Decoding the Content of Delayed Intentions

“For each series, the variance in the BOLD signal was decomposed with a set of regressors in a general linear model (Friston et al., 1995), using a mixed blocked and event-related design (Visscher et al., 2003).”

“According to some authors (Gollwitzer, 1999), intentions can often be conceptualized as mental links between an anticipated future cue (e.”

“The mask described above, indicating regions within RLPFC showing significant activation for the PM store baseline store contrast, was used as the seed region for a psychophysiological interaction (PPI) analysis (Friston et al., 1997).”

“Given the well known role of the hippocampus in memory encoding (Eichenbaum and Cohen, 2001; Simons and Spiers, 2004; Squire et al., 2004), and suggestions that the medial temporal lobes play a role in representing PM intentions (Adda et al.”

“By contrast, the more posterior region identified by Haynes et al. (2007) may be involved in higher-level representation of cognitive operations (i.e., task sets) (Monsell, 2003) rather than specific perceptual or motor representations.”

“A searchlight approach was used (Kriegeskorte et al., 2006), investigating decoding accuracy from a sphere of voxels centered on each voxel in the brain in turn (radius, three voxels).”

“Rostrolateral prefrontal cortex (RLPFC), corresponding to the lateral aspects of Brodmann area (BA) 10, is widely considered to occupy the top of a hierarchy of lateral prefrontal regions supporting behavioral regulation (Badre and D’Esposito, 2009; Christoff et al., 2009; Koechlin et al., 2003).”

“A similar region was also identified by Soon et al. (2008), from which it was possible to decode which of two motor responses participants were preparing to make.”

“These findings are consistent with other evidence showing that cortical regions specialized for processing particular types of low-level perceptual and motor information can also play content-specific mnemonic functions (Chieffi et al., 1999; Eschen et al., 2007; Harrison and Tong, 2009).”

“This is sometimes termed “prospective memory” (PM) (Brandimonte et al., 1996; McDaniel and Einstein, 2000; Kliegel et al., 2008; West, 2008).”

“Neuroimaging and neuropsychological evidence implicates RLPFC in our ability to hold an overarching goal in mind while engaged in subordinate tasks: “cognitive branching” (Koechlin et al., 1999) or “goal– subgoal integration” (Braver and Bongiolatti, 2002).”

“Given the well known role of the hippocampus in memory encoding (Eichenbaum and Cohen, 2001; Simons and Spiers, 2004; Squire et al., 2004), and suggestions that the medial temporal lobes play a role in representing PM intentions (Adda et al., 2008), it is notable that bilateral hippocampal…”

“Given the well known role of the hippocampus in memory encoding (Eichenbaum and Cohen, 2001; Simons and Spiers, 2004; Squire et al., 2004), and suggestions that the medial temporal lobes play a role in representing PM intentions (Adda et al.”

“Rostrolateral prefrontal cortex (RLPFC), corresponding to the lateral aspects of Brodmann area (BA) 10, is widely considered to occupy the top of a hierarchy of lateral prefrontal regions supporting behavioral regulation (Badre and D’Esposito, 2009; Christoff et al., 2009; Koechlin et al., 2003).”

“This is sometimes termed “prospective memory” (PM) (Brandimonte et al., 1996; McDaniel and Einstein, 2000; Kliegel et al., 2008; West, 2008).”

“DOI:10.1523/JNEUROSCI.5336-10.2011 Copyright © 2011 the authors 0270-6474/11/312888-07$15.00/0 ration of the maintenance period, it was possible to disambiguate these three phases (cf. Rowe et al., 2000) (Fig.”

“RTs on PM storage trials were significantly slower than baseline storage trials (PM, 878 ms; baseline, 842 ms; F(1,30) 29.2, p 0.0001), indicating a RT cost associated with maintaining PM intentions and/or monitoring ongoing trials for PM cues (Smith, 2003; Einstein et al., 2005).”

“In this way, rather than intentions being represented in just one brain area, multiple distinct regions are proposed to represent different dimensions of our future intentions: perceptual triggers for action, specific responses, and higher-level task sets or representations of action schemas (Wood and Grafman, 2003).”

“For each phase, contrast images representing conditions of interest were entered into a repeated-measures ANOVA using nonsphericity correction (Friston et al., 2002).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“It has previously been reported that representations of future intentions (which of two cognitive operations: addition vs subtraction) can be decoded from medial prefrontal cortex (Haynes et al., 2007).”

“The region of medial prefrontal cortex identified by Haynes et al. (2007) was considerably posterior to the region identified here, in a part of the brain known to exhibit considerable fine-grained functional specialization (Gilbert et al., 2006, 2010).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“RTs on PM storage trials were significantly slower than baseline storage trials (PM, 878 ms; baseline, 842 ms; F(1,30) 29.2, p 0.0001), indicating a RT cost associated with maintaining PM intentions and/or monitoring ongoing trials for PM cues (Smith, 2003; Einstein et al., 2005).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“, 1999) or “goal– subgoal integration” (Braver and Bongiolatti, 2002).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“According to this latter model, RLPFC might play a non- content-specific role via its interactions with distinct brain regions that themselves represent the content of intentions (Sakai and Passingham, 2003; Rowe et al., 2007).”

“The idea that RLPFC plays a “content-free” role in PM is congruent with repeated findings of RLPFC activation in PM paradigms using a wide variety of different stimulus types and task demands (Okuda et al., 1998; Burgess et al., 2001, 2003; Simons et al., 2006; Gilbert et al., 2009).”

“For each series, the variance in the BOLD signal was decomposed with a set of regressors in a general linear model (Friston et al., 1995), using a mixed blocked and event-related design (Visscher et al., 2003).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and…”

“Rostrolateral prefrontal cortex (RLPFC), corresponding to the lateral aspects of Brodmann area (BA) 10, is widely considered to occupy the top of a hierarchy of lateral prefrontal regions supporting behavioral regulation (Badre and D’Esposito, 2009; Christoff et al., 2009; Koechlin et al., 2003).”

“The idea that RLPFC plays a “content-free” role in PM is congruent with repeated findings of RLPFC activation in PM paradigms using a wide variety of different stimulus types and task demands (Okuda et al., 1998; Burgess et al., 2001, 2003; Simons et al., 2006; Gilbert et al., 2009).”

“This suggests that the two-back task was sufficiently distracting to prevent continuous rehearsal of PM intentions (Einstein et al., 2003).”

“In addition, some patients derive considerable benefit from nonspecific cues (e.g., an auditory beep) that remind them of the existence of a delayed intention but not its content (Fish et al., 2007).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“According to this latter model, RLPFC might play a non- content-specific role via its interactions with distinct brain regions that themselves represent the content of intentions (Sakai and Passingham, 2003; Rowe et al., 2007).”

“, 2004), and suggestions that the medial temporal lobes play a role in representing PM intentions (Adda et al., 2008), it is notable that bilateral hippocampal activation was observed for this contrast (Fig.”

“The region of medial prefrontal cortex identified by Haynes et al. (2007) was considerably posterior to the region identified here, in a part of the brain known to exhibit considerable fine-grained functional specialization (Gilbert et al., 2006, 2010).”

“These findings are consistent with other evidence showing that cortical regions specialized for processing particular types of low-level perceptual and motor information can also play content-specific mnemonic functions (Chieffi et al., 1999; Eschen et al., 2007; Harrison and Tong, 2009).”

“These findings are consistent with other evidence showing that cortical regions specialized for processing particular types of low-level perceptual and motor information can also play content-specific mnemonic functions (Chieffi et al., 1999; Eschen et al., 2007; Harrison and Tong, 2009).”

“…stored PM intentions, consistent with previous neuroimaging and neuropsychological studies indicating the involvement of this region in PM (Okuda et al., 1998; Burgess et al., 2000, 2001, 2003; Simons et al., 2006; West, 2008; Gilbert et al., 2009; Reynolds et al., 2009; Uretzky and Gilboa, 2010).”

“, 2007 showed that it is possible to decode from activity n medial and lateral prefrontal cortex which of two tasks particiants had freely chosen to do); (c) the form of the instruction given: ilbert et al. (2009) found rostral PFC activation differences during PM task according to whether the task instruction was eit…”

“Other models, such as the Gaussian Naïve Bayes approach [14], rely on differnces in activity patterns without capturing any of the interactions among brain regions that are likely critical for differentiating affective states [20,22,23].”

“Participation took approximately 20–30 minutes, for which participants received $2.”

“…prospective memory have suggested that univariate activity in the anterior prefrontal cortex is correlated with event-based (Burgess et al., 2007; Gilbert, 2011; Gilbert et al., 2005, 2006; Okuda et al., 2007; Simons et al., 2006) as well as time-based prospective memory (McFarland and Glisky,…”

“Similarly, in Gilbert (2011), the overall procedure was specifically created such that “correct responses on PM retrieve trials had to be self-initiated on the occurrence of the appropriate item, rather than being strongly cued by the stimulus Frontiers in Human Neuroscience | www.frontiersin.org 6…”

“, 2001)​, and prospective memory ​(Gilbert, 2011; Haynes & Rees, 2006; Momennejad & Haynes, 2012, 2013)​.”

“A few recent brain imaging studies have investigated delayed intentions (Gilbert et al. 2009; Gilbert 2011; Momennejad and Haynes 2012) and indicate that they are processed in the most anterior part of the prefrontal cortex, namely the frontopolar cortex. this finding is consistent with research on…”

“Sachdeva and Gilbert (2020) provided evidence that one such factor is a preference to avoid cognitive effort.”

“Furthermore, a re- cent study by Gilbert (2011) found that although lateral rPFC (lrPFC) exhibited robust activation during maintenance of delayed intentions, the content of those intentions (i.e., the nature of the target stimuli and appropriate PM responses) could not be decoded from this region.”

“One was an analysis that discriminated a dual from single task (Gilbert, 2011), which was excluded from the strict categorization because of the obvious confound with difficulty (for discussion, see Woolgar, Golland, & Bode, 2014; Todd, Nystrom, & Cohen, 2013), and the other pertained to discrimination of task set whe…”

“The authors proposed that, while mPFC plays a role in representing the content of 5 delayed intentions, lateral PFC has instead a content-fr e role in PM tasks (Gilbert, 2011), likely 6 mediating a general predisposition to act.”

“It can also apply to higher-order brain regions, such as prefrontal cortex, and higher-level representations, such as the content of participants’ delayed intentions (Gilbert, 2011; Gilbert, Armbruster, & Panagiotidi, 2012; Haynes et al. 2007).”

“Despite the strong impact that the multiprocess framework (McDaniel & Einstein, 2007a) has had in PM research and despite findings suggesting RLPFC involvement in strategic monitoring during PM (Benoit et al., 2011; Burgess et al., 2001, 2011; Gilbert, 2011; Reynolds et al., 2009), there has been little investigation…”

“The rostrolateral prefrontal cortex is involved in delayed, context-specific implementation of acquired rules [Gilbert, 2011; Koechlin et al., 1999; Sakai and Passingham, 2003].”

“, 2007), in PM paradigms where delays were occupied with an ongoing task (Gilbert, 2011; Momennejad and Haynes, 2012), and even during unconscious preparation of task-related choices (Soon et al.”

“According to Gilbert et al. [Gilbert, 2011; Gilbert et al., 2012], activation of the lateral RPFC is specific to the storage stage of PM but does not directly support the con- tent of delayed intentions.”

“Such representations are encoded in local spatial activation patterns in the lateral prefrontal, dorsal anterior cingulate, and posterior parietal cortex (Bode and Haynes, 2009; Gilbert, 2011; Woolgar et al., 2011; Wisniewski et al., 2015).”

“One such study (Gilbert, 2011) examined intention formation, maintenance, and retrieval of prospective memory.”

“A growing number of studies which investigated intention representations usingMVPA indicate that the anterior medial PFC (Gilbert, 2011), lateral PFC, and parietal cortex (Bode and Haynes, 2009; Momennejad and Haynes, 2012, 2013; Waskom et al.”

“Exploiting its information-mapping nature, a number of studies have applied this approach to a vast range of research topics([15,16,22,25,26,30,58-64]).”

“In the earlier study by Gilbert (2011), also analyzing the present data set, medial rostral pFC was found to maintain a representation of the forthcoming cue for action during store periods.”

“Finally human lesion studies [19] and functional neuroimaging with the suggestion that representations of the PM may be stored in occipital sites [23].”

“This frontopolar region was more active during production of highly coherent speech, which is in line with the idea that RLPFC is involved in the planning and sequencing of complex, goal-directed behaviours (Desrochers, Chatham, & Badre, 2015; Fuster, 2004; Gilbert, 2011; Koechlin & Summerfield, 2007; Momennejad & Hay…”

“Greater involvement of PFC in monitoring processes and prospective memory has been proposed (Floresco et al., 1997; Fuster, 2001; Gilbert, 2011), and thus extending the temporal requirements for information maintenance may require greater strength of connections from this region (Buchsbaum et al.”

“Previous studies have successfully employed multivariate analyses to decode the content of specific mental states and have pointed towards an increased sensitivity of this approach (Kamitani and Tong 2005; Haynes and Rees 2006; Haynes et al. 2007; Bode and Haynes 2009; Gilbert 2011).”

“…in neuroimaging paradigms investigating participants' ability to fulfill delayed intentions (Burgess et al., 2001, 2003; Gilbert et al., 2009; Gilbert, 2011; McDaniel et al., 2013; Momennejad & Haynes, 2012, 2013; Okuda et al., 1998; Reynolds et al., 2009; Simons et al., 2006; reviewed by…”

“…Passingham, 2002, 2006], storing situationally contingent intentions in the service of prospective memory [Beck, et al., 2014; Burgess, et al., 2011; Gilbert, 2011; Momennejad and Haynes, 2012], making counterfactual inferences that facilitate hypothesistesting [Donoso, et al., 2014], and…”

“The finding in Gilbert [42] that functional coupling between RLPFC and intention-representing brain regions was increased during intention storage would be trivially simple to simulate in the present modelling framework.”

“This hypothesis is inconsistent with the idea that involvement of BA 10 in cognitive tasks is substantially content-independent [3,16].”

“Several prospective memory studies assessing the correlates of future intentions have shown the involvement of brain areas recruited by our task, mainly the ventral RPFC (Gilbert, 2011; Landsiedel and Gilbert, 2015; Momennejad and Haynes, 2013, 2012).”

“, waiting for a word versus a picture to reappear) can be decoded from posterior cortical regions (Gilbert, 2011).”

“Glucocorticoids bind to glucocorticoid receptors that are widespread in the PFC [12,13] and alter PFC activity [14,15].”

“Medial and lateral rostral prefrontal cortex are involved in maintaining specific intentions over delays (Burgess et al., 2003; Gilbert, 2011; Haynes et al., 2007) and may use such information to guide future responses (Karnath et al., 1991; Lepage and Richer, 2000).”

“Content memory is traditionally linked with the temporal lobes of the brain (Gordon, Rissman, Kiani, & Wagner, 2014; Squire & Zola-Morgan, 1991; Wixted & Squire, 2011), whereas the frontal lobes have been shown to play a key role in temporal order and prospective memory (DeVito & Eichenbaum, 2011; Gilbert, 2011; Ruben…”

“…Quayle, & Frith, 2001; Burgess et al., 2003; den Ouden, Frith, Frith, & Blakemore, 2005; Gilbert, Gollwitzer, Cohen, Burgess, & Oettingen, 2009; Gilbert, 2011; Hashimoto, Umeda, & Kojima, 2011; Haynes et al., 2007; Okuda et al., 1998, 2007; Poppenk, Moscovitch, McIntosh, Ozcelik, & Craik,…”

“More specifically, activations in lateral rPFC are most often associated with maintaining a delayed intention (den Ouden et al., 2005; Gilbert, 2011; Gilbert et al., 2009; Okuda et al., 2011; Reynolds et al., 2009; Simons et al., 2006), a finding which is in agreement with other suggested theories regarding the role o…”

“…medial PFC responses, mostly in rostral prefrontal areas (Burgess et al., 2001; den Ouden et al., 2005; Simons et al., 2006; Poppenk et al., 2010; Gilbert, 2011; Benoit et al., 2012), whereas non-delayed intention experiments showed activity in the dorsal medial and lateral prefrontal regions…”

“…in addition to lateral BA10, activity is also higher in the bilateral medial frontal gyrus (BA 8/32), the left precuneus and left parietal cortex (BA7) (Benoit, Gilbert, Frith, & Burgess, 2012), as well as a region in BA46 extending to the insular cortex and the anterior cingulate (Gilbert, 2011).”

“…event-and time-based PM tasks (Okuda et al., 2007), and measures of different stages of PM processing such as encoding, storage, cue identification and intention retrieval (Gilbert , 2011; Poppenk, Moscovitch, McIntosh, Ozcelik, & Craik, 2010; Simons, Scholvinck, Gilbert, Frith, & Burgess, 2006).”

“…in brain activity between having any intention versus no intention, these studies have attempted to identify differences in brain activity between two specific intentions (Gilbert, 2011; Haynes et al., 2007; Momennejad and Haynes, 2012, 2013; Soon et al., 2013, 2008; Wisniewski et al., 2016).”

“It enables us to decode fMRI data by focussing the analysis around a single voxel at a time, while combining the signals within a certain radius from the centred voxel to compute a multivariate effect statistic at every location (Haynes & Rees, 2006; Alink, Euler, Kriegeskorte, Singer, & Kohler, 2012; Corradi-Dell’Acq…”

“…adopt the designation “delayed intention” for intentions about specific actions that are formed, then stored in memory, and later realized (following Gilbert, 2011). binding is defined as a shift in perceived time of an action toward the effect of the action, and a complementary shift in…”

“Contrary to previous studies that have indicated a key role for the anterior prefrontal cortex (aPFC) in representing prospective intentions (Gilbert, 2011; Momennejad and Haynes, 2012, 2013), we found that regions known to support perception and working memory for the PM intentions used here…”

“Most research has focused on strategic monitoring and maintenance of prospective memory cues and have implicated the rostral prefrontal cortex (rPFC; BA10) as an important region for that process (Burgess et al., 2003, 2011; Gilbert et al., 2006; Simons et al., 2006; Haynes et al., 2007; Okuda et al., 2007; Soon et al…”

“…important problem for the models of prospective memory closely tied to the PM paradigm is the finding from fMRI experiments that relevant areas of the prefrontal cortex often are activated during the maintenance phase of the PM paradigm (Cona et al., 2016; Gilbert, 2011; Momennejad & Haynes, 2013).”

“It appears that one can obtain surprising dissociations between the regions activated and those showing content specific effects using voxel pattern classification procedures, as in the study of Gilbert (2011) on the role of frontopolar cortex on the encoding, storage and retrieval of intentions.”

“As in our study, the same network did not necessarily encode the outcome of the decision (Gilbert, 2011).”

“Specifically, prospective memory tasks require maintaining intentions while simultaneously performing unrelated secondary tasks during a delay period (Burgess, Scott, and Frith 2003; Gilbert 2011), an absent feature in our task.”

“These identified regions have been previously associatedwith cognitive processing such as retrieval, maintenance, the process of rules, or demands during task switching (Reverberi, Görgen, & Haynes, 2012; Gilbert, 2011; Woolgar et al., 2011; Bunge, Kahn, Wallis, Miller, & Wagner, 2003; Sakai & Passingham, 2003).”

“Nonetheless, while Gilbert (2011) decoded simpler stimulus-response mappings, in our study we classified abstract task-set information, as previous studies that also decoded intention from lPFC did (e.g. Haynes et al., 2007; Momennejad & Haynes, 2013).”

“Although these findings are seemingly contrary to the content-free role of the RLPFC proposed by Gilbert (2011), Wendelken et al. (2011) acknowledge a large amount of overlap between the dorsal and ventral regions. Thus, Gilbert’s (2011) spotlight implementation of MVPA may not have been sensitive to content-specific…”

“lobules bilaterally (BA 7 and BA 40), a region that has been implicated in encoding and maintenance/retrieval of PM intentions (Gilbert, 2011; Poppenk, Moscovitch, McIntosh, Ozcelik, & Craik, 2010; Reynolds et al., 2009).”

“The study by Gilbert (2011) is of particular interest for the investigation of motivational modulations, as it is one of the rare studies using an event-related functional Magnetic Resonance Imaging (fMRI) paradigm to separate different phases of processing delayed intentions: (1) encoding of the…”

“These identified regions have been previously associated with 14 cognitive processing such as retrieval, maintenance, the process of rules or demands during task switching 15 (Bunge et al., 2003; Sakai and Passingham, 2003; Gilbert, 2011; Woolgar et al., 2011; Reverberi et al., 16 2012).”

“Cognitive processes supporting intention-related event recognition are associated with activity in the rostrolateral prefrontal cortex (rlPFC), lateral parietal and temporal cortices, and medial occipital cortex (Burgess, Quayle, & Frith, 2001; Gilbert, 2011; Reynolds, West, & Braver, 2009).”

“We adapted a cognitive offloading task from Sachdeva and Gilbert 11 , measuring aspects of prospective memory in a way that requires remembering and acting delayed intentions (Fig.”

“The rostrolateral prefrontal cortex interacts with the putamen to maintain representations of future intentions [19], whereas the connectivity between the anterior prefrontal cortex and the precuneus has been reported to support the top-down sustained monitoring process [20].”

“Later researchers called such preserved memories of future planned activities “a prospective memory” (Burgess, Gonen-Yaacovi & Volle, 2011; Gilbert, 2011; Momennejad & Haynes, 2012).”

“(p. 127) Later investigators (e.g., Burgess, Gonen-Yaacovi, & Volle, 2011; Gilbert, 2011; Momennejad & Haynes, 2012) have called such stored memories of planned future actions prospective memory.”

“This does not exclude that information about intentions might also be stored at different levels, e.g. in the pattern of inter-regional correlations of BOLD time-courses (Gilbert, 2011), which can be assessed using graph theory (Cole et al., 2013; Dosenbach et al., 2007; Kouneiher et al., 2009).”

“This formulation can be solved by the following optimization problem [5]: L, S = argmin !,! X − L − S ! , subject to rank L ≤ r, S ! ≤ k × n × m ,”

“68 activity in situations that require maintenance of information across periods of unrelated tasks (Koechlin et al., 1999; Pisapia et al., 2007; Reynolds et al., 2009; Gilbert, 2011).”