Medial Prefrontal Cortex: Adding Value - IA de Investigación especializada en el MIT

Medial Prefrontal Cortex: Adding Value
to Imagined Scenarios

Wen-Jing Lin, Aidan J. Horner, James A. Bisby, and Neil Burgess

Abstracto

■ The medial prefrontal cortex (mPFC) is consistently impli-
cated in the network supporting autobiographical memory.
Whereas more posterior regions in this network have been re-
lated to specific processes, such as the generation of visuospa-
tial imagery or the association of items and contexts, el
functional contribution of the mPFC remains unclear. Cómo-
alguna vez, the involvement of mPFC in estimation of value during
decision-making suggests that it might play a similar role in
memory. We investigated whether mPFC activity reflects the
subjective value of elements in imagined scenarios. Participantes
in an MRI scanner imagined scenarios comprising a spatial con-
texto, a physiological state of need (p.ej., thirst), and two items that
could be congruent (p.ej., drink) or incongruent (p.ej., alimento) con
the state of need. Memory for the scenarios was tested outside
the scanner. Our manipulation of subjective value by imagined

need was verified by increased subjective ratings of value for
congruent items and improved subsequent memory for them.
Consistent with our hypothesis, fMRI signal in mPFC reflected
the modulation of an item’s subjective value by the imagined
physiological state, suggesting the mPFC selectively tracked
subjective value within our imagination paradigm. Further anal-
yses showed uncorrected effects in non-mPFC regions, incluir-
ing increased activity in the insula when imagining states of
need, the caudate nucleus when imagining congruent items,
and the anterior hippocampus/amygdala when imagining subse-
quently remembered items. We therefore provide evidence that
the mPFC plays a role in constructing the subjective value of the
components of imagined scenarios and thus potentially in
reconstructing the value of components of autobiographical
recollection. ■

INTRODUCCIÓN

Autobiographical memories (AMs) define who we are
and depend on a network of brain regions including
the hippocampus, parahippocampal gyrus, retrosplenial
corteza, posterior parietal cortices, and medial prefron-
tal cortex (mPFC; p.ej., Addis, Moscovitch, Crawley, &
McAndrews, 2004; Piolino et al., 2004; Maguire, 2001; Nadel
& Moscovitch, 1997). Research into the neural mecha-
nisms underlying AM has focused on closely related con-
cepts of imagery for spatial context (p.ej., Burgess, Maguire,
& O’Keefe, 2002), “scene construction” (p.ej., Hassabis,
Kumaran, & Maguire, 2007), “episodic future thinking”
(p.ej., Addis, Wong, & Schacter, 2007), “self-projection”
(Buckner & Carroll, 2007), and item-to-context binding
(Eichenbaum, Yonelinas, & Ranganath, 2007). Además
to the long-recognized hippocampal role in AM (Howard &
Eichenbaum, 2013; Squire & Zola-Morgan, 1991; O’Keefe &
Nadel, 1978; Scoville & Milner, 1957), this research has pro-
posed specific functional roles for posterior brain regions.
The parahippocampus, retrosplenial cortex, and the rest of
Papez’s circuit have been ascribed roles in the genera-
tion of visuospatial imagery (Byrne, Becker, & Burgess,
2007), whereas medial-temporal regions have been impli-
cated in storing items and context beyond the spatial

University College London

domain (Eichenbaum et al., 2007). Además, lateral pa-
rietal and prefrontal areas have been ascribed roles in atten-
tional and working memory components of AM tasks
( Johnson, suzuki, & Rugg, 2013; Rugg & Vilberg, 2013;
Cabeza, Ciaramelli, Olson, & Moscovitch, 2008; simons
et al., 2008; Wagner, shannon, Kahn, & Buckner, 2005).

Sin embargo, less is known regarding the functional role
of mPFC in AM. In decision-making, mPFC responses are
believed to represent the subjective value of chosen items
relative to potential alternatives (Rushworth, Noonan,
Boorman, Walton, & Behrens, 2011). Activity in mPFC is
correlated with the value of the chosen item, irrespective
of whether the items are food (Gross et al., 2014; Hare,
Camerer, & Rangel, 2009), agua (Bouret & Richmond,
2010), monetary reward (Nicolle et al., 2012; Boorman,
Behrens, lana rica, & Rushworth, 2009), physical action,
engaging activities (Gross et al., 2014), or abstract figures
(Glascher, Hampton, & O’Doherty, 2009). The mPFC is
also associated with self-referential thought, incluido
memory (Levin, 2004; Macrae, Moran, Heatherton,
Banfield, & kelly, 2004; Vogeley et al., 2004; Johnson
et al., 2002; Gusnard, Akbudak, Shulman, & Raichle, 2001),
leading to the recent suggestion that ventromedial pFC
(vmPFC) helps to establish the personal value, affective
quality, or significance of self-related information (Benoit,
Szpunar, & Schacter, 2014; D’Argembeau, 2013; Lebreton
et al., 2013).

© 2015 Massachusetts Institute of Technology Published under a
Creative Commons Attribution 3.0 no portado (CC POR 3.0) licencia

Revista de neurociencia cognitiva 27:10, páginas. 1957–1967
doi:10.1162/jocn_a_00836

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Given the association between memory and imagery, él
is interesting that imagery can interact with subjective
value and can influence our motivation for satisfying
basic needs, such as food consumption (larson, Redden,
& Elder, 2014; Morewedge, Huh, & Vosgerau, 2010).
Además, imagining future scenarios can influence
decision-making by changing the subjective value of
choices (Lebreton et al., 2013; Benoit, Gilbert, &
Burgess, 2011; Peters & Büchel, 2010). De este modo, imagining
oneself in a hungry state may raise the subjective value
of food items. En cambio, human memory can be influ-
enced by the value or motivational salience of the to-be-
remembered stimuli (Erwin & Ferguson, 1979). Para
instancia, fasting people have enhanced memory for food
pictures (morris & Dolan, 2001). De este modo, we infer that
memory for items could also be modulated by their value
in imagined scenarios.

Building on these previous studies, we hypothesized
that mPFC plays a role in AM and self-related imagery by
providing the subjective value of elements of a scene—a
function not ascribed to more posterior parts of the AM
network. To test this hypothesis, we designed a paradigm
in which the subjective value of items within imagined
scenarios could be manipulated experimentally during
resonancia magnética funcional. We required participants to imagine being in a cur-
rent context and state (as opposed to imaging a future
scenario, see Benoit et al., 2014) and subsequently imag-
ine seeing, but importantly not consuming (cf. Bruto
et al., 2014), objects that were congruent or incongruent
with the imagined state of need. We reasoned that the
imagined current state of need would modulate the sub-
jective value of the unconsumed objects and that mPFC
activity would correlate with this state-modulated subjec-
tive value.

MÉTODOS

Participantes

Twenty right-handed participants were recruited from
the University College London student population. Uno
did not finish the task, so the data reported here concern
the remaining 19 Participantes (12 women). The mean age
of the remaining participants was 21.7 años (DE = 2.68,
range = 19–27). All participants gave written informed
consent to participate, in accordance with the local ethics
committee (1825/003). One participant did not complete
the postscan memory task, so the results from the mem-
ory analyses are based on 18 Participantes.

Stimuli and Design

Four different physiological states of need were used:
thirst, coldness, hunger, and tiredness. A neutral state
was used as a baseline condition (instruction for neutral
estado: Imagine you are just fine. You are not in any state

of need but just in an ordinary condition.). Twelve spatial
contexts were used: beach, cocina, desert, campos, class-
habitación, airplane, forest, oficina, library, playground, church,
and ship. These were included to make the imagined sce-
narios more realistic and because, without instruction,
participants would be likely to imagine uncontrolled
backgrounds to facilitate imagery. Había 60 state–
context combinations, with each appearing only once
durante el 60 trials of the imagery task.

Pictures from four categories were used as items; cada
category contained items that were usually used to satisfy
one of the four physiological states of need. The first cat-
egory contained water, juice, beer, and other beverages
used to quench thirst. The second category contained
items that were able to be used to help people resist cold
clima, such as fireplace, hot drink, and winter clothes.
Another category contained food, and the final category
contained items used for taking a rest or relieving tired-
ness included a bed, couch, bathtub, etcétera. Allá
eran 180 item pictures in total, consisting of 45 pictures
per category. Among these pictures, 120 appeared in the
imagery task and another 60 served as new items during
an old–new recognition test. The assignment of pictures
to old items and new items was counterbalanced across
Participantes. All pictures were obtained from FreeDigital-
Photos.net (www.freedigitalphotos.net/).

In the imagery task, each trial contained one state–context
combination presented as cue words and also two item
pictures (see Figure 1A for an example of trial presenta-
tion order). The relationship between the participant’s
current imagined state and each item picture during a
single trial could either be congruent or incongruent.
For a congruent item, the type of item presented would
meet the participant’s current need created by the imag-
ined state. Por ejemplo, a food picture would be classified
as congruent if the state was hunger, but incongruent if
the state was tired, cold, or thirsty. Note that “incongru-
ent” items were irrelevant rather than opposite to the
current state of need. Ambiguous items were never used
as “incongruent items” (p.ej., a hot drink was not used
in thirst trials). From the two item pictures, sequentially
presented during each trial, either item could be congru-
ent or incongruent with the current state. This provided
four possible combinations: congruent–congruent,
incongruent–incongruent, congruent–incongruent, y
incongruent–congruent. En tono rimbombante, all four combina-
tions of items occurred in pseudorandom order across
ensayos, allowing us to identify the effects of an individual
items’ subjective value, as modulated by its congruency
with the imagined state. Among the 120 item pictures
presented during the imagery task, 24 served as neutral
pictures as they occurred in a neutral state. An alternative
would be to use items unrelated to any of the physiolog-
ical states, but such items would be intrinsically different
to the congruent items in the study. El restante 96
pictures were equally assigned as congruent or incongru-
ent items.

1958

Revista de neurociencia cognitiva

Volumen 27, Número 10

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Cifra 1. Procedure of the imagery task (A) and the memory task (B).

Procedimiento

Imagery Task

Participants were provided with task instructions before
scanning and completed a number of practice trials out-
side the scanner. The entire imagery task, consisting of
60 imagery trials, was equally divided into two sessions,
and scanning lasted for 1 hr in total, including acquisition
of a structural scan. See Figure 1A for an illustration of
stimulus presentation for the imagery task. Each trial be-
gan with a fixation cross at the center of the screen,
which was replaced by a pair of state–context cue words
después 0.5 segundo. Participants were instructed to vividly imag-
ine the context and state according to the cue words pro-
vided. The state–context cue words were presented for
4 segundo, and then a fixation cross appeared again (for 8–
12 segundo, jittered), during which the participants were in-
structed to continue imagining. Próximo, two pictures were
presented sequentially, each for 4 sec separated by a
0.5-sec blank screen. Participants were required to incor-
porate each presented item into their imagined scenario
during the trial. Participants were explicitly instructed
to not imagine consuming these items to satisfy their
imagined state and its associated need. Por ejemplo, ellos
were required to imagine seeing (but not consuming) a
chicken burger in a forest while they were thirsty (as in
Figura 1A). After a further blank screen (1–4 sec, jit-
tered), participants made four simple ratings, one at a
tiempo. The first two ratings asked participants to rate
how much they had wanted each item when they initially
saw it during the trial. The last two separately rated how
vividly they had imagined the current state and context.
All ratings used a 4-point scale (1 = not at all, 4 = very
mucho). Each trial ended with a final blank screen (3–
6 segundo, jittered). Visual stimuli were presented by MATLAB
(The MathWorks, Natick, MAMÁ) and COGENT 2000 toolbox
(www.vislab.ucl.ac.uk/cogent.php).

Memory Task

The memory task took place outside the scanner after
the imagery task was completed. Each trial consisted of
a 500-msec fixation cross followed by a picture of an item,
and participants were required to judge whether the pic-
ture had been presented in the imagery task or not (es decir.,
old/new item recognition judgment) and how confident
they were of their answer (Figure 1B shows an illustration
of the memory task). If participants answered “new,” par-
ticipants were then asked how much they like that item
in their daily lives. If an item was judged “old,” two fur-
ther source memory questions were presented to the
participant to test memory for the associated state and
contexto. To test state, one of the state words (hunger,
thirst, tired, cold, or neutral) was presented, and partici-
pants judged whether that state was the one they had
been asked to imagine when the recognized item picture
had appeared in the imagery task. The correct answer
was yes for 50% of trials, and within these trials, 40% de
the state words were congruent with the tested item,
40% were incongruent, y 20% were neutral. Para el
context source memory test, todo 12 of the contexts were
listed to allow participants to select the one which had
accompanied the recognized item picture. The trial ended
with the daily subjective rating. Había 180 memory
trials in total (120 with “old” items and 60 with “new”
elementos). Twelve alternative forced choice is an efficient
way to test memory for the spatial context of an item’s
presentation but could not be used to test memory for
the physiological state, because a simple strategy of
guessing the congruent state would artificially inflate per-
rendimiento (p.ej., choosing “thirst” when presented with a
drink). In this situation, choosing a congruent state
would be correct in 40% of trials, a neutral state would
be correct in 20% of trials, and the three incongruent
states would be correct in 13% of trials. To avoid this,

Lin et al.

1959

we tested participants with yes/no cued recognition of a
single state that was chosen to be correct 50% del
tiempo, irrespective of its congruence with the item.

fMRI Data Acquisition and Preprocessing

Functional imaging was performed on a 3T scanner
(Siemens TIM Trio, Siemens, Berlina, Alemania) durante
the imagery task. The functional data were acquired with
a gradient-echo EPI sequence (repetition time = 3.36 segundo,
echo time = 30 mseg, flip angle = 90°, resolution = 3 × 3 ×
3 mm, 64 × 74, 48 slices per volume). El numero total de
volumes in each run varied across participants because of
the variation of RT for each rating (the mean number of vol-
umes was 332 per session). A high-resolution T1-weighted
3-D structural image (1 mm3) was acquired after two ses-
sions of functional scans. A double-echo FLASH fieldmap
sequence was also recorded.

Functional images were processed and analyzed with
SPM8 (Wellcome Trust Centre for Neuroimaging, Londres,
Reino Unido, www.fil.ion.ucl.ac.uk/spm/software/spm8/). The first
five volumes of each scan were discarded for T1 equilibra-
ción. Preprocessing procedures included bias correction,
realignment, unwarping, coregistration, slice timing cor-
rection, and normalization to the MNI template using the
Dartel toolbox. EPI images were smoothed with an iso-
tropic 8 mm FWHM Gaussian kernel. One of the partici-
pant’s fieldmap scan was not collected, so the unwarping
procedure was skipped in their data.

Análisis de los datos

The preprocessed functional images were analyzed with
general linear models (GLMs). We estimated five GLMs
for different purposes. All GLMs included six movement
regressors for each session, estimated during realign-
mento, as well as two further regressors modeling each
session. On the basis of our strong a priori hypothesis
about the mPFC and vmPFC, we performed small-volume
correction (SVC) within a combined anatomical mask
of these regions: bilateral mPFC and vmPFC (volumen
∼ 53,493 mm3). This mask was derived from the AAL atlas
(Tzourio-Mazoyer et al., 2002), as implemented in
the WFU PickAtlas Tool (Maldjian, Laurienti, kraft, &
Burdette, 2003). This mask contained superior frontal gy-
rus, medial frontal gyrus, anterior cingulate, and cingulate
gyrus. Within this small volume, we report effects that sur-
vive p < .05 FWE correction. For completeness, we also report effects at p < .001 uncorrected across the whole brain; however, caution is needed in interpretation of these effects. The first model (GLM1) was a parametric modulation analysis, searching for regions that correlated with the subjective value of an item during imagined states of need. The first-level model contained seven regressors per session: (1) imagining a state of need, (2) imagining a neutral state, (3) imagining an item in a state of need, (4) a parametric modulator of the item regressor based on the participant’s subjective value of each item, (5) imagin- ing an item in a neutral state, (6) intertrial interval (ITI) periods, and (7) key presses. Trial periods were modeled with a boxcar function for the entire length of each period (e.g., the 4 sec of imaging an item), convolved with the canonical hemodynamic response function. The second- level analysis was a one-sample t test on the parameter estimates from the parametric modulator (Regressor 4) averaged across the two sessions. For the parametric modulation, we used the subjective rating of each item when imagined in the state of need of the current trial mi- nus the subjective rating of the item in the participant’s daily life, given after the scanning session. This calculation allowed us to control for variations in the participants’ baseline preference for the various items. The range of these normalized subjective ratings was from −3 to 3. The second model (GLM2) was used for comparing imagination of congruent items versus incongruent items (given that the first GLM collapsed across these condi- tions to maximize power in our parametric modulation analysis) and also for comparing imagining states of needs versus neutral states. This model included seven regressors per session: (1) imagining a state of need, (2) imagining a neutral state, (3) imagining a congruent item in a state of need, (4) imagining an incongruent item in a state of need, (5) imagining an item in a neutral state, (6) ITI periods, and (7) key presses. Parameter es- timates for regressors (1) to (4) were averaged across the two sessions and entered into a second-level model. A separate regressor was also included for each individual subject that consisted of a “1” for each condition for that specific participant (i.e., subject effects). A third model (GLM3) aimed to test the subsequent memory effect for imagined items. The model was similar to GLM1 but replaced the subjective value parametric modulator with a modulator based on subsequent memory. The model included six regressors per session: (1) imagining a state of need, (2) imagining a neutral state, (3) imagining an item (in either a state of need or neutral state), (4) a para- metric modulator of the previous regressor based on subsequent memory for the item, (5) ITI periods, and (6) key presses. Note that the parametric modulator for subjective value was applied to item imagination during a state of need, not during neural states, as we were specif- ically interested in how states of need modulated subjec- tive value. The parametric modulator for subsequent memory was applied to all item imagination trials (includ- ing neutral states) to maximize power. Subsequent mem- ory was parameterized as a transformed confidence rating to maximize sensitivity. Participants’ 1–4 confidence rat- ings for old and new items at test were transformed into a measure of successful memory performance by com- bining ratings for item “hits” with negative ratings for item “misses” (e.g., a “miss” given a confidence rating of 4 would become −4 in the parametric modulator). The second-level analysis was a one-sample t test on 1960 Journal of Cognitive Neuroscience Volume 27, Number 10 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t f . / u s e r o n 1 7 M a y 2 0 2 1 the parameter estimates from the parametric modulator (Regressor 4) averaged across the two sessions. The final two models (GLM4 and GLM5) aimed to test the subsequent memory effect for the state of need (GLM4) and the context (GLM5) in which items were imagined (i.e., two types of source memory). GLM4 con- tained seven regressors per session: (1) imagining a state of need, (2) imagining a neutral state, (3) item imagina- tion trials for which the item and state of need are sub- sequently remembered, (4) item imagination trials for which the item but not the state is remembered, (5) item imagination trials for which the item is not remembered, (6) ITI periods, and (7) key presses. GLM5 was similar to GLM4 but split item imagination trials (Regressors 3– 5) by whether the context (rather than the state) was re- membered. Second-level models for each GLM were paired t tests comparing either state or context hits ver- sus misses (Regressors 3 and 4) averaged across the two sessions. Note that we built separate GLMs for each analysis of interest. This was due to the overlapping nature of cer- tain regressors. In particular, the categorical congruent versus incongruent contrast correlated with the related, but more sensitive, item-by-item parametric modulation of value by state. Furthermore, the parametric modula- tors relating to subsequent memory and subjective value were also correlated. Despite the overlapping nature of these regressors of interest, our separate GLMs revealed distinct patterns of activity. RESULTS Behavioral Results The Subjective Value of Items in Imagery To demonstrate that our manipulation of imagined state worked, a three-way repeated-measure ANOVA with Situation (two levels: everyday rating and rating during imagery), Rating (1–4), and category (congruent, incongruent, and neutral) was performed. The three- way interaction was significant (F(6, 102) = 18.70, p < .001), so we performed further analyses that revealed that the distributions of ratings differed between categories for ratings during imagery (Rating × Category, F(6, 102) = 27.40, p < .001), but not for everyday ratings (Rating × Category, F(6, 102) = .66, p = .68). Thus, it was only when participants imagined being in a specific state of need that the subjective value of the objects differed between our “congruent” and “incongruent” conditions. Table 1 shows that a greater proportion of congruent items had positive subjective value (controlling for baseline value, i.e., rating of imagined value—everyday rating; 39.67%) whereas most incongruent items had negative subjective values (60.25%). This suggests that our participants indeed followed the instruction to imagine the assigned state of need and that those imagined states influenced the subjec- tive value of the item on that trial. We also carried out a two-way repeated-measure ANOVA with Congruency between the state question word and item (congruent and incongruent) and Rating (1–4) as within-subject variables to test whether the preceding state question might bias ratings (e.g., “hungry” increasing ratings for food items). There was no significant in- teraction between Congruency and Rating (F(3, 51) = .40; p = .75), suggesting that the everyday value ratings were not influenced by the preceding source memory questions. Old–New Recognition A one-way repeated-measure ANOVA across Congruency (congruent, incongruent, and neutral) was carried out to test for differences in hit rate among different categories of items. The results revealed a significant main effect of congruency (F(2, 34) = 9.01, p < .001; see Figure 2A for memory performance). Pairwise comparisons showed that hit rate was higher for congruent items than for in- congruent (t(17) = 5.16, p < .001) and neutral (t(17) = 3.14, p = .006) items. However, there was no signif- icant difference between incongruent and neutral items (t(17) = .35, p = .73). This result suggests that partici- pants had better memory for items that were able to ful- fill their needs in the imagined state. Participants showed a high correct rejection rate for new items (87%). Table 2 shows confidence ratings across all responses. For completeness, we checked whether our results varied with the order in which items were presented within a trial. We ran a two-way repeated-measure ANOVA with Order of presentation (two levels: first or second) and Category (three levels: congruent, incongruent, and neu- tral) as within-subject factors on the subjective ratings and subsequent memory scores. The results show that the order of presentation during encoding did not affect item memory (Order, F(1, 17) = .20, p = .66; Category, Table 1. Percentage of Subjective Values of Items during Imagery (−3 to +3), according to Whether They Were Imagined in a Congruent, Incongruent, or Neutral State of Need, Controlling for Baseline Value ( Value during Imagery—Everyday Value) Congruent Incongruent Neutral −3 2.36% 6.63% 6.35% −2 5.20% 18.64% 18.03% −1 17.66% 34.98% 30.90% 0 35.10% 27.31% 28.32% +1 23.27% 8.93% 11.43% +2 11.95% 2.94% 4.97% +3 4.45% 0.58% 0.00% Lin et al. 1961 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j . t / f u s e r o n 1 7 M a y 2 0 2 1 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t f . u s e r o n 1 7 M a y 2 0 2 1 Figure 2. Behavioral results for the memory task. (A) Mean values of hit rate in the item recognition memory task. (B and C) Mean performance in the source recognition task for the state of need (B) and the spatial context (C). Error bars represent ±1 SEM. F(2, 34) = 9.22, p = .001; Order × Category, F(2, 34) = 1.89, p = .17), and there was a nonsignificant trend to- ward lower ratings for the first item versus the second item (Order × Category, F(2, 34) = 1.05, p = .36; order, F(1, 17) = 3.92, p = .06). Source Memory Source memory performance for correctly associating the imagined state with the recognized item was analyzed using a one-way ANOVA across levels of Congruency. We found a significant main effect of Item congruency (F(2, 34) = 17.30, p < .001). Pairwise comparisons showed that the conditional state source performance hit rate (% correct source memory for the state associ- ated with items correctly recognized as “old”) for congru- ent items was significantly higher than for incongruent items (t(17) = 6.16, p < .001) and neutral items (t(17) = 5.44, p < .001), whereas there was no significant differ- ence between the latter two categories (t(17) = .17, p = .864; see Figure 2B). Although participants showed a response bias toward accepting the state (answering “yes”) when it was congruent (55.6% responses were yes) or neutral (54.2% yes) relative to the item and “no” when it was incongruent (41% responses for incon- gruent items were no), this response bias could not ac- count for our results (the correct proportion of “yes” responses being 50% in both cases). Analysis of source memory performance for the imag- ined spatial context (e.g., “beach”) within the recognized Table 2. Percentage of Hits, Misses, False Alarms, and Correct Rejections across Confidence Ratings (1–4) in the Old–New Item Recognition Task Hit 3.9 14.1 20.1 61.9 Miss 16.7 19.8 26.8 36.7 1 2 3 4 False Alarm Correct Rejection 17.0 29.1 29.1 24.8 10.4 15.3 23.5 50.7 item showed no significant main effect of Item congru- ency (F(2, 34) = .889, p = .42; see Figure 2C for context source memory performance). It is possible that this re- flects the irrelevance of spatial context to the subjective ratings that the participants are required to give on each trial or that any small effects of congruency on context– source memory were obscured by low levels of per- formance (chance = 8%) although performance was above chance in each category (congruent: t(17) = 3.96, p = .001; incongruent: t(17) = 4.48, p < .001; neu- tral: t(17) = 2.14, p = .047). In general, behavioral results supported our predic- tion. Subjective values of items support the validity of our imagined need paradigm. We also saw greater recog- nition performance for congruent than incongruent items and better memory for the imagined state of con- gruent than incongruent items. Thus, we observed better memory performance for items when their value was congruent with the imagined state. fMRI Results Subjective Value of Items in Imagery (GLM1) First, we focused on the main prediction of our study: that the subjective value of items in imagined scenarios would correlate with the BOLD response in the mPFC. To isolate imagined value from differences in the intrinsic values of the items used, we calculated the participant’s subjective value for the item when imagining it in the cur- rent state of need minus their subjective value for the same item in their daily life. This parametric modulator revealed an effect in the mPFC (+9, +57, +12, Z = 3.98; p < .05 FWE SVC). We therefore provide evidence that mPFC represents the values of elements in imagined scenarios, controlling for variations in their intrinsic value in other situations (Figure 3). Given the complexity of our imagination task, it is im- portant to rule out other explanations for our main mPFC result. This is particularly important given the overlapping nature of certain experimental factors (see Methods). In short, none of our subsequent analyses showed an effect 1962 Journal of Cognitive Neuroscience Volume 27, Number 10 Figure 3. The activation of mPFC during imagination of an object during a state of need varied according to the extent to which the subjective value of item was modulated by the imagined state of need. All peaks significant at p < .001, uncorrected (color bar indicates t statistic). (The red line depicts the area of mPFC mask used in SVC analysis.) in mPFC, even at a lenient p < .001 uncorrected thresh- old. However, these analyses did reveal effects in other re- gions at this threshold. We report these results for completeness but note that they should be treated with caution given that they do not survive correction for mul- tiple comparisons. and second item, finding a nonsignificant trend toward a greater effect of state congruency for the first versus sec- ond item in the vmPFC (−3, 33, −12; p = .083 FWE SVC). However, these could not influence the findings themselves, as our manipulation of state congruency was counterbalanced across items. Imagining States of Need and Item Congruency with Need (GLM2) Compared with imagination of a neutral state, imagina- tion of states of physiological need showed greater activa- tion in bilateral insula (MNI coordinates of peak activations: −39, −6, −3, Z = 3.27; +45, +15, +3, Z = 3.15; p < .001, uncorrected; Figure 4A). By contrasting imagery for congruent versus incongruent items, we iden- tified a region in the basal ganglia—the caudate nucleus (+3, +9, +6, Z = 3.60; −6, +9, +6, Z = 3.56, p < .001, uncorrected; Figure 4B). Because congruent items had higher subjective value than incongruent ones, we also car- ried out an SVC analysis for the congruent–incongruent contrast in the mPFC ROI but found no significant effect. We also investigated whether the fMRI correlates of an item’s value or state congruency varied between the first Subsequent Memory Effects (GLM3) This parametric modulation analysis showed that BOLD signal in the right amygdala (+33, −3, −30; Z = 3.27) and left anterior hippocampus (−21, −12, −18; Z = 3.33), when participants were imagining items, were sig- nificantly correlated with participants’ subsequent mem- ory ( p < .001, uncorrected; Figure 5). Note that our subsequent memory modulator combined categorical subsequent memory status (i.e., hits and misses) with subjective confidence, revealing linear increases in BOLD response from −4 (high confidence misses) to +4 (high confidence hits). No other significant activity was re- vealed in this analysis. No significant activations were found corresponding to subsequent source memory effects for state (GLM4) or for context (GLM5), that is, the comparisons of imagery Figure 4. (A) Bilateral insula showed higher activation when participants were imagining states of need compared to imagining the neutral state. (B) The caudate showed greater activation for imagining a state-congruent item than a state-incongruent item. All peaks significant at p < .001, uncorrected (color bars indicate t statistic). D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f t . / u s e r o n 1 7 M a y 2 0 2 1 Lin et al. 1963 Figure 5. Greater activity was seen during encoding in left hippocampus (top row) and right amygdala (bottom row) for items that were subsequently correctly recognized with high confidence compared to subsequently nonrecognized items. All peaks significant at p < .001, uncorrected (color bars indicate t statistic). D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f . t / u s e r o n 1 7 M a y 2 0 2 1 for items that became source hits versus source misses. This may reflect a lack of power, given the relatively low trial numbers in specific conditions (i.e., source misses for state), and the absence of a parametric measure like the confidence ratings used for item memory. DISCUSSION We were interested in the potential role of mPFC in con- tributing subjective value to the contents of imagery. Our paradigm provides a way to measure this by manipulating subjective value of imagined items with respect to imag- ined physiological need. The behavioral results suggest that the manipulation was valid, and the imaging results support the hypothesis that mPFC activity reflects the subjective value of elements in imagined scenarios. The manipulation of imagined need succeeded in al- tering the subjective value of elements within imagined scenarios in that participants indicated higher ratings for items congruent with (i.e., likely to satisfy) the state of need. Subsequent recognition memory for items also supports the success of our manipulation. Items that were able to fulfill people’s imaginary needs showed greater subsequent memory, both in being better recog- nized and being better associated to the state of need in which they were presented. This could be because to imagine a congruent item in the imagined scenario is more consistent with our daily life experiences and this enabled participants to have a richer imagination. Simi- larly, congruent items might fit more readily into a preex- isting “schema” allowing for a more rapid integration of the item and imagined state (Tse et al., 2007; Bransford & Johnson, 1972; Bartlett, 1932). Equally, congruent items might have been better remembered because more valu- able scenarios tend to be more strongly represented in memory-related areas (Lebreton, Jorge, Michel, Thirion, & Pessiglione, 2009; Wittmann et al., 2005). The instruction to imagine states of physiological need was accompanied by increased activity in the insula com- pared to neutral states, albeit at an uncorrected threshold. This would be consistent with studies showing insular activation corresponding to interoception of actual physi- ological states (Craig, 2003), including thermo sensa- tion (Craig, Chen, Bandy, & Reiman, 2000) and hunger (Tataranni et al., 1999). One might wonder whether peo- ple are able to imagine themselves in different physiolog- ical states, because physiological states are not usually Table 3. Whole-brain fMRI Analysis Results Region Cluster Size x y z Peak Z Score The Subjective Value of Items in Imagery Right mPFC Anterior cingulate Left ventral mPFC 18 6 8 9 0 57 12 27 −9 −12 45 3 3.98* 3.1 3.63 Imagine States of Need >Imagine Neutral State

Left insula

Right insula

−39 −6 −3

3.27

3.15

Congruent >Incongruent

Caudate

3.6

Subsequent Memory Effect

Left hippocampus

Right amygdala

−21 −12 −18

33 −3 −30

3.33

3.27

All peaks reached an uncorrected significant level of p = .001.

*p value was <.05 at the cluster level with SVC. 1964 Journal of Cognitive Neuroscience Volume 27, Number 10 thought to be under cognitive control. However, involun- tary physiological signs can be influenced by imagination, for example, pupil dilation can be affected by imagining dark or light environments (Laeng & Sulutvedt, 2014). We were interested in the process by which subjective value is afforded to an item within an imagined scenario. To investigate this, we looked for an fMRI signal match- ing the modulation of an item’s subjective value by the imagined state of need, that is, a regressor formed from the subjective rating of the item when imagined as part of a specific scenario minus the subjective rating of that item in daily life. We found activity following this pattern in mPFC, both in a more superior region and the ventral region of mPFC (albeit at an uncorrected threshold for the latter region; see Table 3). This is consistent with our hypothesis for the role of mPFC in imagery. Thus, beyond the representation of the subjective value of choices in decision-making, the mPFC may also play a role in representing the value of items in imagined sce- narios more generally. This more general role might be- gin to explain its involvement in AM retrieval or episodic future thinking, as well as tasks with an implied compo- nent of choice such as planning. Indeed, mPFC activation has been seen together with hippocampal activation during the imagination of rewarding future situations in a decision task (Lebreton et al., 2013). In general, congruent items were rated as more valu- able than incongruent ones. Congruent items might be valuable because of their utility in a specific context (i.e., a congruent state; Hare, Malmaud, & Rangel, 2011) or because congruent items are more self-relevant in a congruent state (D’Argembeau, 2013). Could the re- sults we observed in mPFC be caused by semantic con- gruency effect? To examine the effect of semantic congruency itself, we simply compared the imagination of explicitly congruent or incongruent items, finding ac- tivity in the caudate nucleus (but not in mPFC, where the difference in activity was some way below threshold, at p = .06 uncorrected). Thus, there is little support for a se- mantic interpretation of the mPFC activity we observed. The representation of the combined scenario may involve the striatum, via increased consolidation of the congruent state–item association, consistent with some rodent stud- ies of consolidation (Pennartz et al., 2004). Alternatively, the striatal activation may reflect the involvement of these areas in reward-related processing (e.g., Knutson, Rick, Wimmer, Prelec, & Loewenstein, 2007), in the sense that the imagined interaction with the congruent item seems more rewarding in nature (although we forbade imagined consummation of items). The behavioral results demonstrate a higher recogni- tion rate for congruent items. This memory effect could relate to schema theory: perhaps the encoding of new information (i.e., a congruent item) benefits from being congruent rather than incongruent with the existing sce- nario. The mPFC has been implicated in incorporating new information into existing knowledge structures (van Kesteren et al., 2013; van Kesteren, Ruiter, Fernández, & Henson, 2012; Tse et al., 2011; Benchenane et al., 2010; van Kesteren, Fernández, Norris, & Hermans, 2010). How- ever, mPFC did not show a significant subsequent memory effect. Subsequent memory for items was related to activ- ity in the anterior medial-temporal lobe during encoding, consistent with several previous studies implicating the hippocampus (e.g., Wagner et al., 1998). Our subsequent memory effects also extended into the amygdala. This may be consistent with a role for the amygdala in item memory (Farovik, Place, Miller, & Eichenbaum, 2011; Kensinger, Addis, & Atapattu, 2011; Ranganath, 2010; Kensinger & Schacter, 2006) or with amygdala involvement in enhanc- ing memory for items with affective salience (Hamann, Ely, Grafton, & Kilts, 1999) or intrinsic value as a reinforcer (Rolls, 2005). Unfortunately, we did not have enough sta- tistical power to analyze subsequent memory effects sep- arately in congruent, neutral, and incongruent items to address these possibilities. The recollection of autobiographical information has been associated with a network of brain regions. Al- though many posterior regions have a hypothesized func- tional role within this network (e.g., Schacter et al., 2012; Hassabis & Maguire, 2009; Byrne et al., 2007; Cabeza & St Jacques, 2007), the mPFC has received somewhat less attention. AMs tend to be highly personal and value-laden. For example, we are more likely to remember the experi- ence of having a cup of hot tea after walking outdoors for hours on a cold winter day than having a cup of tea on an ordinary afternoon. Given its association with value in decision-making and with the value afforded by imagined scenarios in this study and related studies (Benoit et al., 2014; Gross et al., 2014; Winecoff et al., 2013; Nieuwenhuis & Takashima, 2011), mPFC activity may reflect the value of recollected information (see also D’Argembeau, 2013). This is perhaps one reason why mPFC is typically not seen in more traditional episodic memory tasks, such as word recognition, where memory for such items may be high, but little value is associated with the retrieved items. Indeed, the subjective value associated with items may be one critical difference between typical autobiographical and episodic memory tasks. To conclude, we have developed a new paradigm for looking at the interaction of imagery and value. We have validated it behaviorally via subjective value ratings and subsequent memory effects. Supporting our hypothesis, we found activity in the mPFC corresponding to the sub- jective value that an item is afforded by the imagined sce- nario. This suggests an extension of the well-known role of mPFC in representing value during decision-making and offers a potential explanation of its involvement in imagery and AM retrieval. Acknowledgments We thank the Medical Research Council U.K., the Wellcome Trust, and the Taiwan Government scholarship for studying Lin et al. 1965 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t . f u s e r o n 1 7 M a y 2 0 2 1 abroad for funding, and the Wellcome Trust Centre for Neuro- imaging at University College London for providing facilities. Reprint requests should be sent to Neil Burgess, Institute of Cognitive Neuroscience, University College London, 17 Queen Square, London WC1N 3AR, United Kingdom, or via e-mail: n.burgess@ucl.ac.uk. REFERENCES Addis, D. R., Moscovitch, M., Crawley, A. 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Lin et al. 1967 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 7 / 1 2 0 7 / 1 1 9 0 5 / 7 1 1 9 9 5 4 7 9 / 5 1 3 8 7 8 o 3 c 3 n 0 _ 6 a / _ j 0 o 0 c 8 n 3 6 _ a p _ d 0 0 b 8 y 3 g 6 u . e p s t d o f n b 0 y 7 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t f / . u s e r o n 1 7 M a y 2 0 2 1 Medial Prefrontal Cortex: Adding Value image

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