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Does "the motor system" play "a role" in speech perception? If so, where, how, and when? We conducted a systematic review that addresses these questions using both qualitative and quantitative methods. The qualitative review of behavioural, computational modelling, non-human animal, brain damage/disorder, electrical stimulation/recording, and neuroimaging research suggests that distributed brain regions involved in producing speech play specific, dynamic, and contextually determined roles in speech perception. The quantitative review employed region and network based neuroimaging meta-analyses and a novel text mining method to describe relative contributions of nodes in distributed brain networks. Supporting the qualitative review, results show a specific functional correspondence between regions involved in non-linguistic movement of the articulators, covertly and overtly producing speech, and the perception of both nonword and word sounds. This distributed set of cortical and subcortical speech production regions are ubiquitously active and form multiple networks whose topologies dynamically change with listening context. Results are inconsistent with motor and acoustic only models of speech perception and classical and contemporary dual-stream models of the organization of language and the brain. Instead, results are more consistent with complex network models in which multiple speech production related networks and subnetworks dynamically self-organize to constrain interpretation of indeterminant acoustic patterns as listening context requires. [ABSTRACT FROM AUTHOR]

Abstract: A critical assumption underlying the practice of functional localization is that the voxels identified by functional localization are essentially the same as those activated in the main experiment for a particular anatomical area. Violations of this assumption bias the resulting analyses and can dramatically increase the likelihood of both Type I and Type II errors. Here we investigated how the amount of data affects the reliability of a set of common functionally-defined regions-of-interest (fROIs). Four participants were scanned ten times each to functionally localize extrastriate regions sensitive to visually presented words, objects and faces. A within-subject random-effects analysis was used as the “gold standard” for identifying the fROIs and the results were compared to within-subject, fixed-effect analyses typically used for functional localization. By varying the quantity of data included in the analyses, we empirically assessed the amount needed to ensure reliable identification of the fROIs. The results demonstrated that the most consistent fROIs were based on either stringent statistical thresholding (Z>5.0) of large quantities of data or on lenient thresholding (Z>2.3) of a modest amount of data, with both methods yielding 70–80% overlap between the functional localization results and the “gold standard.” Stringent statistical thresholds on typical quantities of localizer data led to the poorest reliability (<20% overlap). These findings suggest that the most reliable and cost-efficient method for functional localization involves collecting a relatively small amount of data (~10min) and using a lenient statistical threshold to identify all voxels in a given region that are sensitive to the process-of-interest. [Copyright &y& Elsevier]

Abstract: The left ventral premotor cortex (PMv) is preferentially activated by exemplars of tools, suggestive of category specificity in this region. Here we used state-dependent transcranial magnetic stimulation (TMS) to investigate the causal role of such category-specific neuronal representations in the encoding of tool words. Priming to a category name (either “Tool” or “Animal”) was used with the objective of modulating the initial activation state of this region prior to application of TMS and the presentation of the target stimulus. When the target word was an exemplar of the “Tool” category, the effects of TMS applied over PMv (but not PMd) interacted with priming history by facilitating reaction times on incongruent trials while not affecting congruent trials. This congruency/TMS interaction implies that the “Tool” and “Animal” primes had a differential effect on the initial activation state of the left PMv and implies that this region is one neural locus of category-specific behavioral priming for the “Tool” category. TMS applied over PMv had no behavioral effect when the target stimulus was an exemplar of the “Animal” category, regardless of whether the target word was congruent or incongruent with the prime. That TMS applied over the left PMv interacted with a priming effect that extended from the category name (“Tool”) to exemplars of that category suggests that this region contains neuronal representation associated with a specific semantic category. Our results also demonstrate that the state-dependent effects obtained in the combination of visual priming and TMS are useful in the study of higher-level cognitive functions. [Copyright &y& Elsevier]

Abstract: Listeners show remarkable flexibility in processing variation in speech signal. One striking example is the ease with which they adapt to novel speech distortions such as listening to someone with a foreign accent. Behavioural studies suggest that significant improvements in comprehension occur rapidly — often within 10–20 sentences. In the present experiment, we investigate the neural changes underlying on-line adaptation to distorted speech using time-compressed speech. Listeners performed a sentence verification task on normal-speed and time-compressed sentences while their neural responses were recorded using fMRI. The results showed that rapid learning of the time-compressed speech occurred during presentation of the first block of 16 sentences and was associated with increased activation in left and right auditory association cortices and in left ventral premotor cortex. These findings suggest that the ability to adapt to a distorted speech signal may, in part, rely on mapping novel acoustic patterns onto existing articulatory motor plans, consistent with the idea that speech perception involves integrating multi-modal information including auditory and motoric cues. [Copyright &y& Elsevier]

Abstract: Along with meaning and form, words can be described on the basis of their grammatical properties. Grammatical gender is often used to investigate the latter as it is a grammatical property that is independent of meaning. The left inferior frontal gyrus (IFG) has been implicated in the encoding of grammatical gender, but its causal role in this process in neurologically normal observers has not been demonstrated. Here we combined verbal satiation with transcranial magnetic stimulation (TMS) to demonstrate that subpopulations of neurons within Broca''s area respond preferentially to different classes of grammatical gender. Subjects were asked to classify Italian nouns into living and nonliving categories; half of these words were of masculine and the other half of feminine grammatical gender. Prior to each test block, a satiation paradigm (a phenomenon in which verbal repetition of a category name leads to a reduced access to that category) was used to modulate the initial state of the representations of either masculine or feminine noun categories. In the No TMS condition, subjects were slower in responding to exemplars to the satiated category relative to exemplars of the nonsatiated category, implying that the neural representations for different classes of grammatical gender are partly dissociable. The application of TMS over Broca''s area removed the behavioral impact of verbal (grammatical) satiation, demonstrating the causal role of this region in the encoding of grammatical gender. These results show that the neural representations for different cases of a grammatical property within Broca''s area are dissociable. [Copyright &y& Elsevier]

Abstract: Functional neuroimaging is fundamentally a tool for mapping function to structure, and its success consequently requires neuroanatomical precision and accuracy. Here we review the various means by which functional activation can be localised to neuroanatomy and suggest that the gold standard should be localisation to the individual’s or group’s own anatomy through the use of neuroanatomical knowledge and atlases of neuroanatomy. While automated means of localisation may be useful, they cannot provide the necessary accuracy, given variability between individuals. We also suggest that the field of functional neuroimaging needs to converge on a common set of methods for reporting functional localisation including a common “standard” space and criteria for what constitutes sufficient evidence to report activation in terms of Brodmann’s areas. [Copyright &y& Elsevier]

Abstract: We investigated the ability to orient attention to a complex, non-perceptual attribute of stimuli—semantic category. Behavioral consequences and neural correlates of semantic orienting were revealed and compared with those of spatial orienting, using event-related functional magnetic-resonance imaging. Semantic orienting significantly shortened response times to identify word stimuli, showing that it is possible to focus attention on non-perceptual attributes of stimuli to enhance behavioral performance. Semantic-orienting cues engaged parietal and frontal areas that were also involved in spatial orienting, but in addition engaged brain areas associated with semantic analysis of words, such as the left anterior inferior frontal cortex. These findings show that attentional orienting selectively engages brain areas with functional specialization for the predicted attributes. They also support the existence of a core frontoparietal network, which controls attentional orienting in speeded response tasks independently of the type of expectations, interacting with task-relevant functionally specialized areas to optimize perception and action. [Copyright &y& Elsevier]

Recent functional imaging studies have referred to a posterior region of the left midfusiform gyrus as the “visual word form area” (VWFA). We review the evidence for this claim and argue that neither the neuropsychological nor neuroimaging data are consistent with a cortical region specialized for visual word form representations. Specifically, there are no reported cases of pure alexia who have deficits limited to visual word form processing and damage limited to the left midfusiform. In addition, we present functional imaging data to demonstrate that the so-called VWFA is activated by normal subjects during tasks that do not engage visual word form processing such as naming colors, naming pictures, reading Braille, repeating auditory words, and making manual action responses to pictures of meaningless objects. If the midfusiform region has a single function that underlies all these tasks, then it does not correspond to visual word form processing. On the other hand, if the region participates in several functions as defined by its interactions with other cortical areas, then identifying the neural system sustaining visual word form representations requires identification of the set of regions involved. We conclude that there is no evidence that visual word form representations are subtended by a single patch of neuronal cortex and it is misleading to label the left midfusiform region as the visual word form area. [Copyright &y& Elsevier]

Primary motor (M1) areas for speech production activate during speechperception. It has been suggested that such activation may be dependent upon modulatory inputs from premotor cortex (PMv). If and how PMv differentially modulates M1 activity during perception of speech that is easy or challenging to understand, however, is unclear. This study aimed to test the link between PMv and M1 during challenging speech perception in two experiments. The first experiment investigated intra-hemispheric connectivity between left hemisphere PMv and left M1 lip area during comprehension of speech under clear and distorted listening conditions. Continuous theta burst stimulation (cTBS) was applied to left PMv in eighteen participants (aged 18-35). Post-cTBS, participants performed a sentence verification task on distorted (imprecisely articulated), and clear speech, whilst also undergoing stimulation of the lip representation in the left M1 to elicit motor evoked potentials (MEPs). In a second, separate experiment, we investigated the role of inter-hemispheric connectivity between right hemisphere PMv and left hemisphere M1 lip area. Dual-coil transcranial magnetic stimulation was applied to right PMv and left M1 lip in fifteen participants (aged 18-35). Results indicated that disruption of PMv during speech perception affects comprehension of distorted speech specifically. Furthermore, our data suggest that listening to distorted speech modulates the balance of intra- and inter-hemispheric interactions, with a larger sensorimotor network implicated during comprehension of distorted speech than when speech perception is optimal. The present results further understanding of PMv-M1 interactions during auditory-motor integration. [ABSTRACT FROM AUTHOR]

Abstract: Orienting spatial attention to locations in the extrapersonal world has been intensively investigated during the past decades. Recently, it was demonstrated that it is also possible to shift attention to locations within mental representations held in working memory. This is an important issue, since the allocation of our attention is not only guided by external stimuli, but also by their internal representations and the expectations we build upon them. The present experiment used behavioural measures and event-related functional magnetic resonance imaging to investigate whether spatial orienting to mental representations can modulate the search and retrieval of information from working memory, and to identify the neural systems involved, respectively. Participants viewed an array of coloured crosses. Seconds after its disappearance, they were cued to locations in the array with valid or neutral cues. Subsequently, they decided whether a probe stimulus was presented in the array. The behavioural results indicated that orienting of spatial attention within working memory attenuates the well-known effect of decreasing performance when memory load is increased. So “internal” spatial orienting seems to highlight information or facilitate search within working memory, which leads to advantages in retrieval. Imaging enabled the separation of brain areas supporting spatial orienting functions from those sensitive to working-memory load. Orienting of spatial attention to the contents of working memory activated posterior parietal cortex bilaterally, the insula, and lateral and medial frontal cortices. [Copyright &y& Elsevier]

It has become increasingly evident that human motor circuits are active during speech perception. However, the conditions under which the motor system modulates speech perception are not clear. Two prominent accounts make distinct predictions for how listening to speech engages speech motor representations. The first account suggests that the motor system is most strongly activated when observing familiar actions (Pickering and Garrod, 2013). Conversely, Wilson and Knoblich's account asserts that motor excitability is greatest when observing less familiar, ambiguous actions (Wilson and Knoblich, 2005). We investigated these predictions using transcranial magnetic stimulation (TMS). Stimulation of the lip and hand representations in the left primary motor cortex elicited motor evoked potentials (MEPs) indexing the excitability of the underlying motor representation. MEPs for lip, but not for hand, were larger during perception of distorted speech produced using a tongue depressor, relative to naturally produced speech. Additional somatotopic facilitation yielded significantly larger MEPs during perception of lip-articulated distorted speech sounds relative to distorted tongue-articulated sounds. Critically, there was a positive correlation between MEP size and the perception of distorted speech sounds. These findings were consistent with predictions made by Wilson & Knoblich (Wilson and Knoblich, 2005), and provide direct evidence of increased motor excitability when speech perception is difficult. [ABSTRACT FROM AUTHOR]

Abstract: In Japanese, the same word can be written in either morphographic Kanji or syllabographic Hiragana and this provides a unique opportunity to disentangle a word’s lexical frequency from the frequency of its visual form – an important distinction for understanding the neural information processing in regions engaged by reading. Behaviorally, participants responded more quickly to high than low frequency words and to visually familiar relative to less familiar words, independent of script. Critically, the imaging results showed that visual familiarity, as opposed to lexical frequency, had a strong effect on activation in ventral occipito-temporal cortex. Activation here was also greater for Kanji than Hiragana words and this was not due to their inherent differences in visual complexity. These findings can be understood within a predictive coding framework in which vOT receives bottom-up information encoding complex visual forms and top-down predictions from regions encoding non-visual attributes of the stimulus. [Copyright &y& Elsevier]

Abstract: Although interactivity is considered a fundamental principle of cognitive (and computational) models of reading, it has received far less attention in neural models of reading that instead focus on serial stages of feed-forward processing from visual input to orthographic processing to accessing the corresponding phonological and semantic information. In particular, the left ventral occipito-temporal (vOT) cortex is proposed to be the first stage where visual word recognition occurs prior to accessing nonvisual information such as semantics and phonology. We used functional magnetic resonance imaging (fMRI) to investigate whether there is evidence that activation in vOT is influenced top-down by the interaction of visual and nonvisual properties of the stimuli during visual word recognition tasks. Participants performed two different types of lexical decision tasks that focused on either visual or nonvisual properties of the word or word-like stimuli. The design allowed us to investigate how vOT activation during visual word recognition was influenced by a task change to the same stimuli and by a stimulus change during the same task. We found both stimulus- and task-driven modulation of vOT activation that can only be explained by top-down processing of nonvisual aspects of the task and stimuli. Our results are consistent with the hypothesis that vOT acts as an interface linking visual form with nonvisual processing in both bottom up and top down directions. Such interactive processing at the neural level is in agreement with cognitive and computational models of reading but challenges some of the assumptions made by current neuro-anatomical models of reading. [Copyright &y& Elsevier]

Abstract: A critical assumption underlying the use of functional localiser scans is that the voxels identified as the functional region-of-interest (fROI) are essentially the same as those activated by the main experimental manipulation. Intra-subject variability in the location of the fROI violates this assumption, reducing the sensitivity of the analysis and biasing the results. Here we investigated consistency and variability in fROIs in a set of 45 volunteers. They performed two functional localiser scans to identify word- and object-sensitive regions of ventral and lateral occipito-temporal cortex, respectively. In the main analyses, fROIs were defined as the category-selective voxels in each region and consistency was measured as the spatial overlap between scans. Consistency was greatest when minimally selective thresholds were used to define “active” voxels (p <0.05 uncorrected), revealing that approximately 65% of the voxels were commonly activated by both scans. In contrast, highly selective thresholds (p <10−4 to 10−6) yielded the lowest consistency values with less than 25% overlap of the voxels active in both scans. In other words, intra-subject variability was surprisingly high, with between one third and three quarters of the voxels in a given fROI not corresponding to those activated in the main task. This level of variability stands in striking contrast to the consistency seen in retinotopically-defined areas and has important implications for designing robust but efficient functional localiser scans. [Copyright &y& Elsevier]