Verb generation in patients with focal frontal lesions: A neuropsychological test of neuroimaging findings

“…semanic processing, as performance on a variety of semantic tasks is isrupted by damage to left VLPFC (Martin & Cheng, 2006; etzler, 2001; Thompson-Schill et al., 1998), by intraoperative timulation of left VLPFC (Klein et al., 1997), and by applicaion of transcranial magnetic…”

“…et al., 2004; Metzler, 2001; Noppeney and Price, 2004; Petersen et al., 1988; Poldrack et al., 1999; Sohn et al., 2003; Swick and Knight, 1996; Thompson-Schill et al., 1998; Wagner et al., 2001; Zhang et al., 2004), though the functional character and topographic organization of processing…”

“Rather, the posterior and dorsal extent of LIPC ( Brodmann’s area [BA] 44) has been observed in studies of selection or interference resolution (Thompson-Schill et al., 1997; Jonides et al., 1998; Thompson-Schill et al., 1998).”

“This parallel area of research has examined the role of the posterior left inferior frontal gyrus (LIFG), or Broca's area, in executive functions involving the contribution of inhibitory control processes, such as semantic selection [30] and the resolution of proactive interference in working memory [31].”

“…alternatives—that is, as the number of intervening semantically related items rises (see also Thompson- Schill et al., 1997, who reported LIFG activation in healthy adults performing a similar task; and Thompson-Schill et al., 1998, who suggested that LIFG is necessary for such a task).”

“…long been associated with cognitive control and there is strong neuropsychological evidence linking left inferior frontal gyrus (LIFG) to deficits in the control of linguistic and semantic processing (Schnur et al., 2009; Thompson-Schill et al., 1998; Novick et al., 2009; Robinson et al., 2010).”

“Thompson-Schill et al. (1998) observed that among aphasia patients with lesion that included the L inferior frontal gyrus (L IFG), there was a direct correlation between extent of lesion within L BA 44, and selection-related errors on a task requiring the subject to generate a verb for a written…”

“Thompson-Schill and colleagues have proposed that LIFG is necessary for the selection of semantic information from competing alternatives and not semantic retrieval per se[101,102,104] .”

“The experiment reported here examined the selection demands (Thompson-Schill et al., 1998).”

“Left prefrontal regions have been implicated in interference tasks involving verbal stimuli (Taylor et al., 1994; Thompson-Schill et al., 1997, 1998; Desmond et al., 1998; Jonides et al., 1998; D'Esposito et al., 1999; Leung et al., 2000; Macdonald et al., 2000), but right lateral prefrontal…”

“…deficits in situations characterized by strong competition between potential responses, increasing the need for semantic selection (e.g., during sentence completion tasks with low vs. high predictive endings) (Robinson et al. 1998, 2005; Novick et al. 2009; see also Thompson-Schill et al. 1998).”

“…for language, per se, but is more generally important for selecting among competing representations, especially when there is a need to override a prepotent response or when there are a large number of equally probable response options (Novick et al., 2005; Thompson-Schill et al., 1998).”

“Although often associated with semantic retrieval, recent evidence suggests that this region is involved in tasks requiring selection among strongly competing alternatives (Thompson-Schill et al., 1997, 1998 ).”

“The left inferior frontal gyrus may serve as a hub that mediates between conflicting representations (Thompson-Schill et al., 1998), and this feature may be represented in its fluctuating network betweenness centrality (Qin et al.”

“…explain its activation by both semantic and phonologic tasks as these processes usually place high demands on the selection of a particular phonologic or semantic feature, respectively, among many other potentially relevant options [Barde and Thompson-Schill, 2002; Thompson-Schill et al, 1998].”

“Consistent with our claim that belief-laden reasoning involves a left temporal system are findings that implicate this system in the semantic retrieval and selection of information (Thompson-Schill, D’Esposito, Aguirre, & Farah, 1997; Thompson-Schill et al., 1998).”

“…Wise et al., 1991; McCarthy et al., 1993; Petersen and Fiez, 1993; Raichle et al., 1994; Buckner et al., 1995; Passingham, 1996; Seger et al., 1997; Thompson-Schill et al., 1998; Barch et al., 2000; Holland et al., 2001; Ojemann et al., 2002; Schlaggar et al., 2002b; Burton et al., 2003; Corina et…”

“Damage to the left frontal lobe, especially to left inferior frontal cortex (LIFC) has consistently been shown to impair verbal fluency, even in patients who are not overtly aphasic (e.g., Baldo and Shimamura, 1998; Milner, 1964; Thompson-Schill et al., 1998).”

“…have been under continuous inquiry in the neuroimaging (for reviews see Fiez and Petersen 1998; Gabrieli et al. 1998) and neuropsychology literature (Illes et al. 1999; Thompson-Schill et al. 1998) since the verb generation task was first employed in functional imaging (Petersen 1988, 1989).”

“…with LIFC lesions do not typically have semantic memory deficits, although their ability to retrieve specific semantic information, select among multiple options, or inhibit competing alternatives can be impaired (Costello and Warrington, 1989; Robinson et al., 1998; Thompson-Schill et al., 1998).”

“(A) In the ROI analyses of lesion-deficit associations, patients who exhibited a large (HIGH) growth of interference across cycles of blocked naming (n 7) exhibited a greater extent of damage to the LIFG (expressed as a percentage of ROI volume) than did patients with a small (LOW) growth effect [n 5; t(10) 3.”

“Future research with targeted transcranial magnetic stimulation (e.g. Mottaghy et al., 2002) or consideration of select lesion populations (e.g. Thompson-Schill et al., 1998) may provide further evidence for the multi-component nature of analogical reasoning.”

“…line of studies of LIPC functioning has been Semantic Words 1108 (198)conducted by Thompson-Schill and her colleagues Phonological Words 1203 (254)(Thompson-Schill et al., 1997, 1998, 1999), who have Pseudowords 1359 (251) suggested that a portion of pLIPC (near BA 44) is critical Letter Words…”

“Moreover, in a PET study (Frith, Friston, Liddle, & Frackowiak, 1991) verbal fluency performance was associated with activation in the prefrontal cortex (see also Thompson-Schill et al., 1998).”

“Furthermore, patients with VLPFC damage were impaired at retrieving verbs only under conditions of increased competition [49].”

“The posterior part of Broca's area has been described as essential for the retrieval and selection of factual knowledg”

“…with many competing alternatives (e.g., cat); further- ore, the magnitude of this effect was correlated specifically ith the extent of LIFG damage (Thompson-Schill et al., 1998). group of patients with Alzheimer’s disease who exhibited his same competition effect on verb generation, were also…”

“…structures in the left premotor/prefrontal region and in the left posterior temporal regions (Caramazza and Hillis, 1991; Damasio and Tranel, 1993; Daniele et al., 1994; Hillis and Caramazza, 1995; Miceli et al., 1988; Miozzo et al., 1994; Thompson-Schill et al., 1998; Zingneser and Berndt, 1990).”

“…Pollman, Muller, & von Cramon, 2002; Monchi, Petrides, Petre, Worsley, & Dagher, 2001) and controlled retrieval tasks (Sohn, Goode, Stenger, Carter, & Anderson, 2003; Wheeler & Buckner, 2003; Gold & Buckner, 2002; Thompson-Schill et al., 1998; Thompson-Schill, D’Esposito, Aguirre, & Farah, 1997).”

“…2005; Aron and Poldrack 2006; Li et al. 2006; Chikazoe et al. 2007; Leung and Cai 2007), and also with those of other types of inhibitory control (Thompson-Schill et al. 1997, 1998; Monchi et al. 2001; Braver et al. 2003; Hazeltine et al. 2003; Konishi et al. 2003, 2005; Brass and von Cramon…”

“, 1991), and corroborated by lesion-based work (Thompson-Schill et al., 1998).”

“It should also be clarified that the lesion overlap approach taken here differs from other published approaches that define patient groups on the basis of their lesion overlap and then assess any behavioral difference”

“Recent fMRI and patient studies suggest that left prefrontal regions have a role in selecting information from semantic memory (Thompson-Schill, D’Esposito, Aguirre, & Farah, 1997; Thompson-Schill et al., 1998; Thompson-Schill, D’Esposito, & Kan, 1999).”

“Unfortunately, many standard language tasks, especially those that are popular in clinical practice (such as verb generation [83]), confound linguistic and general cognitive demands, making the resulting activations difficult to interpret.”

“Patients with damage to LIFG after SA are impaired across a variety of semantic control manipulations, including some of the tasks used in the current study (i.e., relatedness judgments with strong and weakly related targets; Novick et al., 2009; Jefferies & Lambon Ralph, 2006; Thompson-Schill et al., 1998).”

“…of left lateral frontal involvement when selection among verbal alternatives is required (although the areas involved here may be somewhat different from those involved in tasks that include explicit interference: for example, Desmond, Gabrieli, & Glover, 1998; Thompson-Schill et al., 1998).”

“Moreover, Thompson-Schill and colleagues have reported that patients with LIPC lesions demonstrate impaired performance under conditions associated with high, but not low, selection demands (Thompson-Schill et al., 1998).”

“The ROI was introduced to the analyses in the form of mask, constructed from the Talairach and Tournoux (1988) atlas of the human brain, by outlining the regions on the atlas and transforming the outlines to a digital image.”

“…competing semantic alternatives activates ventrolateral prefrontal cortex in the inferior frontal gyrus (Thompson-Schill et al., 1997; Crosson et al., 2001; Tremblay and Gracco, 2006, but see Thompson-Schill et al., 1998; Fletcher and Henson, 2001; Rowe and Passingham, 2001; Nagel et al., 2008).”

“The association between IFG and semantic control is supported by TMS studies (Whitney, Kirk, et al., 2011, Whitney et al., 2012) and investigations of patients with IFG lesions (Bedny, Hulbert, & Thompson-Schill, 2007; Noonan et al., 2010; Robinson et al., 2010; Thompson-Schill et al., 1998).”

“For example, although damage to the left inferior frontal gyrus results in word-finding deficits, it does not disrupt conceptual knowledge per se (Baldo and Shimamura 1998; Thompson-Schill et al. 1998; Price et al. 1999).”

“functional neuroimaging studies also implicate fronto-temporal cortex ( Thompson-Schill et al., 1998 ).”

“…the same semantic features that were initially retrieved are required for the subsequent decision and declines when different semantic representations from those initially retrieved are subsequently required (Thompson-Schill & Gabrieli, 1999; Thompson-Schill et al., 1998; Vriezen et al., 1995).”

“For example, VLPFC but not DLPFC activity has been reported during tasks that require maintaining, retrieving and/or selecting detailed item information (Thompson-Schill et al., 1998 ),.”

“…associated with semantic association and speech production, verb generation also involves a speech perception phase that involves posterior temporal regions (Edwards et al., 2010; Herholz et al., 1997; Ojemann et al., 2002; Petersen et al., 1988; Thiel et al., 1998; Thompson-Schill et al., 1998).”

“The available evidence consistently implicates inferior and middle frontal gyri (IFG and MFG), in addition to other widely distributed cortical areas, in the semantic association/selection aspects of the verb generation task (Petersen et al., 1988; Herholz et al., 1997; Thiel et al., 1998; Thompson-Schill et al., 1998…”

“With increasing demands on selection between competing alternatives, IFG activity in normal subjects increases and performance of patients with IFG lesions deteriorates (Thompson-Schill et al. 1998; Kan et al. 2006).”

“This view is in line with functional neuroimaging (Fiez, 1997; Thompson-Schill et al., 1997; Badre et al., 2005) and neuropsychological studies (Thompson-Schill et al., 1998; Jefferies and Lambon Ralph, 2006; Noonan et al., 2010) linking left VLPFC with executive regulation of semantic knowledge.”

“These results suggest that the LIFG may be a candidate region for conceptual selection (see also Kan & Thompson-Schill, 2004; ThompsonSchill, D’Esposito, & Kan, 1999; Thompson-Schill et al., 1998).”

“ably to normal people when the cue noun had few or only one dominate associate (Thompson-Schill et al., 1998).”

“In patients, lesions impair performance on tasks that require selection amongst many possible alternative responses (Thompson-Schill et al., 1998).”

“…patients with damage to PFC demonstrate deficits in memory tasks that involve strategic control of memory, goaldirected manipulation of mnemonic information, or overcoming interference (Moscovitch, 1992; Stuss et al., 1994; Wheeler et al., 1995; Aly et al., 2011; Thompson-Schill et al., 1998).”

“Thompson-Schill et al. (1998) examined the ability of patients with focal frontal lesions to perform a verb generation task under high and low selection demands.”

“SA patients perform poorly on semantic tasks that require selection among competing responses, particularly when the most obvious or prepotent response is not the correct one (Jefferies & Lambon Ralph, 2006; Thompson-Schill et al., 1998).”

“In patients with focal brain lesions, impairment of both phonemic and semantic fluency has been found after frontal lobe damage [2,3].”

“These questions are closely related to the normal contribution of left IFG to word retrieval and the interpre-Thompson-Schill et al., 1998).”

“Moreover, evidence suggests that brain-damaged patients with deficient inhibiting abilities have difficulty producing words under conditions of high lexical competition in a word generation task (e.g., Badre, Poldrack, Paré-Blagoev, Insler, & Wagner, 2005; Thompson-Schill et al., 1998).”

“semantic memory and language processing (Metzler, 2001; Thompson-Schill et al., 1998).”

“There is also evidence that focal brain damage involving IFG selectively disrupts performance on verb generation tasks that require high demands on cognitive control (Thompson-Schill et al., 1998).”

“…comprehension, while those with damage to LIFG show semantic retrieval problems but relatively intact knowledge of word meaning in tasks that minimize executive control demands (Hart and Gordon 1990; Chertkow et al. 1997; Thompson-Schill et al. 1998; Robinson et al. 2005; Novick et al. 2009).”

“…subcategories (Rosen and Engle, 1997) and actively shifting between them (Troyer et al., 1997; Reverberi et al., 2006), semantic fluency may thus require additional control processes such as selection of appropriate items from competing alternatives (Thompson-Schill et al., 1997, 1998).”