Evidence Accumulation and the Moment of Recognition: Dissociating Perceptual Recognition Processes Using fMRI

“As a means of accommodating more diverse patterns of data, UPGMA was developed (Eisen et al., 1998).”

“, 2003), attention for action (Posner et al., 1988; Posner and Petersen, 1990; Bush et al., 1998), response conflict/competition (Bush et al.”

“, 1998), response conflict/competition (Bush et al., 1998; Botvinick et al., 1999; Botvinick et al., 2001), confidence (Fleck et al.”

“, geons) provide a higher degree of object level information than straight lines (Biederman, 1987).”

“…proven highly effective in describing human performance in recognition memory, lexical decisions, economic decisions, and sensory discriminations (Ratcliff, 1978; Busemeyer, 1985; Bundesen, 1990; Nosofsky and Palmeri, 1997; Ratcliff and Rouder, 1998; Logan and Gordon, 2001; Ratcliff et al.,…”

“Consistent with our data, several other fMRI studies have also associated the aPFC, dACC/ msFC and aI/fO with important task control functions [24,27,33,36,40].”

“Similarly, there is extensive overlap between the VA-related network and regions activated by the sudden recognition of a slowly revealed target (Ploran et al., 2007).”

“…to increase with the degree of difficulty or uncertainty in a decision (Critchley et al. 2001; Grinband et al. 2006; Thielscher and Pessoa 2007), and Ploran et al. (2007) found a spike of activity in this network time-locked to the ‘‘moment of recognition’’ in a perceptual discrimination task.”

“In another study (Ploran et al., 2007), as a subject viewed pictures that were revealed from a blank screen gradually in eight steps (2 s each), the areas that exhibited a gradual buildup in activity peaking in correspondence with the time of recognition were the parietal and frontal areas as well…”

“ness of the percept (Ploran et al. 2007).”

“If in the same experiment, we could demonstrate regions that show clear error-like signals without the recognition-like profile of the Ploran et al. (2007) experiment, these regions could be more unequivocally ascribed to feedback processing rather than recognition per se.”

“More recently Ploran et al. (2007) found in an experiment employing a behavioral paradigm in which objects gradually emerge from noise that the activity of anterior insula was strongly linked to the moment when the subjects became aware of the identity of the object (allowing for the delays…”

“For example, in ‘‘extended reveal’’ paradigms that progressively reveal a stimulus out of visual noise, the anterior insula responds strongly at the moment of recognition when the participant perceives the stimulus as a salient object [Ploran et al., 2007; Wheeler et al., 2008].”

“Likewise, recent data demonstrated that ventral and anterior regions of the insula representing relevant ‘‘interoceptive sites’’ also support processes related to focal attention [76–77].”

“…used a novel analytic technique to demonstrate that errorrelated activity in dACC during a stop signal task followed inhibitory failures, in accord with characterization of the region as being involved in postresponse monitoring, evaluation, and adjustment signaling (see also, Ploran et al. 2007).”

“2005), as well as with evidence suggesting that regions of IPS accumulate evidence in service of perceptual recognition decisions (Ploran et al. 2007).”

“…evidence during vibrotactile decision making (Pleger et al., 2006), increase at the moment of a perceptual decision in an image recognition task (Ploran et al., 2007), and correlate with a non-monotonic RT function during an auditory discrimination task, implying a role in the decision process…”

“Thus, the degree to which relevant VOTC structures are engaged corresponds to the strength of visual evidence, whether that evidence comes from external inputs that drive current perception (15, 19) or episodic remembering.”

“Similarly, results of human neuroimaging studies suggest that the integration of sensory evidence might involve regions of the parietal and prefrontal cortex [8, 9, 13, 22].”

“001), we expected to find an earlier and stronger increase of activity (Heekeren et al., 2004; Ploran et al., 2007; Ho et al., 2009).”

“One could argue, however, that an earlier activation is not necessarily related to the perceptual decision process itself, but could just as well reflect the perceived stimulus salience at the moment of recognition, in line with the above-mentioned study by Ploran et al. (2007). Together, findings from perceptual deci…”

“Numerous studies report activity in this region for nonword and nonletter stimuli (Price and Devlin 2003; Xue et al. 2006; Ben-Shachar et al. 2007; Ploran et al. 2007; Starrfelt and Gerlach 2007; Xue and Poldrack 2007; Kherif et al. 2011; Mei et al. 2010; Van Doren et al. 2010).”

“Insular activations unique to Trained subjects are in an anterior region associated with interoception, subjective feeling and perceptual awareness (Kikyo et al., 2002; Ploran et al., 2007; Craig, 2009).”

“The first is that these areas serve as accumulators of evidence for perceptual decision making (Shadlen and Newsome, 2001; Ploran et al., 2007).”

“To dissociate accumulation from sustained activity, a recent study has artificially prolonged the period of evidence accumulation by revealing objects gradually in a stepwise fashion (Ploran et al., 2007).”

“In controls high gamma-band activity in the face condition was found in a distributed network, such as the temporal cortex and IFG, which have been related to face perception and perceptual decision making (Kanwisher et al., 1997; Ploran et al., 2007).”

“…variance N97%), areaswhich have previously been implicated in perceptual decision making and evidence accumulation, both in the human (Heekeren et al., 2006; Ploran et al., 2007; Tosoni et al., 2008) and primate (Kiani and Shadlen, 2009; Kim and Shadlen, 1999; Shadlen and Newsome, 2001) brains.”

“We note, however, that previous research has emphasized the role of intraparietal, not ventral parietal, regions in information accumulation in both memory (Yago and Ishai, 2006; Ploran et al., 2007) and nonmemory domains (Colby and Goldberg, 1999; Platt and Glimcher, 1999).”

“…is supported by a number of functional imaging studies (Tagamets et al., 2000; Price and Devlin, 2003; Xue et al., 2006; Ben-Shachar et al., 2007; Ploran et al., 2007; Starrfelt and Gerlach, 2007; Xue and Poldrack, 2007; Mei et al., 2010; Van Doren et al., 2010; Kherif et al., 2011), as well as…”

“Another innovative approach, designed to compensate for the poor temporal resolution of MRI, has been to use tasks in which the sensory evidence is gradually strengthened as the trial progresses and then to identify regions whose BOLD response scales with the evidence time course (Ploran et al., 2007; Ivanoff et al.,…”

“Recent work has suggested that these areas accumulate information to the time of recognition (Ploran et al., 2007), support the persistence of a percept (Philiastides and Sajda, 2007), and therefore may contribute to the comparison of sensory evidence during decision making.”

“Various functional imaging studies have demonstrated that the AIC is uniquely activated during all bodily feelings and all emotional feelings, and in direct correlation with subjective awareness of one’s body and its movements [24–26], with recognition of one’s own image [27], with one’s subjective visual and auditory…”

“…Shadlen, 1999; Shadlen and Newsome, 2001; Roitman and Shadlen, 2002; Schall, 2002; Mazurek et al., 2003; Huk and Shadlen, 2005; Hanks et al., 2006; Gold and Shadlen, 2007) and neuroimage studies (Ploran et al., 2007; Heekeren et al., 2008; Ho et al., 2009; Kayser et al., 2010; Zhang et al., 2012).”

“…dot motion direction decisions (Kayser et al., 2010a,b; Liu and Pleskac, 2011; Rahnev et al., 2011); recognition of gradually revealed objects (Ploran et al., 2007, 2011); categorization of distorted dot-pattern configurations (Vogels et al., 2002); categorical decisions of animated…”

“…of perceptual decisionmaking processing in humans using neuroimaging, such as fMRI, EEG, and MEG (Donner et al., 2007; Philiastides and Sajda, 2007; Ploran et al., 2007; Heekeren et al., 2008; Tosoni et al., 2008; Sajda et al., 2009; Noppeney et al., 2010; Philiastides et al., 2011; Siegel et al.,…”

“However, perceptual processing takes time (Schyns and Oliva, 1994; Ploran et al., 2007) and encoding can be easily disrupted by new input (Breitmeyer and Ganz, 1976; Potter et al., 2004).”

“…neurophysiological and behavioral experiments, as well as from modeling, suggesting that metacognitive judgments could be based on evidence unavailable to a type 1 response (Petrusic and Baranski, 2003; Ploran et al., 2007; Resulaj et al., 2009; Hilgenstock et al., 2014; Graziano et al., 2015).”

“The activity of neuronal populations in parietal cortex correlates with perceptual decisions in nonhuman primates (Shadlen and Newsome, 1996, 2001; Hanks et al., 2006; Kiani and Shadlen, 2009) and human subjects (Heekeren et al., 2004; Ploran et al., 2007; Tosoni et al., 2008; Ho et al., 2009).”

“Using the processGLM, we created group voxelwise statistical maps corresponding to each process in which participant was treated as a random effect.”

“Human imaging studies have begun using simple two-choice tasks to explore the neural substrates of visual perceptual decision-making (Heekeren et al., 2004; Huettel et al., 2005; Ivanoff et al., 2008; Noppeney et al., 2010; Ploran et al., 2007; Tosoni et al., 2008).”

“Here, there is substantial evidence that decision termination is mediated by application of a threshold to the neural representation of accumulating evidence [8, 46, 47], and this also holds in experiments that do not allow for free responses, as in our controlled-duration experiment [13, 16, 48].”

“One key aspect of decision processing is clearly the detection of errors, and evidence suggests that frontal retrieval success regions are likely to play a role in the overall decision process (see Dosenbach et al., 2006, 2008; Wheeler et al., 2008; Ploran et al., 2007).”

“S1, the maps also pointed to other brain regions that past work suggests are associated with criterion setting and evidence accumulation in decision making, including areas of posterior parietal cortex as well as the caudate (27, 28, 31, 32, 34, 35).”

“This interpretation is in line with a recent study that investigated perceptual recognition processes with fMRI and found that activation of the IFG was related to the moment of recognition (Ploran et al., 2007).”

“Of course, this does not mean that other regions (in left superior lateral posterior parietal cortex or elsewhere) may not show the more continuous gradations in retrieval-related activity that would be expected of a region supporting accumulation of generic evidence about the study status of a test item (for e.g., se…”

“In addition, human neuroimaging studies have localized patterns of evidence accumulation in the dorsolateral prefrontal cortex (dlPFC) (Bowman, Kording, & Gottfried, 2012; Heekeren, Marrett, Bandettini, & Ungerleider, 2004; Ploran et al., 2007; Philiastides, Auksztulewicz, Heekeren, & Blankenburg, 2011; Ploran, Tremel…”

“…online at or near the end of the trial, suggesting that these responses serve as feedback into the task set, whereas the signals in the frontoparietal network effects begin early in the trial and may be more related to accumulating information over the course of the trial (Ploran et al., 2007).”

“However, there has been evidence from fMRI studies that the human parietal cortex is also involved in accumulating the decision variable (Ploran et al., 2007; Heekeren et al., 2008).”

“2000; Menon and Uddin 2010) or by behaviorally relevant events (Dosenbach et al. 2006; Ploran et al. 2007; Nelson et al. 2010) is widely accepted.”

“Recent work on perceptual recognition found that activity in the AI/FO region remained near baseline until the moment of recognition, suggesting a relation to the moment of the decision itself (Ploran et al., 2007).”

“…2010; Heekeren et al., 2004, 2006; Ho et al., 2009; Kayser et al., 2010a, 2010b; Kovács et al., 2010; Li et al., 2009; Liu and Pleskac, 2011; Ploran et al., 2007; Tosoni et al., 2008), the vast majority of these studies focused on which brain areas are " active " during the decision process.”

“…baseline and also similar to those seen previously (Binder et al., 2004; Grinband et al., 2006; Heekeren et al., 2006; Philiastides and Sajda, 2007; Ploran et al., 2007; Thielscher and Pessoa, 2007; Kayser et al., 2010), including the bilateral IPS, IFS, frontal eye fields (FEFs), and anterior…”

“…method is not limited to the application of action decisions, as similar approaches have been successfully used in previous fMRI studies in understanding analogous accumulation processes during perceptual decisions (Heekeren et al., 2008; Ho et al., 2009; Kayser et al., 2010; Ploran et al., 2007).”

“The ACC has been implicated in neurobiological models of cognitive control, the inhibition of competing or prepotent responses, the mediation of conflict, and reward and motivation [46,47], particularly in decision making [48-50].”

“…was employed to see if regions segregated into groups on the basis of their profile of time courses for the five conditions (high-frequency words, oneand three-syllable low-frequency words and one- and threesyllable nonwords; Ploran et al., 2007; Salvador et al., 2005; Cordes et al., 2002).”

“, 2007) and fMRI (Ploran et al., 2007) in healthy subjects suggest that visual recognition involves a progressive accumulation of perceptual evidence before reaching threshold for accurate stimulus identification, which may require more time when stimuli are degraded (see also Thorpe, Fize, & Marlot, 1996).”

“This method has been extended to other perceptual domains, including recognition of simple line drawings (Ploran et al., 2007), detection of net orientation of a set of stimuli (de Lange et al., 2011) and discrimination between two ambiguous visual object classes, e.g. faces versus houses (Heekeren…”

“Third, this region may play the role of a mnemonic accumulator, such that it accumulates evidence until a criterion for the decision-making of recognition is reached [28].”

“The IPS has been associated with accumulation of information for perceptual decisions in many functional imaging studies (Ploran et al., 2007, 2011; Kayser et al., 2010; Sestieri et al., 2014).”

“The anterior temporal activation converges with the results of neuropsychological studies showing permanent retrograde amnesia following anterior temporal cortical damage and with the results of neuroimaging studies on semantic knowledge (Kapur et al., 1992; Damasio et al., 1996; Gorno-Tempini et al., 1998; Cabeza and…”

“The perceptual closure tests included degraded visual objects and word forms, which are processed in inferior temporal areas rather than in early visual cortex (Grützner et al., 2010; Ploran et al., 2007; Sehatpour et al., 2006).”

“Previous research has demonstrated that similar left VLPFC areas are involved in evidence accumulation in the perceptual domain (Ploran et al., 2007, 2011).”

“Another recent fMRI study also showed how the fMRI signal can be used to examine the process of accumulation of sensory evidence [40].”

“Some recent works (Ploran et al., 2007; Craig, 2010; Nelson et al., 2010) directly suggest that the insular cortex may be involved in awareness.”

“(Ploran et al., 2007; Sun et al., 2012), as was recently shown using a CI task similar to the JOVI in healthy subjects in a study of”

“Additionally, the clear functional dissociations observed in left parietal cortex are consistent with the idea that the region likely subserves highly distinct cognitive processes, a finding which corresponds well with graph analytic techniques also suggesting considerable functional heterogeneity in parietal cortex (…”

“There are striking similarities between the patterns found in our data and the patterns reported in studies of perceptual recognition (42, 43).”

“…activation (Craig, 2005) and supports the role of the left insular cortex as a pivotal region in higher level cognitive control, attention (Ploran et al., 2007), emotional regulation (Goldin et al., 2008) and emotional processes (Craig, 2003; Critchley et al., 2002; Kong et al., 2006;…”