Predicting events of varying probability: uncertainty investigated by fMRI

“There is evidence that aspects of outcome probability are coded separately to value in the brain (e.g. Critchley et al., 2001; Volz et al., 2003; Dreher et al., 2006; Tobler et al., 2007).”

“, 2001), and varies as a function of the risk in a decision (Huettel et al., 2005; Paulus et al., 2002; Volz et al., 2003).”

“We also excluded studies that reported contrasts that focused only on risk (i.e. in which models focused on probability rather than magnitude; Volz et al. 2003, 2004; Fukui et al. 2005; Huettel et al. 2006), as well as studies that included a dynamic learning component (i.e. in which anticipation…”

“Common activations of the aesthetic and the symmetry judgment reflected that participants encountered decisions under uncertainty, as indicated by activation of mesial BA 8, anterior insula, and ventral tegmental area (Volz et al., 2003, 2004).”

“Phelps and co-workers (2001) assume that the insula cortex conveys a cortical representation of fear to the amygdala [52], and that uncertainty about the advent of the aversive stimulus during intermittent pairing is reflected by insula and dorsal prefrontal cortex activation [31,32,76].”

“Furthermore, these findings are in accordance with recent studies investigating underdetermined responding (Elliott and Dolan, 1998; Volz et al., 2003).”

“We will conclude with a number of open questions and suggest approaches to address them in future research.”

“…found caudate activity in many tasks ranging in complexity from learning to respond to individual stimuli with arbitrary responses (Toni et al., 2002) to classifying multiple stimuli with varying features into categories (Poldrack et al., 1999, 2001; Seger and Cincotta, 2002; Volz et al., 2003).”

“…1999; Shadlen and Newsome, 2001; Heekeren et al., 2004; Ridderinkhof et al., 2004), with several studies specifically addressing the issue of decision difficulty and uncertainty (Critchley et al., 2001; Paulus et al., 2002; Volz et al., 2003, 2004; Huettel et al., 2005; Grinband et al., 2006).”

“, 2001), ventromedial and ventrolateral frontal cortex (Elliott et al., 2000; Ernst et al., 2002; Rogers et al., 1999; Volz et al., 2003), anterior cingulate (Elliott et al.”

“Furthermore, a number of studies of decision-making have shown insular activity in the absence of reward modulation and/or minimal integration demands (Huettel et al., 2005; Volz et al., 2003, 2004; Critchley et al., 2001; Paulus et al., 2002).”

“This so-called situation model integrates the current language input with both general world knowledge and the prior discourse context (Kintsch, 1998; Zwaan & Radvansky, 1998; Graesser, Millis, & Zwaan, 1997; van Dijk & Kintsch, 1983 ).,In this study we investigated language processing in context, and in particular th…”

“Increased activity in the ventral striatum during a decision-making task involving choices with different probabilities was reported in healthy volunteers (Volz et al., 2003).”

“The insula is ideally placed to integrate the temporal and affective features of reinforcement, given its involvement in reward (Cardinal, 2006; Cardinal et al., 2002; Elliott et al., 2000; Volz et al., 2003; Wittmann et al., 2007), temporal”

“…of this focus (0, 30, 48; for the comparison of negative and positive feedback) were very close to those identified in a study of uncertain versus certain predictions (Volz et al. 2003) (coordinates: 4, 30, 46), as well as a further fMRI study of hypothesis testing (Elliott and Dolan 1998).”

“…studies on the neurobiological bases of choice under uncertainty also reported cuneus activation (Huet- tel et al., 2005; Schlund and Ortu, 2010; Volz et al., 2003), but the activation was not linked to parametric changes in the level of uncertainty or to changes in the learning rate induced…”

“SpeciWc anterior insula activation was reported in decisionmaking under uncertainty (Paulus et al. 2001; Volz et al. 2003), in risk-taking situations (Paulus et al. 2003), in studies on preference judgments (Paulus and Frank 2003), and when receiving monetary rewards (Knutson et al. 2000).”

“…anterior cingulate cortex (ACC), insula, and amygdala are recruited under conditions of uncertainty (Critchley et al. 2001; Davis and Whalen 2001; Volz et al. 2003; Hsu et al. 2005; Paulus 2005; Grinband et al. 2006; Krain et al. 2006; Rosen and Donley 2006; Belova et al. 2007; Dunsmoor et al.…”

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“The greater recruitment of ACC in children, a region involved in conflict monitoring (van Veen and Carter, 2002) and anticipation of uncertain outcome (Volz et al., 2003) suggests that children are less efficient than adolescents or adults at processing uncertainty.”

“…with this finding, studies examining probability in the context of " guess " , " certainty " , or " predictability " have also found increased dACC/dorsomedial prefrontal cortical activity in the lower probability conditions (Elliott et al., 1999; Koechlin et al., 2000; Volz et al., 2003, 2005).”

“First, activity in this region is known to be modulated by ambiguity both in social ( Jenkins & Mitchell, 2010) and nonsocial (Volz, Schubotz, & von Cramon, 2003) contexts.”

“In their study, Volz et al. (2003) showed participants pairs of pictures showing comic figures which were systematically associated with a particular reward probability (either 0.6, 0.7, 0.8, 0.9, or 1.0), and these associations were consistent throughout the experiment.”

“, 2004), prediction tasks (Paulus et al., 2002; Volz et al., 2003), working memory tasks (Jansma et al.”

“In addition to OFC, several other regions, including anterior cingulate cortex and midbrain regions (in particular, the nucleus accumbens and ventral striatum), are reported as being important for uncertain decision-making (Paulus, Hozack, Frank, & Braun, 2002; Rodriguez, Aron, & Poldrach, 2005;…”

“An important future direction for this area of research is to include multiple uncertain cues with varying associated probabilities of negative outcomes (Bach et al., 2009; Volz et al., 2003).”

“The percent area under the second through fourth images of the hemodynamic response curve (AUC), which occur prior to UCS presentation, was used as an index of the response magnitude produced by each CS. Functional maps reflecting the AUC were converted to a standard stereotaxic coordinate system ( Talairach & Tournou…”

“The RCZ is involved in unsuccessful humor processing or the processing of pictures containing an irresolvable incongruity: This activation might reflect conflict monitoring as described in several studies (see Botvinick, Cohen, & Carter, 2004 , for a review) or increasing uncertainty (Volz, Schubotz, & von Cramon, 200…”

“Neuroimaging evidence indicates that the DLPFC (Paulus et al., 2002; Huettel et al., 2005), posterior parietal cortex (Volz et al., 2003; Huettel et al., 2005), anterior cingulate cortex (ACC; Elliott and Dolan, 1998; Critchley et al., 2001; Stern et al., 2010), orbito-frontal cortex (OFC; Goel and…”

“These studies investigated hypothesis testing with low restrictions (Elliott and Dolan, 1998), the application of arbitrary SR-rules (Goel and Dolan, 2000), and the detection of arbitrary SR-rules (Knutson et al., 2003; Volz et al., 2003).”

“…relationship between choice uncertainty and activity in the medial prefrontal cortex when subjects learn through trials and errors the probability of making a correct choice (Volz et al., 2003; Huettel et al., 2005; Volz et al., 2005; Grinband et al., 2006; Huettel, 2006; Platt and Huettel, 2008).”

“Other studies have also reported frontal or parietal activations related to prediction errors [O’Doherty et al., 2003; Volz et al., 2003].”

“Recent investigations have also attributed a critical role of the anterior cingulate and medial prefrontal cortex in decision-making situations (Bush et al., 2002; Rogers et al., 2004; Volz et al., 2003).”

“2B) (Critchley et al., 2001; d'Acremont and Bossaerts, 2008; Grinband et al., 2006; Hsu et al., 2005; Huettel et al., 2005; Paulus et al., 2003; Preuschoff et al., 2006; Rolls et al., 2008; Rushworth and Behrens, 2008; Volz et al., 2003, 2004; Weber and Huettel, 2008).”

“Studies in healthy individuals have related greater activity in posterior MFC (pMFC)/dorsal anterior cingulate cortex (Brodmann’s areas [BAs] 6, 8, 24, 32) [Grinband et al., 2006; Krain et al., 2006; Volz et al., 2003] and aI/fO (BAs 13, 47) [Preuschoff et al.”

“…in the midbrain (Fiorillo et al., 2003, 2005), the ventral striatum (Knutson et al., 2005; Preuschoff et al., 2006), the ACC (Amiez et al., 2006), and the medial (Knutson et al., 2005; Matsumoto et al., 2003; McClure et al., 2004; Volz et al., 2003, 2004) and lateral PFC (Matsumoto et al., 2003).”

“…outcome and perceptual difficulty have typically shown increased activity within the dorsal part of the medial prefrontal cortex (DMPFC; Volz et al., 2003, 2004; Hsu et al., 2005; Knutson et al., 2005; Grinband et al., 2006; Krain et al., 2006; Callan et al., 2009; Mohr et al., 2010a),…”

“It is likely that this area generates predictions about the upcoming stimulus, which are then tested in the remaining areas, an interpretation that concurs with previous studies showing that the dmPFC is involved in generating predictions under uncertainty (Volz et al. 2003).”

“Although we did find activity in OFC/IFC, we did not replicate previous findings linking decision uncertainty to nearby anterior insula (Volz et al., 2003; Grinband et al., 2006; Huettel, 2006; Preuschoff et al., 2008), which may be attributable to the fact that greater decision uncertainty in…”

“Such an assumption is highly compatible with the results of studies investigating the functional neuroanatomy of decision-making in the presence of uncertainty (e.g. Volz et al., 2003 ).”

“A number of brain regions have been implicated in processing uncertainty, including the anterior cingulate, anterior insula, amygdala, posterior parietal cortex, and regions within the medial, orbitofrontal and dorsolateral prefrontal cortices (Bechara et al., 1999; Critchley et al., 2001; Elliott et al., 1999; Ernst…”

“, the pre-SMA, the anterior cingulate and the medial frontal gyrus, are involved in both analysis and executive control of movements, especially in probabilistic contexts (Volz et al., 2003; Ridderinkhof et al., 2004).”

“, 2002; Schubotz and von Cramon, 2002) and response uncertainty (Volz et al., 2003).”

“The activation in the pMFC corresponds well with findings from Volz et al. (2003, 2004) who showed that this region reflects the degree of uncertainty and is related to decision conflicts, which in the present study increase with increasing similarity.”

“This requirement for rapid updating was absent in the present study—and in prior studies of uncertainty (Volz et al., 2003, 2004)—suggesting that the specific manner in which uncertainty is generated determines what brain systems are necessary for its resolution.”

“Previous studies showing the involvement of this area in processing events under uncertainty and error evaluation (Volz, Schubotz, & von Cramon, 2003, 2006) provide support for this interpretation.”

“As can be seen in Figure 3, the DMFC is preferentially activated when subjects predict events under varying levels of uncertainty based on natural sampling (Volz et al., 2003).”

“Third, as mentioned before, the dmPFC is very important for the computation and estimation of uncertainty of potential threat (Michael et al., 2015; Volz et al., 2003).”

“The dmPFC and dACC are also crucial brain regions for several important subfunctions that are necessary (but not sufficient) for task selection in dynamic environments, such as action outcome prediction (Alexander and Brown 2011), the processing of uncertainty (Volz et al. 2003; Rushworth and Behrens 2008) and task di…”

“, 2005) and that risk correlates with the BOLD signal in the anterior insula (aINS) (Critchley et al., 2001; Huettel et al., 2005; Paulus et al., 2003; Volz et al., 2003).”

“Other studies have suggested that the pre-SMA regions (e.g., BA 8 and 6) are involved in " decisions under uncertainty " (Volz et al., 2003, 2004), free selection of responses (Lau et al., 2004), and response conflict monitoring (Garavan et al., 2003).”

“Of those, the anterior dorsal insula, ACC and mesial BA 8 have been implicated in situations characterized by uncertainty (Volz et al., 2003; Wager and Feldman, 2004; Volz, 2005), signifying the likelihood of errors and a need to adapt one’s expectations.”

“Indeed, previous studies showed that cerebellar involvement in cognitive tasks increases with the complexity and uncertainty of the task (Boecker et al., 2002; D’Agata et al., 2011; Schubotz and von Cramon, 2002; Volz et al., 2003).”

“…has been investigated in several psychological, neuroimaging and neuropsychological studies (Elliott et al., 1999; Hsu et al., 2005; Krain et al., 2006; Paulus et al., 2001, 2002; Sanfey et al., 2006; Volz et al., 2003, 2004; Yoshida and Ishii, 2006) (see Platt and Huettel (2008) for a review).”

“Activation in the parahippocampal gyrus has been linked to learning effects while processing rewards of uncertain probability (Elliott et al. 2000; Volz et al. 2003) and persistent activation in this region has been associated with higher error rate (Paulus et al. 2002).”

“This work indicates that the PFC, insula, and amygdala respond in anticipation of aversive stimuli and show increased activity when participants have uncertain expectations (Critchley et al., 2001; Davis and Whalen, 2001; Dunsmoor et al., 2007; Volz et al., 2003).”

“…Sarinopoulos et al. 2010 36 20.7 (1.5) 3 High vs. low uncertainty Schlosser et al. 2009 12 26.8 (2.1) 3 Linear relationship with uncertainty Volz et al. 2003 16 24.9 3 Linear relationship with uncertainty Volz et al. 2004 12 25.1 3 Linear relationship with uncertainty (B) Type-B task…”

“ously that prefrontal cortical areas are engaged during processing of uncertain information requiring higher levels of cognitive control (Koch et al., 2008; Krain et al., 2006; Satterthwaite et al., 2007; Volz et al., 2003).”

“For example, Volz et al. (2003, 2004) reported this region when participants where highly uncertain about their decisions compared to situations where participants could recollect their decisions from memory.”

“Neuroimaging studies have demonstrated that the probability with which a stimulus or response occurs modulates activity in the anterior insula, anterior and posterior cingulate cortex, middle frontal gyrus, ventromedial and dorsolateral prefrontal cortex [20,37,46].”

“As certainty decreases, activity increases in the lateral orbitofrontal cortex (LOFC), dorsal anterior cingulate cortex (DACC), dorsomedial prefrontal cortex (DMPFC), LPFC, anterior insula, and ventral striatum (Critchley et al., 2001; Huettel et al., 2005; Paulus et al., 2001; Preuschoff et al., 2006; van Leijenhorst…”

“The functional correlation analysis aimed to depict the activation network coupled with mPFC; a cortical region known to be sensitive to internally generated processing ( Volz, Schubotz, & von Cramon, 2004; Volz et al., 2003 ).”