Consistency influences altered auditory feedback processing.

Previous research on the effect of perturbed auditory feedback in speech production has focused on two types of responses. In the short term, speakers generate compensatory motor commands in response to unexpected perturbations. In the longer term, speakers adapt feedforward motor programmes in response to feedback perturbations, to avoid future errors. The current study investigated the relation between these two types of responses to altered auditory feedback. Specifically, it was hypothesised that consistency in previous feedback perturbations would influence whether speakers adapt their feedforward motor programmes. In an altered auditory feedback paradigm, formant perturbations were applied either across all trials (the consistent condition) or only to some trials, whereas the others remained unperturbed (the inconsistent condition). The results showed that speakers' responses were affected by feedback consistency, with stronger speech changes in the consistent condition compared with the inconsistent condition. Current models of speech-motor control can explain this consistency effect. However, the data also suggest that compensation and adaptation are distinct processes, which are not in line with all current models.

Self-monitoring in the cerebral cortex: Neural responses to small pitch shifts in auditory feedback during speech production.

Speaking is a complex motor skill which requires near instantaneous integration of sensory and motor-related information. Current theory hypothesizes a complex interplay between motor and auditory processes during speech production, involving the online comparison of the speech output with an internally generated forward model. To examine the neural correlates of this intricate interplay between sensory and motor processes, the current study uses altered auditory feedback (AAF) in combination with magnetoencephalography (MEG). Participants vocalized the vowel/e/and heard auditory feedback that was temporarily pitch-shifted by only 25 cents, while neural activity was recorded with MEG. As a control condition, participants also heard the recordings of the same auditory feedback that they heard in the first half of the experiment, now without vocalizing. The participants were not aware of any perturbation of the auditory feedback. We found auditory cortical areas responded more strongly to the pitch shifts during vocalization. In addition, auditory feedback perturbation resulted in spectral power increases in the θ and lower ß bands, predominantly in sensorimotor areas. These results are in line with current models of speech production, suggesting auditory cortical areas are involved in an active comparison between a forward model's prediction and the actual sensory input. Subsequently, these areas interact with motor areas to generate a motor response. Furthermore, the results suggest that θ and ß power increases support auditory-motor interaction, motor error detection and/or sensory prediction processing.

Opposing and following responses in sensorimotor speech control: Why responses go both ways.

When talking, speakers continuously monitor and use the auditory feedback of their own voice to control and inform speech production processes. When speakers are provided with auditory feedback that is perturbed in real time, most of them compensate for this by opposing the feedback perturbation. But some responses follow the perturbation. In the present study, we investigated whether the state of the speech production system at perturbation onset may determine what type of response (opposing or following) is made. The results suggest that whether a perturbation-related response is opposing or following depends on ongoing fluctuations of the production system: The system initially responds by doing the opposite of what it was doing. This effect and the nontrivial proportion of following responses suggest that current production models are inadequate: They need to account for why responses to unexpected sensory feedback depend on the production system's state at the time of perturbation.

Individual variability as a window on production-perception interactions in speech motor control.

An important part of understanding speech motor control consists of capturing the interaction between speech production and speech perception. This study tests a prediction of theoretical frameworks that have tried to account for these interactions: If speech production targets are specified in auditory terms, individuals with better auditory acuity should have more precise speech targets, evidenced by decreased within-phoneme variability and increased between-phoneme distance. A study was carried out consisting of perception and production tasks in counterbalanced order. Auditory acuity was assessed using an adaptive speech discrimination task, while production variability was determined using a pseudo-word reading task. Analyses of the production data were carried out to quantify average within-phoneme variability, as well as average between-phoneme contrasts. Results show that individuals not only vary in their production and perceptual abilities, but that better discriminators have more distinctive vowel production targets-that is, targets with less within-phoneme variability and greater between-phoneme distances-confirming the initial hypothesis. This association between speech production and perception did not depend on local phoneme density in vowel space. This study suggests that better auditory acuity leads to more precise speech production targets, which may be a consequence of auditory feedback affecting speech production over time.

Cognate Costs in Bilingual Speech Production: Evidence from Language Switching.

This study investigates cross-language lexical competition in the bilingual mental lexicon. It provides evidence for the occurrence of inhibition as well as the commonly reported facilitation during the production of cognates (words with similar phonological form and meaning in two languages) in a mixed picture naming task by highly proficient Welsh-English bilinguals. Previous studies have typically found cognate facilitation. It has previously been proposed (with respect to non-cognates) that cross-language inhibition is limited to low-proficient bilinguals; therefore, we tested highly proficient, early bilinguals. In a mixed naming experiment (i.e., picture naming with language switching), 48 highly proficient, early Welsh-English bilinguals named pictures in Welsh and English, including cognate and non-cognate targets. Participants were English-dominant, Welsh-dominant, or had equal language dominance. The results showed evidence for cognate inhibition in two ways. First, both facilitation and inhibition were found on the cognate trials themselves, compared to non-cognate controls, modulated by the participants' language dominance. The English-dominant group showed cognate inhibition when naming in Welsh (and no difference between cognates and controls when naming in English), and the Welsh-dominant and equal dominance groups generally showed cognate facilitation. Second, cognate inhibition was found as a behavioral adaptation effect, with slower naming for non-cognate filler words in trials after cognates than after non-cognate controls. This effect was consistent across all language dominance groups and both target languages, suggesting that cognate production involved cognitive control even if this was not measurable in the cognate trials themselves. Finally, the results replicated patterns of symmetrical switch costs, as commonly reported for balanced bilinguals. We propose that cognate processing might be affected by two different processes, namely competition at the lexical-semantic level and facilitation at the word form level, and that facilitation at the word form level might (sometimes) outweigh any effects of inhibition at the lemma level. In sum, this study provides evidence that cognate naming can cause costs in addition to benefits. The finding of cognate inhibition, particularly for the highly proficient bilinguals tested, provides strong evidence for the occurrence of lexical competition across languages in the bilingual mental lexicon.

Hierarchy is Detrimental for Human Cooperation.

Studies of animal behavior consistently demonstrate that the social environment impacts cooperation, yet the effect of social dynamics has been largely excluded from studies of human cooperation. Here, we introduce a novel approach inspired by nonhuman primate research to address how social hierarchies impact human cooperation. Participants competed to earn hierarchy positions and then could cooperate with another individual in the hierarchy by investing in a common effort. Cooperation was achieved if the combined investments exceeded a threshold, and the higher ranked individual distributed the spoils unless control was contested by the partner. Compared to a condition lacking hierarchy, cooperation declined in the presence of a hierarchy due to a decrease in investment by lower ranked individuals. Furthermore, hierarchy was detrimental to cooperation regardless of whether it was earned or arbitrary. These findings mirror results from nonhuman primates and demonstrate that hierarchies are detrimental to cooperation. However, these results deviate from nonhuman primate findings by demonstrating that human behavior is responsive to changing hierarchical structures and suggests partnership dynamics that may improve cooperation. This work introduces a controlled way to investigate the social influences on human behavior, and demonstrates the evolutionary continuity of human behavior with other primate species.

Effects of parallel planning on agreement production.

An important issue in current psycholinguistics is how the time course of utterance planning affects the generation of grammatical structures. The current study investigated the influence of parallel activation of the components of complex noun phrases on the generation of subject-verb agreement. Specifically, the lexical interference account (Gillespie & Pearlmutter, 2011b; Solomon & Pearlmutter, 2004) predicts more agreement errors (i.e., attraction) for subject phrases in which the head and local noun mismatch in number (e.g., the apple next to the pears) when nouns are planned in parallel than when they are planned in sequence. We used a speeded picture description task that yielded sentences such as the apple next to the pears is red. The objects mentioned in the noun phrase were either semantically related or unrelated. To induce agreement errors, pictures sometimes mismatched in number. In order to manipulate the likelihood of parallel processing of the objects and to test the hypothesized relationship between parallel processing and the rate of agreement errors, the pictures were either placed close together or far apart. Analyses of the participants' eye movements and speech onset latencies indicated slower processing of the first object and stronger interference from the related (compared to the unrelated) second object in the close than in the far condition. Analyses of the agreement errors yielded an attraction effect, with more errors in mismatching than in matching conditions. However, the magnitude of the attraction effect did not differ across the close and far conditions. Thus, spatial proximity encouraged parallel processing of the pictures, which led to interference of the associated conceptual and/or lexical representation, but, contrary to the prediction, it did not lead to more attraction errors.

Modulations of the auditory M100 in an imitation task.

Models of speech production explain event-related suppression of the auditory cortical response as reflecting a comparison between auditory predictions and feedback. The present MEG study was designed to test two predictions from this framework: (1) whether the reduced auditory response varies as a function of the mismatch between prediction and feedback; (2) whether individual variation in this response is predictive of speech-motor adaptation. Participants alternated between online imitation and listening tasks. In the imitation task, participants began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorily-presented vowels varying in acoustic distance from /e/. Results replicated suppression, with a smaller M100 during speaking than listening. Although we did not find unequivocal support for the first prediction, participants with less M100 suppression were better at the imitation task. These results are consistent with the enhancement of M100 serving as an error signal to drive subsequent speech-motor adaptation.

Electrophysiological evidence that inhibition supports lexical selection in picture naming.

We investigated the neural basis of inhibitory control during lexical selection. Participants overtly named pictures while response times (RTs) and event-related brain potentials (ERPs) were recorded. The difficulty of lexical selection was manipulated by using object and action pictures with high name agreement (few response candidates) versus low name agreement (many response candidates). To assess the involvement of inhibition, we conducted delta plot analyses of naming RTs and examined the N2 component of the ERP. We found longer mean naming RTs and a larger N2 amplitude in the low relative to the high name agreement condition. For action naming we found a negative correlation between the slopes of the slowest delta segment and the difference in N2 amplitude between the low and high name agreement conditions. The converging behavioral and electrophysiological evidence suggests that selective inhibition is engaged to reduce competition during lexical selection in picture naming.

Keeping it simple: studying grammatical encoding with lexically reduced item sets.

Compared to the large body of work on lexical access, little research has been done on grammatical encoding in language production. An exception is the generation of subject-verb agreement. Here, two key findings have been reported: (1) speakers make more agreement errors when the head and local noun of a phrase mismatch in number than when they match [e.g., the key to the cabinet(s)]; and (2) this attraction effect is asymmetric, with stronger attraction for singular than for plural head nouns. Although these findings are robust, the cognitive processes leading to agreement errors and their significance for the generation of correct agreement are not fully understood. We propose that future studies of agreement, and grammatical encoding in general, may benefit from using paradigms that tightly control the variability of the lexical content of the material. We report two experiments illustrating this approach. In both of them, the experimental items featured combinations of four nouns, four color adjectives, and two prepositions. In Experiment 1, native speakers of Dutch described pictures in sentences such as the circle next to the stars is blue. In Experiment 2, they carried out a forced-choice task, where they read subject noun phrases (e.g., the circle next to the stars) and selected the correct verb-phrase (is blue or are blue) with a button press. Both experiments showed an attraction effect, with more errors after subject phrases with mismatching, compared to matching head and local nouns. This effect was stronger for singular than plural heads, replicating the attraction asymmetry. In contrast, the response times recorded in Experiment 2 showed similar attraction effects for singular and plural head nouns. These results demonstrate that critical agreement phenomena can be elicited reliably in lexically reduced contexts. We discuss the theoretical implications of the findings and the potential and limitations of studies using lexically simple materials.

Twisting tongues to test for conflict-monitoring in speech production.

A number of recent studies have hypothesized that monitoring in speech production may occur via domain-general mechanisms responsible for the detection of response conflict. Outside of language, two ERP components have consistently been elicited in conflict-inducing tasks (e.g., the flanker task): the stimulus-locked N2 on correct trials, and the response-locked error-related negativity (ERN). The present investigation used these electrophysiological markers to test whether a common response conflict monitor is responsible for monitoring in speech and non-speech tasks. Electroencephalography (EEG) was recorded while participants performed a tongue twister (TT) task and a manual version of the flanker task. In the TT task, people rapidly read sequences of four nonwords arranged in TT and non-TT patterns three times. In the flanker task, people responded with a left/right button press to a center-facing arrow, and conflict was manipulated by the congruency of the flanking arrows. Behavioral results showed typical effects of both tasks, with increased error rates and slower speech onset times for TT relative to non-TT trials and for incongruent relative to congruent flanker trials. In the flanker task, stimulus-locked EEG analyses replicated previous results, with a larger N2 for incongruent relative to congruent trials, and a response-locked ERN. In the TT task, stimulus-locked analyses revealed broad, frontally-distributed differences beginning around 50 ms and lasting until just before speech initiation, with TT trials more negative than non-TT trials; response-locked analyses revealed an ERN. Correlation across these measures showed some correlations within a task, but little evidence of systematic cross-task correlation. Although the present results do not speak against conflict signals from the production system serving as cues to self-monitoring, they are not consistent with signatures of response conflict being mediated by a single, domain-general conflict monitor.

Attention for speaking: domain-general control from the anterior cingulate cortex in spoken word production.

Accumulating evidence suggests that some degree of attentional control is required to regulate and monitor processes underlying speaking. Although progress has been made in delineating the neural substrates of the core language processes involved in speaking, substrates associated with regulatory and monitoring processes have remained relatively underspecified. We report the results of an fMRI study examining the neural substrates related to performance in three attention-demanding tasks varying in the amount of linguistic processing: vocal picture naming while ignoring distractors (picture-word interference, PWI); vocal color naming while ignoring distractors (Stroop); and manual object discrimination while ignoring spatial position (Simon task). All three tasks had congruent and incongruent stimuli, while PWI and Stroop also had neutral stimuli. Analyses focusing on common activation across tasks identified a portion of the dorsal anterior cingulate cortex (ACC) that was active in incongruent trials for all three tasks, suggesting that this region subserves a domain-general attentional control function. In the language tasks, this area showed increased activity for incongruent relative to congruent stimuli, consistent with the involvement of domain-general mechanisms of attentional control in word production. The two language tasks also showed activity in anterior-superior temporal gyrus (STG). Activity increased for neutral PWI stimuli (picture and word did not share the same semantic category) relative to incongruent (categorically related) and congruent stimuli. This finding is consistent with the involvement of language-specific areas in word production, possibly related to retrieval of lexical-semantic information from memory. The current results thus suggest that in addition to engaging language-specific areas for core linguistic processes, speaking also engages the ACC, a region that is likely implementing domain-general attentional control.

Stimulating the brain's language network: syntactic ambiguity resolution after TMS to the inferior frontal gyrus and middle temporal gyrus.

The posterior middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) are two critical nodes of the brain's language network. Previous neuroimaging evidence has supported a dissociation in language comprehension in which parts of the MTG are involved in the retrieval of lexical syntactic information and the IFG in unification operations that maintain, select, and integrate multiple sources of information over time. In the present investigation, we tested for causal evidence of this dissociation by modulating activity in IFG and MTG using an offline TMS procedure: continuous theta-burst stimulation. Lexical-syntactic retrieval was manipulated by using sentences with and without a temporarily word-class (noun/verb) ambiguity (e.g., run). In one group of participants, TMS was applied to the IFG and MTG, and in a control group, no TMS was applied. Eye movements were recorded and quantified at two critical sentence regions: a temporarily ambiguous region and a disambiguating region. Results show that stimulation of the IFG led to a modulation of the ambiguity effect (ambiguous-unambiguous) at the disambiguating sentence region in three measures: first fixation durations, total reading times, and regressive eye movements into the region. Both IFG and MTG stimulation modulated the ambiguity effect for total reading times in the temporarily ambiguous sentence region relative to the control group. The current results demonstrate that an offline repetitive TMS protocol can have influences at a different point in time during online processing and provide causal evidence for IFG involvement in unification operations during sentence comprehension.

Conflict monitoring in speech production: physiological evidence from bilingual picture naming.

Self-monitoring in production is critical to correct performance, and recent accounts suggest that such monitoring may occur via the detection of response conflict. The error-related negativity (ERN) is a response-locked event-related potential (ERP) that is sensitive to response conflict. The present study examines whether response conflict is detected in production by exploring a situation where multiple outputs are activated: the bilingual naming of form-related equivalents (i.e. cognates). ERPs were recorded while German-Dutch bilinguals named pictures in their first and second languages. Although cognates were named faster than non-cognates, response conflict was evident in the form of a larger ERN-like response for cognates and adaptation effects on naming, as the magnitude of cognate facilitation was smaller following the naming of cognates. Given that signals of response conflict are present during correct naming, the present results suggest that such conflict may serve as a reliable signal for monitoring in speech production.

Areas of left perisylvian cortex mediate auditory-verbal short-term memory.

A contentious issue in memory research is whether verbal short-term memory (STM) depends on a neural system specifically dedicated to the temporary maintenance of information, or instead relies on the same brain areas subserving the comprehension and production of language. In this study, we examined a large sample of adults with acquired brain lesions to identify the critical neural substrates underlying verbal STM and the relationship between verbal STM and language processing abilities. We found that patients with damage to selective regions of left perisylvian cortex - specifically the inferior frontal and posterior temporal sectors - were impaired on auditory-verbal STM performance (digit span), as well as on tests requiring the production and/or comprehension of language. These results support the conclusion that verbal STM and language processing are mediated by the same areas of left perisylvian cortex.

The Rhymes that the Reader Perused Confused the Meaning: Phonological Effects during On-line Sentence Comprehension.

Research on written language comprehension has generally assumed that the phonological properties of a word have little effect on sentence comprehension beyond the processes of word recognition. Two experiments investigated this assumption. Participants silently read relative clauses in which two pairs of words either did or did not have a high degree of phonological overlap. Participants were slower reading and less accurate comprehending the overlap sentences compared to the non-overlapping controls, even though sentences were matched for plausibility and differed by only two words across overlap conditions. A comparison across experiments showed that the overlap effects were larger in the more difficult object relative than in subject relative sentences. The reading patterns showed that phonological representations affect not only memory for recently encountered sentences but also the developing sentence interpretation during on-line processing. Implications for theories of sentence processing and memory are discussed.

Increased Alpha-Band Power during the Retention of Shapes and Shape-Location Associations in Visual Short-Term Memory.

Studies exploring the role of neural oscillations in cognition have revealed sustained increases in alpha-band (~8-14 Hz) power during the delay period of delayed-recognition short-term memory tasks. These increases have been proposed to reflect the inhibition, for example, of cortical areas representing task-irrelevant information, or of potentially interfering representations from previous trials. Another possibility, however, is that elevated delay-period alpha-band power (DPABP) reflects the selection and maintenance of information, rather than, or in addition to, the inhibition of task-irrelevant information. In the present study, we explored these possibilities using a delayed-recognition paradigm in which the presence and task relevance of shape information was systematically manipulated across trial blocks and electroencephalographic was used to measure alpha-band power. In the first trial block, participants remembered locations marked by identical black circles. The second block featured the same instructions, but locations were marked by unique shapes. The third block featured the same stimulus presentation as the second, but with pretrial instructions indicating, on a trial-by-trial basis, whether memory for shape or location was required, the other dimension being irrelevant. In the final block, participants remembered the unique pairing of shape and location for each stimulus. Results revealed minimal DPABP in each of the location-memory conditions, whether locations were marked with identical circles or with unique task-irrelevant shapes. In contrast, alpha-band power increases were observed in both the shape-memory condition, in which location was task irrelevant, and in the critical final condition, in which both shape and location were task relevant. These results provide support for the proposal that alpha-band oscillations reflect the retention of shape information and/or shape-location associations in short-term memory.

A common neural substrate for language production and verbal working memory.

Verbal working memory (VWM), the ability to maintain and manipulate representations of speech sounds over short periods, is held by some influential models to be independent from the systems responsible for language production and comprehension [e.g., Baddeley, A. D. Working memory, thought, and action. New York, NY: Oxford University Press, 2007]. We explore the alternative hypothesis that maintenance in VWM is subserved by temporary activation of the language production system [Acheson, D. J., & MacDonald, M. C. Verbal working memory and language production: Common approaches to the serial ordering of verbal information. Psychological Bulletin, 135, 50-68, 2009b]. Specifically, we hypothesized that for stimuli lacking a semantic representation (e.g., nonwords such as mun), maintenance in VWM can be achieved by cycling information back and forth between the stages of phonological encoding and articulatory planning. First, fMRI was used to identify regions associated with two different stages of language production planning: the posterior superior temporal gyrus (pSTG) for phonological encoding (critical for VWM of nonwords) and the middle temporal gyrus (MTG) for lexical-semantic retrieval (not critical for VWM of nonwords). Next, in the same subjects, these regions were targeted with repetitive transcranial magnetic stimulation (rTMS) during language production and VWM task performance. Results showed that rTMS to the pSTG, but not the MTG, increased error rates on paced reading (a language production task) and on delayed serial recall of nonwords (a test of VWM). Performance on a lexical-semantic retrieval task (picture naming), in contrast, was significantly sensitive to rTMS of the MTG. Because rTMS was guided by language production-related activity, these results provide the first causal evidence that maintenance in VWM directly depends on the long-term representations and processes used in speech production.

The effect of concurrent semantic categorization on delayed serial recall.

The influence of semantic processing on the serial ordering of items in short-term memory was explored using a novel dual-task paradigm. Participants engaged in 2 picture-judgment tasks while simultaneously performing delayed serial recall. List material varied in the presence of phonological overlap (Experiments 1 and 2) and in semantic content (concrete words in Experiment 1 and 3; nonwords in Experiments 2 and 3). Picture judgments varied in the extent to which they required accessing visual semantic information (i.e., semantic categorization and line orientation judgments). Results showed that, relative to line-orientation judgments, engaging in semantic categorization judgments increased the proportion of item-ordering errors for concrete lists but did not affect error proportions for nonword lists. Furthermore, although more ordering errors were observed for phonologically similar relative to dissimilar lists, no interactions were observed between the phonological overlap and picture-judgment task manipulations. These results demonstrate that lexical-semantic representations can affect the serial ordering of items in short-term memory. Furthermore, the dual-task paradigm provides a new method for examining when and how semantic representations affect memory performance.