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…”

“Keywords: language production, word production, picture naming, neuroimaging”

“Gorno-Tempini and colleagues (Gorno-Tempini et al., 2004, 2008; Henry and Gorno-Tempini, 2010; Wilson et al., 2010) described a so-called logopenic variant of PPA in which mainly gray matter in the mid to posterior temporal lobe is affected1.”

“…brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann et al., 1989; Schäffler et al., 1993; Haglund et al., 1994; Malow et al., 1996; Hamberger et al., 2001; Stewart et al., 2001).”

“COMPONENT PROCESSES OF WORD PRODUCTION Models of language production (Garrett, 1980; Stemberger, 1985; Dell, 1986; Butterworth, 1989; Levelt, 1989; Caramazza, 1997; Dell et al., 1997; Levelt et al., 1999) agree that there are processing levels of meaning, form, and articulation.”

“The I&L estimate for the duration of lemma retrieval was based on mathematical modeling of the semantic interference effect (Levelt et al., 1991; Roelofs, 1992), suggesting a lemma retrieval duration of 100–150 ms and electrophysiological data by Schmitt et al. (2001) suggesting that grammatical…”

“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.”

“COMPONENT PROCESSES OF WORD PRODUCTION Models of language production (Garrett, 1980; Stemberger, 1985; Dell, 1986; Butterworth, 1989; Levelt, 1989; Caramazza, 1997; Dell et al., 1997; Levelt et al., 1999) agree that there are processing levels of meaning, form, and articulation.”

“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.”

“…brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann et al., 1989; Schäffler et al., 1993; Haglund et al., 1994; Malow et al., 1996; Hamberger et al., 2001; Stewart et al., 2001).”

“…is characterized by anomia and phonemic paraphasias in confrontation naming, an impairment of verbal short-term memory functions, and an absence of the phonological similarity effect on letter recall, whereas conceptual knowledge seems to be relatively unaffected (Gorno-Tempini et al., 2008).”

“The I&L estimate for the duration of lemma retrieval was based on mathematical modeling of the semantic interference effect (Levelt et al., 1991; Roelofs, 1992), suggesting a lemma retrieval duration of 100–150 ms and electrophysiological data by Schmitt et al. (2001) suggesting that grammatical…”

“Gorno-Tempini and colleagues (Gorno-Tempini et al., 2004, 2008; Henry and Gorno-Tempini, 2010; Wilson et al., 2010) described a so-called logopenic variant of PPA in which mainly gray matter in the mid to posterior temporal lobe is affected1.”

“…IFG (Broca’s area) was the only remaining core area that was not more often reported in experiments using overt responses (see Murphy et al., 1997; Wise et al., 1999; Huang et al., 2001; and more recently Ackermann and Riecker, 2004, for the absence of Broca’s area activation in direct…”

“Christoffels et al., 2007b, see also van de Ven et al., 2009) studied self-monitoring using verbal feedback without distortion.”

“To find out which of these regions are in fact necessary for word production (rather than just somehow involved), Indefrey (2007) compared them with brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann…”

“Gaskell and Dumay (2003) showed that pseudowords only become fully integrated in the lexicon, i.e., showing competition effects on phonologically similar words) after consolidation during a sleep phase.”

“To date, four MEG studies (Salmelin et al., 1994; Levelt et al., 1998; Sörös et al., 2003; Hultén et al., 2009) provide timing data on the activation of posterior STG and MTG.”

“Analyzing data from 64 aphasic patients, Schwartz et al. (2009) found a significant association between semantic errors in language production and lesions in three cortical areas: the left prefrontal lobe, the left posterior MTG, and the left anterior to mid MTG. Importantly, after factoring out…”

“It is, therefore, fortunate, that one of the first studies using a hypothesis-testing approach in an fMRI experiment on word production (de Zubicaray et al., 2001) investigated the neural correlates of lemma selection. They used a semantic picture-word interference paradigm. In this paradigm, competition at the lemma…”

“, 2001; and more recently Ackermann and Riecker, 2004, for the absence of Broca’s area activation in direct comparisons of overt and covert responses). The somewhat indirect conclusion that Broca’s area is the most likely candidate area for syllabification has more recently been challenged for different reasons. First…”

“In another study, insular activation increased linearly with syllable repetition rate (Riecker et al., 2005).”

“To date, four MEG studies (Salmelin et al., 1994; Levelt et al., 1998; Sörös et al., 2003; Hultén et al., 2009) provide timing data on the activation of posterior STG and MTG.”

“Both in functional magnetic resonance imaging (fMRI) and in electroencephalography (EEG) methods have been developed that allow for overt speaking during experiments (for fMRI see, e.g., de Zubicaray et al., 2001; Grabowski et al., 2006; Christoffels et al., 2007b; Heim et al., 2009b; Hocking et al., 2009; for EEG see…”

“Davis et al. (2008) linked this behavioral effect to changes in hemodynamic activation.”

“Wheeldon and Levelt (1995) measured a 125 ms difference between RTs for the first and the last phoneme of bisyllabic words with on average six phonemes. van Turennout et al. (1997) measured a corresponding 80 ms difference for words with on average 4.5 phonemes.”

“Based on the comparison of chronometric data from reaction time studies, modeling data, and electrophysiological studies, I&L provided the following estimates for the duration of the different processing components in the picture naming task: Conceptual preparation (from picture onset to selection…”

“The I&L estimate for the duration of conceptual preparation until the selection of a target concept was based on data showing the availability of information about whether a picture showed an animal or not (Thorpe et al., 1996, around 150 ms; Schmitt et al., 2000, around 200 ms).”

“The I&L estimate for the duration of phonological code retrieval was based on the difference between the lateralized readiness potential (LRP) onsets for a grammatical gender compared to a first phoneme decision in a study by van Turennout et al. (1998).”

“Christoffels et al., 2007b, see also van de Ven et al., 2009) studied self-monitoring using verbal feedback without distortion.”

“…the interaction between brain areas (structural equation modeling, SEM; dynamic causal modeling, DCM, independent component analysis, ICA) have begun to be applied to word production (Saur et al., 2008; Tourville et al., 2008; Eickhoff et al., 2009; Heim et al., 2009a; van de Ven et al., 2009).”

“A more recent study by Binder et al. (2005) suggests that this area and the adjacent angular gyrus can even be deactivated for pseudoword reading compared to a fixation condition.”

“…for overt speaking during experiments (for fMRI see, e.g., de Zubicaray et al., 2001; Grabowski et al., 2006; Christoffels et al., 2007b; Heim et al., 2009b; Hocking et al., 2009; for EEG see, e.g., Christoffels et al., 2007a; Koester and Schiller, 2008; Costa et al., 2009; Strijkers et al., 2010).”

“…was the only remaining core area that was not more often reported in experiments using overt responses (see Murphy et al., 1997; Wise et al., 1999; Huang et al., 2001; and more recently Ackermann and Riecker, 2004, for the absence of Broca’s area activation in direct comparisons of overt and…”

“…studies (e.g., Sörös et al., 2003; Hultén et al., 2009) and the use of intracranial electrophysiology in neurosurgical patients (Sahin et al., 2009; Edwards et al., 2010) the main development has been the use of transcranial magnetic stimulation (TMS) as a tool for temporarily stimulating or…”

“…imaging (fMRI) and in electroencephalography (EEG) methods have been developed that allow for overt speaking during experiments (for fMRI see, e.g., de Zubicaray et al., 2001; Grabowski et al., 2006; Christoffels et al., 2007b; Heim et al., 2009b; Hocking et al., 2009; for EEG see, e.g.,…”

“Shuster and Lemieux (2005) compared the overt production of multisyllabic words to the production of monosyllabic words.”

“…information taken from Rodriguez-Fornells et al. (2002), Guo et al. (2005), and Hanulová et al. (2011) were approximately 170, 140, and 90 ms. Schmitt et al. (2000) reported a N200 peak difference of 90 ms. Adding the estimate of 200 ms for conceptual preparation, these numbers suggest an…”

“Acheson et al. (2011) found a significant facilitating TMS effect on response latencies when the left mid MTG but not when the left posterior STG was stimulated between 100 ms before and 200 ms after picture onset.”

“Lexical selection in word production was also targeted in an MEG study on picture naming (Maess et al., 2002) by use of the semantic category interference paradigm.”

“To date only one study (Papoutsi et al., 2009) used an experimental variable (biphone frequency) that convincingly disentangles syllabification from later phonetic and motor processing stages and confirmed a pattern predicted for syllabification in a subregion of Broca’s area.”

“By contrast Shafto et al. (2007) found age-related word finding problems to be correlated with gray matter atrophy in the left insula but not the posterior temporal lobe.”

“A third recent development in fMRI research on word production is the use of repetition suppression or adaptation paradigms (e.g., Graves et al., 2008; Peeva et al., 2010).”

“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.”

“Graves et al. (2007) manipulated three variables (lexical frequency, object familiarity, and word length) to study effects at lexical phonological, semantic, and articulatory processing stages.”

“Carreiras et al. (2006) also favor a role in articulatory planning based on their finding of a syllable frequency effect in the left anterior insula.”

“Peeva et al. (2010) found fMRI adaptation for repeated syllables in the left ventral premotor cortex, suggesting syllablelevel representations in this region.”

“In a complex design combining semantic blocking and picture-word interference, Aristei et al. (2011) found increases in naming latencies of category coordinates for both manipulations.”

“To find out which of these regions are in fact necessary for word production (rather than just somehow involved), Indefrey (2007) compared them with brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann…”

“…for overt speaking during experiments (for fMRI see, e.g., de Zubicaray et al., 2001; Grabowski et al., 2006; Christoffels et al., 2007b; Heim et al., 2009b; Hocking et al., 2009; for EEG see, e.g., Christoffels et al., 2007a; Koester and Schiller, 2008; Costa et al., 2009; Strijkers et al., 2010).”

“The study of Peeva et al. (2010) is a nice example of this approach.”

“…the interaction between brain areas (structural equation modeling, SEM; dynamic causal modeling, DCM, independent component analysis, ICA) have begun to be applied to word production (Saur et al., 2008; Tourville et al., 2008; Eickhoff et al., 2009; Heim et al., 2009a; van de Ven et al., 2009).”

“In a study by Hanulová et al. (2011) N200 nogo responses were contingent on a man-made/natural decision, showing an N200 onset latency of around 300 ms. Note that the latency of N200 responses includes the time needed for the decision to withhold the button press, so that the information on which…”

“In patients with temporal lobe epilepsy, Trebuchon-Da Fonseca et al. (2009) found a relationship between TOT states and reduced resting-state metabolism (measured with 18-fluoro-2-desoxy-dglucose-PET) in the left inferior parietal lobe and the posterior superior and inferior temporal cortex.”

“…interference effect (Levelt et al., 1991; Roelofs, 1992), suggesting a lemma retrieval duration of 100–150 ms and electrophysiological data by Schmitt et al. (2001) suggesting that grammatical gender information is available about 75 ms later than conceptual information about the physical…”

“This study, thus, provides much stronger support than previous studies for an involvement of anterior to mid temporal regions in lemma selection, which had been called into question by some authors (e.g., Wilson et al., 2009).”

“By contrast, Ackermann and Riecker (2004) and Riecker et al. (2000) directly compared overt and covert responses and found insular activation only for overt responses.”

“Two more recent studies (Rodriguez-Fornells et al., 2002; Guo et al., 2005) also used paradigms in which go/nogo responses were contingent on an animal/object decision.”

“In a more recent study, de Zubicaray et al., 2006, see, however, Abel et al., 2009, for a negative finding in left mid MTG in a similar contrast) ruled out this possible concern by showing increased mid STG and posterior STG activation for the naming of target words that were preceded by the…”

“…the bilateral STG in the external loop of self-monitoring based on studies showing additional bilateral superior temporal activations by distorting the subjects’ feedback of their own voice or presenting the subjects with alien feedback while they spoke (McGuire et al., 1996; Hirano et al., 1997).”

“Including more recent MEG and TMS studies there are now five studies providing temporal data on left IFG activation in picture naming (Salmelin et al., 1994; Sörös et al., 2003; Vihla et al., 2006; Hultén et al., 2009; Schuhmann et al., 2009) summarized in Table 3.”

“In an fMRI study using the picture-word interference paradigm, de Zubicaray et al. (2002) targeted lexical word form retrieval by using distractor words that were phonologically related to the picture names.”

“The SMA and/or the anterior cingulum also seem to be involved in internal speech monitoring (Möller et al., 2007).”

“Including more recent MEG and TMS studies there are now five studies providing temporal data on left IFG activation in picture naming (Salmelin et al., 1994; Sörös et al., 2003; Vihla et al., 2006; Hultén et al., 2009; Schuhmann et al., 2009) summarized in Table 3.”

“However, as shown by Abdel Rahman and Sommer (2003), the relatively late availability of more peripheral conceptual information, such as the kind of food an animal prefers, does not delay lemma and word form retrieval, suggesting that conceptual processing continues to run in parallel with subsequent processing stages.”

“In an fMRI study using the picture-word interference paradigm, de Zubicaray et al. (2002) targeted lexical word form retrieval by using distractor words that were phonologically related to the picture names.”

“In a more recent study, de Zubicaray et al., 2006, see, however, Abel et al., 2009, for a negative finding in left mid MTG in a similar contrast) ruled out this possible concern by showing increased mid STG and posterior STG activation for the naming of target words that were preceded by the…”

“Using a similar paradigm in an arterial spin labeling fMRI study with overt naming, Hocking et al. (2009) observed hippocampal and left mid to posterior STG activation but no mid MTG activation in the semantic blocking condition (see Frontiers in Psychology | Language Sciences October 2011 | Volume…”

“To find out which of these regions are in fact necessary for word production (rather than just somehow involved), Indefrey (2007) compared them with brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann…”

“To find out which of these regions are in fact necessary for word production (rather than just somehow involved), Indefrey (2007) compared them with brain areas in which transient lesions induced by electrocortical or TMS stimulation reliably interfered with picture naming across seven studies (Ojemann, 1983; Ojemann…”

“, 2010) the main development has been the use of transcranial magnetic stimulation (TMS) as a tool for temporarily stimulating or interfering with neuronal activity in specific brain regions at a specific time (Schuhmann et al., 2009; Acheson et al., 2011).”

“Including more recent MEG and TMS studies there are now five studies providing temporal data on left IFG activation in picture naming (Salmelin et al., 1994; Sörös et al., 2003; Vihla et al., 2006; Hultén et al., 2009; Schuhmann et al., 2009) summarized in Table 3.”

“…the interaction between brain areas (structural equation modeling, SEM; dynamic causal modeling, DCM, independent component analysis, ICA) have begun to be applied to word production (Saur et al., 2008; Tourville et al., 2008; Eickhoff et al., 2009; Heim et al., 2009a; van de Ven et al., 2009).”

“Indefrey and Levelt (2000, 2004, see also Indefrey, 2007) conducted comprehensive meta-analyses of word production studies that had used the mapping approach. They first analyzed the tasks with respect to lead-in processes preceding word production and core word production processes as assumed by psycholinguistic mode…”

“Considering that this activation was sensitive to word-frequency and that I&L suggested a time window for lemma selection between 175 and 250 ms, the authors (and also Hagoort and Levelt, 2009) interpreted this result as indicating a role of Broca’s area in lexical access in word production.”

“Results from an ERP study comparing the naming of pictures with high and low name agreement (Cheng et al., 2010) also show an early effect (100–150 ms) probably due to object recognition difficulty being one source of low name agreement.”

“(2002), Guo et al. (2005), and Hanulová et al. (2011) also provided N200 onset times for first phoneme decisions around 460, 400, and 390 ms.”

“Three other studies (Christoffels et al., 2007a; Laganaro et al., 2009b; Strijkers et al., 2010) investigated the time course of the electrophysiological effects of lexical frequency and cognate status (the target language name of the depicted object sounds/doesn’t sound similar in another language…”

“However, as shown by Abdel Rahman and Sommer (2003), the relatively late availability of more peripheral conceptual information, such as the kind of food an animal prefers, does not delay lemma and word form retrieval, suggesting that conceptual processing continues to run in parallel with subsequent processing stages.”

“…the interaction between brain areas (structural equation modeling, SEM; dynamic causal modeling, DCM, independent component analysis, ICA) have begun to be applied to word production (Saur et al., 2008; Tourville et al., 2008; Eickhoff et al., 2009; Heim et al., 2009a; van de Ven et al., 2009).”

“In this study, they, furthermore, showed additional activation of the anterior cingulate (see also Hirschfeld et al., 2008), and inferior prefrontal regions suggesting the involvement of top-down control processes in the naming of target words preceded by distractors.”

“…October 2011 | Volume 2 | Article 255 | 11 area may thus serve as a common store of lexical word form representations for word production and perception (see also Hocking and Price, 2009, for a lexical phonological effect on left posterior STG activation in comprehension).”

“…2011 | Volume 2 | Article 255 | 3 Table 1 | Continued Study Manipulation Onset/time window of effect (ms) Voice onset time (ms) Processing stage Schiller et al. (2003), ERP: N200 peak latencies Metrical (stress on first or second syllable) vs. syllabification (consonant in first or second…”

“More recently Schiller (2006) measured an N200 peak difference of 58 ms between a lexical stress decision on the first and the second syllable of bisyllabic words, confirming earlier data by Wheeldon and Levelt (1995) and van Turennout et al. (1997) that suggested a phonological encoding duration…”

“Three other studies (Christoffels et al., 2007a; Laganaro et al., 2009b; Strijkers et al., 2010) investigated the time course of the electrophysiological effects of lexical frequency and cognate status (the target language name of the depicted object sounds/doesn’t sound similar in another language…”

“Bles and Jansma (2008) manipulated the phonological relatedness of unattended distractor pictures and also found activation decreases in the left posterior STG when the distractor pictures were phonologically related.”

“Considering that this activation was sensitive to word-frequency and that I&L suggested a time window for lemma selection between 175 and 250 ms, the authors (and also Hagoort and Levelt, 2009) interpreted this result as indicating a role of Broca’s area in lexical access in word production. Note, however, that the ta…”

“A good example in this respect is an eye-tracking study by Huettig and Hartsuiker (2008) who measured very long naming latencies of more than 2000 ms.”

“…et al. (2002), Guo et al. (2005), and Hanulová et al. (2011) also provided N200 onset times for first phoneme decisions around 460, 400, and 390 ms. Zhang and Damian (2009b) report an N200 onset around 290 ms. Camen et al. (2010) using temporal segmentation analysis observed much earlier ERP…”

“Yagishita et al. (2008) asked their subjects to name famous faces during fMRI scanning.”

“Camen et al. (2010) using temporal segmentation analysis observed much earlier ERP effects related to a decision on the first phoneme (210–290 ms). The median estimate of the five studies is 390 ms. N200 peak differences between the availability of semantic and first phoneme information taken from Rodriguez-Fornells e…”

“Evidence for the retrieval of word form information that does not rely on phoneme monitoring comes from a study by Morgan et al. (2008).”

“…(EEG) methods have been developed that allow for overt speaking during experiments (for fMRI see, e.g., de Zubicaray et al., 2001; Grabowski et al., 2006; Christoffels et al., 2007b; Heim et al., 2009b; Hocking et al., 2009; for EEG see, e.g., Christoffels et al., 2007a; Koester…”

“…the bilateral STG in the external loop of self-monitoring based on studies showing additional bilateral superior temporal activations by distorting the subjects’ feedback of their own voice or presenting the subjects with alien feedback while they spoke (McGuire et al., 1996; Hirano et al., 1997).”

“…et al. (2002), Guo et al. (2005), and Hanulová et al. (2011) also provided N200 onset times for first phoneme decisions around 460, 400, and 390 ms. Zhang and Damian (2009b) report an N200 onset around 290 ms. Camen et al. (2010) using temporal segmentation analysis observed much earlier ERP…”

“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…”