information in musical and language perception (Koel-
During trials in which attention was directed toward searching sch, 2011). In the present study, bilateral IFG increased activity
for associated memory content, sensorimotor regions responsible alongside sensorimotor regions during attend-music trials. Al-
for processing tonal and rhythmic patterns formed a unique com- though the IFG and pre-SMA are anatomically connected and
munity structure. The somatosensory cortex, pre-SMA, and mid- typically coactivated during musical imagery tasks (Alonso et al.,
line motor regions extending to the MCC formed a network 2016; Halpern & Zatorre, 1999), research on cognitive control
distinct from DMN and FPCN communities found during other mechanisms has also linked the IFG to the suppression of mem-
conditions (see Figure 5B red outlines, slices x ⫽ ⫺7 and 5). ories (Benoit et al., 2015), and this area is known to play an
During trials when attention was directed externally, away from important role in inhibiting response tendencies in nonmemory
memory search, the sensorimotor regions failed to form a unique tasks (Aron, Robbins, & Poldrack, 2014), particularly the right
community, instead showing increased connectivity with lateral IFG (but see Swick, Ashley, & Turken, 2008). In the present study,
temporal, prefrontal, and posterior midline DMN ROIs. We pro- bilateral IFG regions increased activity on trials in which MEAMs
pose that this reconfiguration in community structure is best un- were meant to be ignored, and only the right IFG demonstrated an
derstood by considering the different roles representations of mu- increase in negative connectivity with the MPFC community, in
sic serve in the two task conditions. When attending to the music, line with its proposed role in memory inhibition. Future work is
the representations of the temporal and melodic structures are used needed to disentangle the role of the IFG in music perception and
to inform a decision on the number of instruments present; how- cognitive control.
ATTENTION AND MEMORY-EVOKING MUSIC 97

Attention modulates MTL activity during MEAMs. Our shown to correlate with intensely pleasurable responses to person-
results support the interpretation that MTL regions were not ob- ally meaningful music (Salimpoor, Benovoy, Larcher, Dagher, &
served during the experiencing of MEAMs in Janata (2009) be- Zatorre, 2011). Though emotional and reward processes likely
cause of the spontaneous nature of the evoked memories and shaped the reliving of MEAMs in the present study, most of the
absence of task demands that mandated memory retrieval. Corrob- striatal activity was found in the dorsal striatum, including bilateral
orating the findings of Ford et al. (2011), we found increased MTL putamen and caudate nucleus. A recent study by Ford et al. (2016)
activity during active search for MEAMs, particularly in the left found that activity in the dorsal striatum tracked familiarity ratings
hippocampus (Figure 6). Directing attention toward structural as- of the musical cues used in an explicit autobiographical memory
pects of music that triggered MEAMs significantly reduced activ- retrieval task. Given the dorsal striatum’s well-documented role in
ity within the hippocampus compared with focusing on the temporal predictions during the performance and perception of
MEAMs. Although the hippocampus demonstrated activity near music (Kung, Chen, Zatorre, & Penhune, 2013; Zatorre, Chen, &
the resting-state baseline, the behavioral data indicated that par- Penhune, 2007), as well as anticipation of highly pleasurable
ticipants did indeed experience salient MEAMs during attend- chill-evoking moments (Salimpoor et al., 2011), the striatal acti-
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music trials. Interestingly, greater autobiographical salience of a vations in the present study are likely related to representations of
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piece of music corresponded to a significant increase in the prob- the musical cue used for retrieval. Interactions between the hip-
ability of correctly identifying the number of instruments in an pocampus and striatum are known to influence the consolidation of
excerpt (Table S2 in the online supplemental materials). Thus, procedural memories (Albouy et al., 2015; Albouy, King, Maquet,
spontaneous MEAM retrieval, in addition to accessing a represen- & Doyon, 2013), and current models of spatial navigation empha-
tation of familiar music to facilitate a focused perceptual analysis, size contributions from distinct hippocampal- and striatal-
appears to occur without associated activity increases within the dependent learning strategies that are mediated by the MPFC
hippocampus. (Chersi & Burgess, 2015). The present study shows an influence of
Overall, the present results suggest that the hippocampus, attentional focus on the interactions between the left hippocampus,
thought to be a core region supporting non–music-related autobi- bilateral striatum, and MPFC during memory-evoking music. We
ographical memories (Svoboda et al., 2006), is highly susceptible suggest that the role of the hippocampus during the perception of
to attentional modulation, even when the participant experiences autobiographically salient music is to integrate sensorimotor rep-
autobiographical associations. Although directing attention toward resentations of familiar music with episodic and autobiographical
MEAMs resulted in increased subjective reliving of autobiograph- information in the DMN—a role that depends on the attentional
ical memories, this increase cannot be explained entirely by in- focus of the listener. The binding of multiple sources of informa-
creased hippocampal activity. Despite the fact that participants tion may represent the use of music to help identify and recall a
reported no autobiographical salience during nonevoking attend- specific lifetime period and event in one’s past. This account is
memory trials, searching for memories led to an increase in hip- very similar to accounts of episodic memory retrieval in which the
pocampal activity above resting-baseline that was absent when role of the hippocampus is to bind items with contextual informa-
attention was directed away from MEAMs. The present study tion (Eichenbaum, 2000; Ranganath & Ritchey, 2012) and recon-
shows that attentional processes are an important source of vari- struct vivid events (Gilboa et al., 2004).
ability in the MTL, adding to findings that variables such as a Neurological evidence demonstrates memory for a piece of
person’s emotional state during recall (LaBar & Cabeza, 2006) and music remains despite impairments in episodic retrieval (Huijgen
age of a memory (Piefke, Weiss, Zilles, Markowitsch, & Fink, et al., 2015). Recent research on patients with Alzheimer’s disease
2003) also determine the degree of MTL activity. (AD) has shown both spared musical memory and structural in-
We hypothesize that when attending to MEAMs, sensorimotor tegrity of pre-SMA (Jacobsen et al., 2015), a brain region anatom-
regions provide information unique to a piece of music, and that ically connected to the dorsal striatum (Vergani et al., 2014).
the hippocampus uses this information to aid in the construction of Although AD patients show impairments in explicit recollection of
autobiographical memories. Ignoring a memory by focusing atten- memories concurrent with atrophy in the MTL and DMN, implicit
tion on the instruments in a musical excerpt changes the connec- procedural memories often remain intact (Beaunieux et al., 2012;
tivity of sensorimotor regions and is likely to reduce the extent Eldridge, Masterman, & Knowlton, 2002; Willingham, Peterson,
music serves as a cue for autobiographical information, and as a Manning, & Brashear, 1997). Recent research has also shown
result, may explain the decrease in salience and vividness of evidence for spared MEAMs throughout the progression of AD
memories evoked under such conditions. In support of this hy- (Cuddy et al., 2015), which suggests memory for music and
pothesis, we found that directing attention toward MEAMs in- associated autobiographical information may interact without
creased correlations between the left hippocampal time series and MTL involvement. These results are consistent with previous
clusters in DMN, auditory cortex, and sensorimotor regions in- results demonstrating the absence of MTL activity during sponta-
cluding bilateral striatum (Figure 7; Table 3), which we interpret as neously evoked memories (Janata, 2009). We believe that the
stronger integration of sensorimotor and episodic memory retrieval interaction between procedural and episodic memory systems may
processes. When Attentional Focus was directed away from mem- underlie the preservation of MEAMs despite difficulty remember-
ories or when no memory was found despite searching for one, the ing other episodic information in response to less specific or
hippocampal time series did not significantly correlate with any sensorially impoverished cues. Future research on the salience and
other regions beyond those correlations observed during rest. vividness of MEAMs in AD patients may clarify the role of the
Activation of the ventral striatum during recollection of autobi- MTL. Although neurological and neuroimaging evidence suggests
ographical memories has previously been attributed to reward MTL involvement isn’t necessary, the present results suggest the
processes (Speer, Bhanji, & Delgado, 2014) and has also been hippocampus plays a role in the elaboration, and perhaps the
98 KUBIT AND JANATA

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This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

communication between resting-state networks and integration Benoit, R. G., & Anderson, M. C. (2012). Opposing mechanisms support
This document is copyrighted by the American Psychological Association or one of its allied publishers.

across multiple memory systems during trials in which participants the voluntary forgetting of unwanted memories. Neuron, 76, 450 – 460.
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