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.”

“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.”

“…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).”

“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.”

“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.”

“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.”

“…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).”

“…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).”

“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.”

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

“…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…”

“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).”

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

“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…”

“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.”

“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).”

“…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).”

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

“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).”

“…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…”

“…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…”

“…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…”

“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.”

“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.”

“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…”

“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.”

“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).”

“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.”

“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…”

“…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).”

“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).”