The neural correlates of the verbal component of working memory

“…e.g., Marr, 1969; Tach, 1997; Procedural & associative Poldrack & Gabrieli, 2001 learning e.g., Bracke-Tolkmitt et al., 1989 Memory Long-term memory e.g., Appollonio et al., 1993 Phonological short-term memory e.g., Paulesu et al., 1993 Imagery Visuomotor imagery e.g., Decety et al., 1990 Note.”

“First we observed no activation in the left frontal operculum including Broca’s area, whose involvement is usually reported for phonologic processing (Fiez and Petersen, 1998; Price et al., 1996; Pugh et al., 1996) and overt or subvocal articulation (Dogil et al., 2002; Paulesu et al., 1993; Smith and Jonides, 1999).”

“Both tasks also produced activation in left Broca’s area (BA 44), which is a region believed to mediate articulatory rehearsal during operation of the phonological loop (Awh et al., 1996; Paulesu et al., 1993).”

“The dorsolateral prefrontal cortex is implicated in executive control, visual working memory, tasks requiring planning, deriving abstract criteria, and changing criteria in cognitive tasks (Grafman, 1989; Paulesu et al., 1993; D’Esposito et al., 1995).”

“…foci reported within the dorsolateral and ventrolateral frontal cortices during 27 verbal working-memory studies (Grasby et al. 1993, 1994; Paulesu et al. 1993; Petrides et al. 1993b; Becker et al. 1994; Cohen et al. 1994, 1997; Andreasen et al. 1995; Awh et al. 1996; Coull et al. 1996;…”

“Activation of this area has been reported in working memory tasks manipulating information type (Paulesu et al., 1993, 1996; Salmon et al., 1996) and in passive listening paradigms (Fiez et al., 1996; Petersen et al., 1998).”

“…version of this task has been shown to give rise to significant activations in left parietal, inferior frontal, and motor areas (Awh et al., 1996; Paulesu et al., 1993) where the left inferior parietal regions may serve as a phonological store, and the VLPFC as a subvocal rehearsal system…”

“For example, the subvocal rehearsal mechanism appears to involve the right cerebellum, apart from left inferior frontal gyrus, supplementary motor area, and insula (34).”

“, 1999), and left inferior try in the amount of white matter in faster learners frontal regions in and around Broca’s area (Zatorre et may be related to greater myelination allowing more al., 1992, 1996; Paulesu et al., 1993; Fiez et al., 1995; efficient neural processing, which is critical for the Burton et al., 2000…”

“This view is not inconsistent with the interpretation that Broca’s area mediates verbal working memory [Paulesu et al., 1993; Pöppel, 1996].”

“Along with its established role in taste perception and visceral somatosensory processing, more cognitive roles for the insula in selective visual attention (Corbetta, Miezin, Dobmeyer, Shulman, & Petersen, 1991), verbal working memory ( Paulesu, Frith, & Frackowiak, 1993 ), and linguistic processing (Augustine, 1996;…”

“…of brain imaging and the finger tapping, simple auditory monitoring and modified PASAT tasks (Cabeza and Nyberg, 2000; D’Esposito et al., 1995; Desmond et al., 1997; Fletcher and Henson, 2001; Lange et al., 2005; Paulesu et al., 1993; Petrides et al., 1993; Salmon et al., 1996; Smith et al., 1998).”

“The left inferior parietal area contributes to the representation of phonetic sequences (Gelfand and Bookheimer, 2003; Paulesu et al., 1993; Small and Burton, 2002; Smith and Jonides, 1998), independent of subvocal articulatory rehearsal (Logie et al.”

“The fMRI activation patterns of bilinguals included a number of areas previously implicated in phonological and pseudoword processin”

“…judgment on pairs of auditorily presented syllables (Zatorre, Evans, Meyer, & Gjedde, 1992; Zatorre, Meyer, Gjedde, & Evans, 1996), or letters (Paulesu et al., 1993), and other letter-sound tasks (Sergent, Zuck, Levesque, & MacDonald, 1992) have resulted in increased blood ºow in the left…”

“, 1997; Fiez and Petersen, 1998) and the generation of subvocal speech (Paulesu et al., 1993; Awh et al., 1996).”

“[36] as the locus of the phonological store.”

“…have reported pre-SMA activation in motor imagery (Deiber et al., 1998; Jenkins et al., 2000; Cunnington et al., 2002; Malouin et al., 2003) as well as in other mental imagery modalities (Paulesu et al., 1993; Ghaem et al., 1997a; Halpern and Zatorre, 1999; Mellet et al., 2000b; Halpern, 2001).”

“are either held in verbal short-term memory (supramarginal gyrus; Paulesu et al. 1993; Lee et al. 2007; for review, see Jacquemot and Scott 2006) or reflect acoustically ambiguous,”

“…of the medial wall in 14 previous studies that were originally designed to be studies of either spatial or nonspatial visual working memory [PET: Paulesu et al. (1993), Smith et al. (1995), Coull et al. (1996), Courtney et al. (1996), Fiez et al. (1996), Goldberg et al. (1996), Klingberg et al.…”

“Moreover, previous PET studies have indicated that highly significant activations can occur in motor areas in the absence of muscular activity (Paulesu et al. 1993; Decety et al. 1994; Stephan et al. 1995).”

“Regions in the left precentral gyrus have been postulated to be involved in ''inner hearing'' of phonological information (Paulesu et al., 1993; Zatorre et al., 1992) or in covert articulatory rehearsal (Fiez et al., 1995).”

“However, not only have past effects generally been found bilaterally, but activity within the LIFG is usually greater than in the RIFG during memory tasks (Paulesu et al., 1993; Awh et al., 1996; Rypma et al., 1999).”

“Results of functional imaging studies have supported the idea that Broca's area is involved in subvocal rehearsal in verbal working memory (Démonet et al., 1994b; Paulesu et al., 1993).”

“This languagespecific effect can be understood from the conceptualization of the IPL as part of an auditory-motor integration circuit in speech perception (Hickok and Poeppel, 2000; Wise et al., 2001).”

“Recent tomographic and magnetic resonance studies (Decety et al. 1990; Akshoomoff et al. 1992; Paulesu et al. 1993 ) and recent studies of cerebellar patients (Fiez et al. 1992; Silveri et al. 1994) have provided further support for the involvement of the cerebellum in cognitive activities.”

“…activateddition. preferentially during veridical recognition with auditory/ phonological processing and retention (Howard et al., Discussion 1992; Paulesu et al., 1993; Petersen et al., 1989); lesions to the supramarginal gyrus are also known to disrupt This experiment has yielded several novel…”

“Regions including the left supramarginalis gyrus (BA 40) and the posterior part of the left superior temporal gyrus (BA 42) have been associated with the phonological store [Paulesu et al., 1993].”

“…parietal cortex, the inferotemporal cortex, the cingulate gyrus, and hippocampus—all of which have anatomical connections with the prefrontal cortex (Petrides and Pandya 1984; Selemon and Goldman-Rakic 1985; Goldman-Rakic 1988, 1999; Cavada and Goldman-Rakic 1989; Fuster 1997; Jonides et al. 1998).”

“, 2004], as well as the insular cortex [Bamiou et al., 2003; Paulesu et al., 1993].”

“, 1996a), inner speech (Paulesu et al., 1993) or silent reading (Bavelier et al.”

“, 2001), phonological memory storage (Paulesu et al., 1993), phonological analysis, lexical semantic processing (Just et al.”

“Indeed, this area seems to be involved in encoding phonological representations in terms of mouth articulation gestures ( Paulesu, Frith, & Frackowiak, 1993; Zatorre, Evans, Meyer, & Gjedde, 1992) and imitation of arm gestures (Tanaka & Inui, 2002).”

“Neuroimaging studies have also shown that the prefrontal cortex, as well as other areas of association cortex, are active during WM tasks (Jonides et al., 1993; Paulesu et al., 1993; McCarthy et al., 1994; Smith et al., 1995; Courtney et al., 1996; Owen et al., 1996; Smith et al., 1996; Braver et al., 1997; Cohen et a…”

“Paulesu et al. (1993) reported evidence that frontal areas mediate the rehearsal of verbal material, whereas parietal areas mediate the storage of such material.”

“Human imaging studies have shown that the area (1996) also reached the same conclusion using a maze task.corresponding to the pre-SMA was activated when encoding and retrieval of short-term memory is required (Buckner et al. 1996; Fiez et al. 1996; Paulesu et al. 1993).”

“... and cooling studies with primates (Barone & Joseph, 1989; Quintana, Fuster, & Yajeya, 1989; Fuster, Bauer, & Jervey, 1985; Bauer & Fuster, 1976; Fuster & Alexander, 1971; Kubota & Niki, 1971) and functional neuroimaging studies with humans (Nystrom et al., 2000; Smith & Jonides, 1999; Courtney, Petit, Haxby, & Ung…”

“Subvocalization would be expected to activate the rehearsal component of the articulation which has been associated with activation of the left inferior frontal gyrus (BA 44) and ventral premotor cortex (Smith & Jonides, 1999; Cohen et al., 1997; Paulesu, Frith, & Frackowiak, 1993 ).”

“The right inferior frontal cortex is activated in other tasks related to language, such as metaphor comprehension (Bottini et al. 1994) and phonological working memory (Paulesu et al. 1993), as well as in a face matching task (Haxby t al.”

“For example, the activation of primary (BA 41) and secondary (BA 22) auditory regions, premotor cortex (BA 4/6), cerebellum, and brainstem can be tied to the auditory/manual secondary task demands, while the activation of bilateral BA 40 may be linked to the greater working memory demands of the dual task situation (P…”

“The parietal region highlighted in the present study is superior to that associated with phonological rehearsal/storage (Paulesu et al., 1993; Smith et al., 1998) revealed in a prior SD study (Habeck et al., 2004), and we posit that it might be related to the maintenance of attention (Kastner and…”

“Moreover, the phonological loop store is related to the left posterior parietal cortices (BA 40), whereas the articulatory rehearsal process is related to the Broca’s area (BA 6/44) [Baddeley, 2003; Jonides et al., 1996; Paulesu et al., 1993].”

“, articulation) and phonological storage, respectively, during word reading (Paulesu et al., 1993; Stoeckel et al., 2009).”

“Verbal working memory studies have also reported correlated left-frontal and right-cerebellar activations associated with increased working memory demands ( Paulesu et al., 1993, 1995; Grasby et al., 1994; Awh et al., 1996; Fiez et al., 1996; Jonides et al., 1997).”

“Impaired activation was also revealed in the supramarginal gyrus (inferior parietal lobe area 40), which has been related to the phonological store component of working memory (Paulesu et al., 1993) and the auditory association areas of the temporal lobe, related to phonological and lexical processing.”

“These results suggest that the observed activation in Broca’s area is related to subvocal rehearsals (Zatorre et al., 1996; Paulesu et al., 1993).”

“Lesion and imaging studies, mainly based on visual stimuli, have shown that the left IPS plays an important role in the phonological storage during VWM (Awh et al., 1996; Cohen, 1992; Fiez et al., 1996; Henson et al., 2000; Jonides et al., 1998; Paulesu et al., 1993; Shallice, 1988).”

“Its activation has often been reported in language experiments, and has mainly been linked to processing of phonological information [9,10,86,91] and phonological working memory [78,84].”

“The supramarginal gyrus in particular is a key module of the auditory/phonological loop, a circuitry mediating the short-term storage and processing of auditory information and especially language (Paulesu et al., 1993).”

“These efforts have linked articulatory control processes to left inferior frontal regions and phonological storage to left inferior parietal regions ( Awh et al., 1996; Fiez & Raichle, 1997; Fiez et al., 1996; Jonides et al., 1998; Paulesu, Frith, & Frackowiak, 1993; Smith & Jonides, 1998).”

“Scientific literature is also replete with studies that have shown relatively greater activation of the prefrontal cortex due to the increased demands of load, duration or manipulation on working memory [e.g., Baker et al., 1996; Courtney et al., 1997; Jonides et al., 1998; Manoach et al., 1997; Owen et al., 1996; Pau…”

“ticulatory loop of verbal WM (Paulesu et al., 1993) and has also been associated with syntactic processing (Zurif, 1982).”

“…that have reported similar networks underlying working memory, namely, parietal regions and anterior, usually IF, regions (Hickok, Buchsbaum, Humphries, & Muftuler, 2003; Honey et al., 2000; Jonides et al., 1998; Paulesu et al., 1993; Postle, Berger, & D’Esposito, 1999; Salmon et al., 1996).”

“…(Mac- Donald et al., 2000) and anterior cingulate for on-line monitoring, error detection, and response execution (Botvinick et al., 2001; Paus, 2001), whereas the parietal cortex is thought to play a crucial role in short-term storage (Gathercole , 1994; Honey et al., 2000; Paulesu et al., 1993).”

“In more recent neuroimaging studies, various kinds of cognitive tasks induced activity in parts of BA6 (Jonides et al., 1993; Paulesu et al., 1993; Dehaene et al., 1996; Fiez et al., 1996; Mellet et al., 1996).”

“The functional status most commonly attributed to this area is that it is involved in the subvocal rehearsal system (Paulesu et al., 1993).”

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