The Spatial and Temporal Signatures of Word Production Components: A Critical Update

“Secondly, the influential dualstream model of speech processing (Hickok and Poeppel, 2000, 2004, 2007) assumes an area at the boundary between temporal and parietal lobes (area Spt) to function as a sensorimotor interface in language production.”

“Levelt et al. (1999) assume this selection process to take place in a part of the mental lexicon (lemma level) that is linked to the conceptual level and contains information about the grammatical properties of words, such as word class or grammatical gender.”

“…prefrontal cortex, IFG, ventromedial prefrontal cortex, and posterior cingulate gyrus) identified in a recent comprehensive meta-analysis by Binder et al., 2009, see also Schwartz et al., 2009, for an excellent discussion of the clinical evidence on regions involved in conceptual…”

“With respect to the insula, this pattern of reports is better compatible with a role in articulatory planning as suggested by Dronkers (1996) than a role in articulatory execution.”

“, 1996; Hirano et al., 1997). An involvement of the bilateral STG in the internal loop of self-monitoring is suggested by data from Shergill et al. (2002) who manipulated the rate of inner speech. More recently, Tourville et al. (2008) used feedback with a shifted first formant frequency and applied SEM to the resulti…”

“Moreover, the language production system is notoriously slow – preparation before output begins takes 600 ms for a single word if primed [7,8], approximately 1000 ms if not [9], and around 1500 ms for a short clause [10].”

“Indefrey and Levelt viewed Broca’s area as critically involved in phonological processing whereas subsequent articulatory encoding is supported by motor cortices (4, 7).”

“…serial, and proceeds through conceptualization, lemma retrieval, phonological retrieval, and phonetic encoding, following a time course that seems well understood (Indefrey, 2011), extending 600–1200 ms or more before articulation depending on the ease of retrieval and the length of the turn.”

“In addition, analytical methods differ greatly between this study and previous ones: Here we measure response reliability, whereas the majority of previous studies measure signal amplitude using eventrelated averaging methods (12, 23).”

“Second, the definition of and restriction to the few brain regions comprising the classical model dramatically underestimates the number and distribution of brain regions now known to play a critical role in language comprehension (Price, 2012) and production (Indefrey, 2011).”

“In both label-based (Indefrey, 2011; Indefrey & Levelt, 2000, 2004; see also Munding, Dubarry, & Alario, 2016) and activation likelihood estimation (ALE) neuroimaging meta-analyses (Brown, Ingham, Ingham, Laird, & Fox, 2005; Eickhoff, Heim, Zilles, & Amunts, 2009; Turkeltaub, Eden, Jones, &…”

“, 2006; Levinson, 2013), since listeners will often need several hundreds of milliseconds to plan the initial words of their turn (Indefrey & Levelt, 2004; Indefrey, 2011). Under this hypothesis, listeners would need to rely on information located earlier in the speaker's turn. We address this issue in the following s…”

“…& Gagnon, 1997; Levelt, Roelofs, & Meyer, 1999; Schwartz, Dell, Martin, Gahl, & Sobel, 2006), with current neurolinguistic models attempting to map these previously defined linguistic concepts onto cortical networks (e.g., Hickok & Poeppel, 2004, 2007; Indefrey, 2011; Indefrey & Levelt, 2004).”

“In a meta-analysis of neuroimaging studies on word production, Indefrey and Levelt (2004; see also Indefrey, 2011) identified the mid- to posterior middle temporal gyrus (MTG) as important for semantically driven lexical retrieval, suggesting a posterior temporal lobe (including the mid-posterior…”

“…et al., 2012) and activation maxima slowly progressing from posterior towards anterior brain regions are frequently reported in response to object naming (e.g. Hulten et al., 2009; Indefrey, 2011; I&L, 2004; Liljestrom et al., 2009; Salmelin et al., 1994; Soros et al., 2003; Vihla et al., 2006).”

“…(Marinkovic et al., 2003) corresponding to the www.frontiersin.org February 2015 | Volume 6 | Article 71 | 1 network of brain areas observed in functional magnetic resonance imaging (fMRI) studies of speech processing (Binder et al., 2000; Hickok and Poeppel, 2007; Rauschecker and Scott, 2009).”

“, the AG and SMG) also is associated with sentence level semantic representations and sentence comprehension (Indefrey, 2011; Indefrey & Levelt, 2004; Schönberger et al., 2014; Vigneau et al., 2006); though note that our analyses revealed no activation peaks in the SMG for sentence comprehension.”

“Some of the most influental models of linguistic production (e.g., Frederiksen, Bracewell, Breuleux, & Renaud, 1990; Indefrey, 2011) postulate that the process of lexical generation relies on three major stages: (1) a pre-linguistic conceptual phase, where the speaker generates a mental plan of…”

“Keywords: language production, lexical access, MEG”

“…detection in internal speech to take place through speech perception processes (Hartsuiker & Kolk, 2001; Levelt, 1983, 1989; Indefrey and Levelt, 2004; Indefrey 2011; Hickok 2012), but rather supports a conflict monitoring model of error detection in speech, as proposed by Nozari et al. (2011).”

“As Levinson (2013) has argued, short gaps between turns do not provide adequate time to prepare even a simple next turn, which psycholinguistic research has shown requires at least 600 ms (Indefrey and Levelt, 2004; Indefrey, 2011).”

“1 Stages of speech production according to Indefrey’s [3] model.”

“25,26 However, a recent study found stimulating some speech arrest sites in vPrCG also elicited the arrest of ongoing manual movements (negative motor responses).”

“…in these areas was between 0 and 800 ms (Acheson et al., 2011; Hulten et al., 2009; Salmelin et al., 1994; Soros et al., 2003; Vihla et al., 2006) with a median value of 190 ms for mMTG, 320 ms for pSTG, 360 ms for pMTG, 371 ms for mSTG, 280 ms for SMG, and 300 ms for anG (Indefrey, 2011).”

“However, according to time estimates from meta-analyses [37,38], picture name planning reaches the articulatory buffer no earlier than about 145 ms before articulation onset.”

“However, this is not the case, since Indefrey (2011) does rely on studies which solely manipulate one and arguably more ambiguous variables (e.g., SI in Aristei et al., 2011; name agreement in Cheng et al., 2010; etc.) as independent sources of evidence concerning time course. www.frontiersin.org…”

“This differentiation is useful for separating the motor part of language from conceptual, phonological, and semantic processing of language [39].”

“…the neural substrates associated with the processes of regulating and monitoring language production have remained relatively underspecified (cf. Indefrey, 2011; for recent advances, see Nozari et al., 2011; Riès et al., 2011), in part because the manipulations and comparisons within these…”

“…the psycholinguistic models of lexical production (Caramazza 1997; Dell and O’Seaghdha 1992; Levelt 1992) consider three main stages of word retrieval and generation, i.e., conceptual (lexical), meaning (lexical retrieval, semantic), and phonological–phonetic (for details, see Indefrey 2011).”

“EEG studies have been used to study the time courses associated with the processing stages in word production (see Indefrey, 2011 for review).”

“Dynamics of function – interpretation according to Indefrey’s attribution of functional anatomy Table S1 lists all areas mentioned as being responsible for a function in Indefrey and Levelt’s model of speech production (Indefrey, 2011; Indefrey & Levelt, 2004).”

“However, the beta-power decrease in the naming task was more broadly distributed over the left temporal cortex, extending more anteriorly to temporal areas strongly associated with conceptualizing and lexical retrieval in word production [e.g., Baldo et al., 2013; Indefrey, 2011; Indefrey and Levelt, 2004; Lambon Ralp…”

“This proposition was based on considerations and models by Indefrey (2011), Guenther and Vladusich (2012), Hickok (2012), and Tian and Poeppel (2013), among others.”

“The modal conversational response time is extremely short, around 200 ms (18, 19)—three times faster than the average speed with which people can name an object (20, 21) and too rapid to rely on deliberative conscious control (22).”

“…localizing the functional processes assumed by the psycholinguistic model (i.e., WEAVER þþ ) to anatomical gray-matter areas of the human brain (Indefrey, 2011a; Indefrey & Levelt, 2004); (3) tracto-graphic evidence concerning the structure and anatomical white-matter connections of the AF…”

“Additionally, given that the aMTG region is sometimes implicated in effortful retrieval attempts (Han, O'Connor, Eslick, & Dobbins, 2012; Lind ın, D ıaz, Capilla, Ortiz, & Maest u, 2010), especially the retrieval and selection of more peripheral thematic relations (Indefrey, 2011; Taylor, Rastle, & Davis, 2013) and se…”

“This assumed timeline is often borrowed from Indefrey and Levelt (2004) or its upgraded form (Indefrey, 2011).”

“The color bar at the bottom limits the temporal influence to − 400 ms prior to PSO, consistent with previous reports about speech planning 39 and articulatory representations 19 .”

“It is now known that the anatomical foundations of speech comprehension and production and their interrelation are more complex (Hickok & Poeppel, 2007; Indefrey, 2011).”

“…are thought to involve various areas in the MTG and superior temporal gyrus (STG) (Damasio et al. 2004; Indefrey 2007; Dronkers et al. 2000; Indefrey 2011; Indefrey and Levelt 2004; Philipose et al. 2007; Hickok and Poeppel 2000; Indefrey and Levelt 2000; Poeppel and Hickok 2004), with…”

“…Janssen et al., 2011; Maess et al., 2002; although see Janssen et al., 2014 for activity in a later time-window: 500–750 ms post-stimulus), which is in the time-window typically attributed to lexical selection (Indefrey, 2011; Indefrey & Levelt, 2004; Levelt et al., 1999; Strijkers & Costa, 2011).”

“A large body of psycho-and neuro-linguistic have shown the effect of specific lexical and image properties on word production (e.g., Indefrey, 2011).”

“Lexical access takes place between about 200 ms post picture-onset and about 145 ms before articulation onset (Indefrey, 2011; Indefrey & Levelt, 2004).”

“, 2013; Zhang, Yu, Zhang, Jin, & Li, 2018), arguing for a role of this region in syllabification (Indefrey, 2011).”

“These included the middle STS—an area associated with lexical (lemma) selection and retrieval in studies of spoken languages (29)—and the posterior STS.”

“Following the proposals made by Indefrey (2011), they argued that cognitive processes are common up to the point of lexeme encoding.”

“Obviously, rapid turn taking between speakers requires quick cognitive processing, considering that it takes at least 600ms to prepare a single word production (Indefrey and Levelt, 2004; Indefrey, 2011).”

“In general, the brain areas engaged during picture naming include the left posterior inferior frontal gyrus, the left precentral gyrus, the supplementary motor area, the left middle and posterior parts of the STG and middle temporal gyri, the right middle STG, the left fusiform gyrus, the left anterior insula, the lef…”

“This resulted in a map of the involved brain areas and the time course of their activation (see also the update by Indefrey, 2011).”

“Background The restriction of language function to the classical Broca’s or Wernicke’s area is not compatible anymore [1-3].”

“The finding that word frequency modulated activity in the left anterior and posterior-superior temporal cortex, and the left IFG with a similar temporal profile already calls for a revision of serial models of language production which assume that early lexical processes are confined to the temporal cortex (Indefrey,…”

“Theoretically, competition models of lexical selection assume that early effects resulting from lexical competition should emerge in the time window of around 200-250ms (Indefry & Levelt, 2004; Indefrey, 2011).”

“5 s, we applied rTMS pulses 300 ms after the picture presentation onset, according to our current knowledge of the timing of naming-related activity reported in previous studies (Indefrey, 2011; Salmelin et al., 2000; Wheat et al., 2013).”

“…language processing, including the inferior frontal gyrus (Broca's area) and superior temporal cortex (Wernicke's area) (Hickok and Poeppel, 2007; Indefrey, 2011; Price, 2010), and those for language control such as the dorsal lateral prefrontal cortex, dorsal anterior cingulate cortex (dACC 1 )…”

“…speech monitoring (Alario, Chainay, Lehericy, & Cohen, 2006; Chauvel, Rey, Buser, & Bancaud, 1996; Crosson et al., 2001; Fried et al., 1991; Gabarrós et al., 2011; Indefrey, 2011; Indefrey & Levelt, 2004; Krainik et al., 2003; Laplane, Talairach, Meininger, Bancaud, & Orgogozo, 1977; Pai, 1999).”

“The positive associations between the phonological processing parameter p and both left AF and MdLF are also consistent with findings from meta-analyses of functional activation studies of word production (Indefrey and Levelt, 2004; Indefrey, 2011).”

“Seven participants were excluded from the EEG analysis because of large electrode drift and excessive artifacts.”

“However, there is also a reason to assume that a certain amount of rescaling 23 should be used (see also Indefrey, 2011).”

“In a meta-analysis of word production studies, Indefrey and Levelt (2004) and Indefrey (2011), see also Costa et al. (2009), estimated that lexical selection takes place between about 200 to 275 ms after picture onset and phonological code retrieval between 275 and 355 ms. Based on these estimates,…”

“Accordingly, psycholinguistic research has availed itself of electrophysiological and neuroimaging data in order to develop theory relevant to combined cognitive and brain operations, beginning with the meta-analysis of Indefrey and Levelt (2004; also Indefrey, 2011).”

“2) and our cognitive interpretation of the data (Table 1) are in agreement with many previous findings (Edwards et al., 2010; Indefrey, 2011; Leuthardt et al., 2012; Llorens et al., 2011; see also Munding et al., 2015; Price, 2012).”

“We hypothesized that pause distribution by word class would predict atrophy in regions supporting lemma retrieval, such as the MTG (Indefrey, 2011; Indefrey and Levelt, 2004; Schwartz et al., 2009; de Zubicaray et al., 2002) and/or the IFG, which has been claimed to exert top-down control over…”

“We note that activity in medial frontal regions is not included in the model proposed by [5] contrarily to that proposed by [3].”

“…duration), it is possible that the facilitation occurred already at the initiation phase when the phonological planning of the grammatical item retrieval occurred, as articulation takes place at an even later point after stimulus onset and involves more posterior cortical regions (Indefrey, 2011).”

“ing on language processing unveiled the existence of an articulated cognitive architecture implemented in an extensive array of neural networks (Indefrey, 2011; Indefrey & Levelt, 2004).”

“This is because, as discussed earlier and as pointed out by both Papoutsi et al. (2009) and Indefrey (2011), the left frontal lobe not only implements phonetic encoding and articulation, but also subserves the more abstract process of syllabification.”

“In terms of models proposed for speech production [Indefrey, 2011; Indefrey and Levelt, 2004], differences between the two tasks in the early time window would be associated primarily with perceptual or lexical processing, while processes related to motor planning of speech should be captured within the late time wind…”

“As is evident from previous reviews (Indefrey and Levelt, 2004; Indefrey, 2011) this issue is particularly complex and our current knowledge, which relies on the comparison of disparate sources of evidence, is still incomplete.”

“Given that experimental research has shown that speakers need at least 600 ms to plan even a simple word (Indefrey and Levelt, 2004; Indefrey, 2011), processes of language production in conversation must begin well before the current turn ends (Levinson, 2013).”

“In the model, self monitoring occurs via an inner loop that utilizes speech perception areas in the superior temporal gyrus (Indefrey, 2011), but the precise mechanism of monitoring is not critical for the current study (for further discussion of self-monitoring in IS, see Discussion Section 4.4).”

“Nevertheless, the early onset of the nTMS may also influence conceptual preparation, lexical concept selection, and lemma retrieval attributed to the early stages of picture naming processes (Indefrey and Levelt, 2004; Indefrey, 2011).”

“Unfortunately, incomplete coverage did not allow us to test existing neuroanatomical picture naming (Indefrey and Levelt, 2004; Indefrey, 2011; Roelofs, 2014) and semantic processing (Binder and Desai, 2011; Gainotti, 2011; Lambon Ralph et al., 2017; Pulvermüller, 2013) accounts that attribute…”