A neuroanatomical and cognitive model of impaired social behaviour in frontotemporal dementia.

Impaired social cognition is a core deficit in frontotemporal dementia (FTD). It is most commonly associated with the behavioural-variant of FTD, with atrophy of the orbitofrontal and ventromedial prefrontal cortex. Social cognitive changes are also common in semantic dementia, with atrophy centred on the anterior temporal lobes. The impairment of social behaviour in FTD has typically been attributed to damage to the orbitofrontal cortex and/or temporal poles and/or the uncinate fasciculus that connects them. However, the relative contributions of each region are unresolved. In this review, we present a unified neurocognitive model of controlled social behaviour that not only explains the observed impairment of social behaviours in FTD, but also assimilates both consistent and potentially contradictory findings from other patient groups, comparative neurology and normative cognitive neuroscience. We propose that impaired social behaviour results from damage to two cognitively- and anatomically-distinct components. The first component is social-semantic knowledge, a part of the general semantic-conceptual system supported by the anterior temporal lobes bilaterally. The second component is social control, supported by the orbitofrontal cortex, medial frontal cortex and ventrolateral frontal cortex, which interacts with social-semantic knowledge to guide and shape social behaviour.

Graded functional organization in the left inferior frontal gyrus: evidence from task-free and task-based functional connectivity.

The left inferior frontal gyrus has been ascribed key roles in numerous cognitive domains, such as language and executive function. However, its functional organization is unclear. Possibilities include a singular domain-general function, or multiple functions that can be mapped onto distinct subregions. Furthermore, spatial transition in function may be either abrupt or graded. The present study explored the topographical organization of the left inferior frontal gyrus using a bimodal data-driven approach. We extracted functional connectivity gradients from (i) resting-state fMRI time-series and (ii) coactivation patterns derived meta-analytically from heterogenous sets of task data. We then sought to characterize the functional connectivity differences underpinning these gradients with seed-based resting-state functional connectivity, meta-analytic coactivation modeling and functional decoding analyses. Both analytic approaches converged on graded functional connectivity changes along 2 main organizational axes. An anterior-posterior gradient shifted from being preferentially associated with high-level control networks (anterior functional connectivity) to being more tightly coupled with perceptually driven networks (posterior). A second dorsal-ventral axis was characterized by higher connectivity with domain-general control networks on one hand (dorsal functional connectivity), and with the semantic network, on the other (ventral). These results provide novel insights into an overarching graded functional organization of the functional connectivity that explains its role in multiple cognitive domains.

Quantifying social semantics: An inclusive definition of socialness and ratings for 8388 English words.

It has been proposed that social experience plays an important role in the grounding of concepts, and socialness has been proffered as a fundamental organisational principle underpinning semantic representation in the human brain. However, the empirical support for these hypotheses is limited by inconsistencies in the way socialness has been defined and measured. To further advance theory, the field must establish a clearer working definition, and research efforts could be facilitated by the availability of an extensive set of socialness ratings for individual concepts. Therefore, in the current work, we employed a novel and inclusive definition to test the extent to which socialness is reliably perceived as a broad construct, and we report socialness norms for over 8000 English words, including nouns, verbs, and adjectives. Our inclusive socialness measure shows good reliability and validity, and our analyses suggest that the socialness ratings capture aspects of word meaning which are distinct to those measured by other pertinent semantic constructs, including concreteness and emotional valence. Finally, in a series of regression analyses, we show for the first time that the socialness of a word's meaning explains unique variance in participant performance on lexical tasks. Our dataset of socialness norms has considerable item overlap with those used in both other lexical/semantic norms and in available behavioural mega-studies. They can help target testable predictions about brain and behaviour derived from multiple representation theories and neurobiological accounts of social semantics.

The role of the ventrolateral anterior temporal lobes in social cognition.

A key challenge for neurobiological models of social cognition is to elucidate whether brain regions are specialised for that domain. In recent years, discussion surrounding the role of anterior temporal regions epitomises such debates; some argue the anterior temporal lobe (ATL) is part of a domain-specific network for social processing, while others claim it comprises a domain-general hub for semantic representation. In the present study, we used ATL-optimised fMRI to map the contribution of different ATL structures to a variety of paradigms frequently used to probe a crucial social ability, namely 'theory of mind' (ToM). Using multiple tasks enables a clearer attribution of activation to ToM as opposed to idiosyncratic features of stimuli. Further, we directly explored whether these same structures are also activated by a non-social task probing semantic representations. We revealed that common to all of the tasks was activation of a key ventrolateral ATL region that is often invisible to standard fMRI. This constitutes novel evidence in support of the view that the ventrolateral ATL contributes to social cognition via a domain-general role in semantic processing and against claims of a specialised social function.

Establishing a role of the semantic control network in social cognitive processing: A meta-analysis of functional neuroimaging studies.

The contribution and neural basis of cognitive control is under-specified in many prominent models of socio-cognitive processing. Important outstanding questions include whether there are multiple, distinguishable systems underpinning control and whether control is ubiquitously or selectively engaged across different social behaviours and task demands. Recently, it has been proposed that the regulation of social behaviours could rely on brain regions specialised in the controlled retrieval of semantic information, namely the anterior inferior frontal gyrus (IFG) and posterior middle temporal gyrus. Accordingly, we investigated for the first time whether the neural activation commonly found in social functional neuroimaging studies extends to these 'semantic control' regions. We conducted five coordinate-based meta-analyses to combine results of 499 fMRI/PET experiments and identified the brain regions consistently involved in semantic control, as well as four social abilities: theory of mind, trait inference, empathy and moral reasoning. This allowed an unprecedented parallel review of the neural networks associated with each of these cognitive domains. The results confirmed that the anterior left IFG region involved in semantic control is reliably engaged in all four social domains. This supports the hypothesis that social cognition is partly regulated by the neurocognitive system underpinning semantic control.

Establishing task- and modality-dependent dissociations between the semantic and default mode networks.

The default mode network (DMN) and semantic network (SN) are two of the most extensively studied systems, and both are increasingly used as clinical biomarkers in neurological studies. There are strong theoretical reasons to assume a relationship between the networks, as well as anatomical evidence that they might rely on overlapping cortical regions, such as the anterior temporal lobe (ATL) or angular gyrus (AG). Despite these strong motivations, the relationship between the two systems has received minimal attention. We directly compared the SN and DMN using a large (n = 69) distortion-corrected functional MRI (fMRI) dataset, spanning a range of semantic and nonsemantic tasks that varied input modality. The results showed that both networks fractionate depending on the semantic nature of the task, stimulus type, modality, and task difficulty. Furthermore, despite recent claims that both AG and ATL are semantic hubs, the two areas responded very differently, with results supporting the role of ATL, but not AG, in semantic representation. Specifically, the left ATL was positively activated for all semantic tasks, but deactivated during nonsemantic task performance. In contrast, the left AG was deactivated for all tasks, with the level of deactivation related to task difficulty. Thus, ATL and AG do not share a common interest in semantic tasks, but, rather, a common "disinterest" in nonsemantic tasks. The implications for the variability in the DMN, its cognitive coherence, and interpretation of resting-state fMRI data are discussed.

Healthy brain connectivity predicts atrophy progression in non-fluent variant of primary progressive aphasia.

Neurodegeneration has been hypothesized to follow predetermined large-scale networks through the trans-synaptic spread of toxic proteins from a syndrome-specific epicentre. To date, no longitudinal neuroimaging study has tested this hypothesis in vivo in frontotemporal dementia spectrum disorders. The aim of this study was to demonstrate that longitudinal progression of atrophy in non-fluent/agrammatic variant primary progressive aphasia spreads over time from a syndrome-specific epicentre to additional regions, based on their connectivity to the epicentre in healthy control subjects. The syndrome-specific epicentre of the non-fluent/agrammatic variant of primary progressive aphasia was derived in a group of 10 mildly affected patients (clinical dementia rating equal to 0) using voxel-based morphometry. From this region, the inferior frontal gyrus (pars opercularis), we derived functional and structural connectivity maps in healthy controls (n = 30) using functional magnetic resonance imaging at rest and diffusion-weighted imaging tractography. Graph theory analysis was applied to derive functional network features. Atrophy progression was calculated using voxel-based morphometry longitudinal analysis on 34 non-fluent/agrammatic patients. Correlation analyses were performed to compare volume changes in patients with connectivity measures of the healthy functional and structural speech/language network. The default mode network was used as a control network. From the epicentre, the healthy functional connectivity network included the left supplementary motor area and the prefrontal, inferior parietal and temporal regions, which were connected through the aslant, superior longitudinal and arcuate fasciculi. Longitudinal grey and white matter changes were found in the left language-related regions and in the right inferior frontal gyrus. Functional connectivity strength in the healthy speech/language network, but not in the default network, correlated with longitudinal grey matter changes in the non-fluent/agrammatic variant of primary progressive aphasia. Graph theoretical analysis of the speech/language network showed that regions with shorter functional paths to the epicentre exhibited greater longitudinal atrophy. The network contained three modules, including a left inferior frontal gyrus/supplementary motor area, which was most strongly connected with the epicentre. The aslant tract was the white matter pathway connecting these two regions and showed the most significant correlation between fractional anisotropy and white matter longitudinal atrophy changes. This study showed that the pattern of longitudinal atrophy progression in the non-fluent/agrammatic variant of primary progressive aphasia relates to the strength of connectivity in pre-determined functional and structural large-scale speech production networks. These findings support the hypothesis that the spread of neurodegeneration occurs by following specific anatomical and functional neuronal network architectures.

Mapping the Multiple Graded Contributions of the Anterior Temporal Lobe Representational Hub to Abstract and Social Concepts: Evidence from Distortion-corrected fMRI.

A growing body of recent convergent evidence indicates that the anterior temporal lobe (ATL) has connectivity-derived graded differences in semantic function: the ventrolateral region appears to be the transmodal, omni-category center-point of the hub whilst secondary contributions come from the peripheries of the hub in a manner that reflects their differential connectivity to different input/output modalities. One of the key challenges for this neurocognitive theory is how different types of concept, especially those with less reliance upon external sensory experience (such as abstract and social concepts), are coded across the graded ATL hub. We were able to answer this key question by using distortion-corrected fMRI to detect functional activations across the entire ATL region and thus to map the neural basis of social and psycholinguistically-matched abstract concepts. Both types of concept engaged a core left-hemisphere semantic network, including the ventrolateral ATL, prefrontal regions and posterior MTG. Additionally, we replicated previous findings of weaker differential activation of the superior and polar ATL for the processing of social stimuli, in addition to the stronger, omni-category activation observed in the vATL. These results are compatible with the view of the ATL as a graded transmodal substrate for the representation of coherent concepts.

A Neuropsychological Perspective on Abstract Word Representation: From Theory to Treatment of Acquired Language Disorders.

Natural languages are rife with words that describe feelings, introspective states, and social constructs (e.g., liberty, persuasion) that cannot be directly observed through the senses. Effective communication demands linguistic competence with such abstract words. In clinical neurological settings, abstract words are especially vulnerable to the effects of stroke and neurodegenerative conditions such as Alzheimer's disease. A parallel literature in cognitive neuroscience suggests that abstract and concrete words are at least partially neuroanatomically dissociable. Much remains to be learned about the nature of lexical-semantic deficits of abstract words and how best to promote their recovery. Here, we review contemporary theoretical approaches to abstract-concrete word representation with an aim toward contextualizing patient-based dissociations for abstract words. We then describe a burgeoning treatment approach for targeting abstract words and suggest a number of potential strategies for future interventions. We argue that a deeper understanding of is essential for informing language rehabilitation.

Frontal white matter tracts sustaining speech production in primary progressive aphasia.

In primary progressive aphasia (PPA), speech and language difficulties are caused by neurodegeneration of specific brain networks. In the nonfluent/agrammatic variant (nfvPPA), motor speech and grammatical deficits are associated with atrophy in a left fronto-insular-striatal network previously implicated in speech production. In vivo dissection of the crossing white matter (WM) tracts within this "speech production network" is complex and has rarely been performed in health or in PPA. We hypothesized that damage to these tracts would be specific to nfvPPA and would correlate with differential aspects of the patients' fluency abilities. We prospectively studied 25 PPA and 21 healthy individuals who underwent extensive cognitive testing and 3 T MRI. Using residual bootstrap Q-ball probabilistic tractography on high angular resolution diffusion-weighted imaging (HARDI), we reconstructed pathways connecting posterior inferior frontal, inferior premotor, insula, supplementary motor area (SMA) complex, striatum, and standard ventral and dorsal language pathways. We extracted tract-specific diffusion tensor imaging (DTI) metrics to assess changes across PPA variants and perform brain-behavioral correlations. Significant WM changes in the left intrafrontal and frontostriatal pathways were found in nfvPPA, but not in the semantic or logopenic variants. Correlations between tract-specific DTI metrics with cognitive scores confirmed the specific involvement of this anterior-dorsal network in fluency and suggested a preferential role of a posterior premotor-SMA pathway in motor speech. This study shows that left WM pathways connecting the speech production network are selectively damaged in nfvPPA and suggests that different tracts within this system are involved in subcomponents of fluency. These findings emphasize the emerging role of diffusion imaging in the differential diagnosis of neurodegenerative diseases.

The anterior temporal lobes support residual comprehension in Wernicke's aphasia.

Wernicke's aphasia occurs after a stroke to classical language comprehension regions in the left temporoparietal cortex. Consequently, auditory-verbal comprehension is significantly impaired in Wernicke's aphasia but the capacity to comprehend visually presented materials (written words and pictures) is partially spared. This study used functional magnetic resonance imaging to investigate the neural basis of written word and picture semantic processing in Wernicke's aphasia, with the wider aim of examining how the semantic system is altered after damage to the classical comprehension regions. Twelve participants with chronic Wernicke's aphasia and 12 control participants performed semantic animate-inanimate judgements and a visual height judgement baseline task. Whole brain and region of interest analysis in Wernicke's aphasia and control participants found that semantic judgements were underpinned by activation in the ventral and anterior temporal lobes bilaterally. The Wernicke's aphasia group displayed an 'over-activation' in comparison with control participants, indicating that anterior temporal lobe regions become increasingly influential following reduction in posterior semantic resources. Semantic processing of written words in Wernicke's aphasia was additionally supported by recruitment of the right anterior superior temporal lobe, a region previously associated with recovery from auditory-verbal comprehension impairments. Overall, the results provide support for models in which the anterior temporal lobes are crucial for multimodal semantic processing and that these regions may be accessed without support from classic posterior comprehension regions.

Mapping semantic space: Exploring the higher-order structure of word meaning.

Multiple representation theories posit that concepts are represented via a combination of properties derived from sensorimotor, affective, and linguistic experiences. Recently, it has been proposed that information derived from social experience, or socialness, represents another key aspect of conceptual representation. How these various dimensions interact to form a coherent conceptual space has yet to be fully explored. To address this, we capitalized on openly available word property norms for 6339 words and conducted a large-scale investigation into the relationships between 18 dimensions. An exploratory factor analysis reduced the dimensions to six higher-order factors: sub-lexical, distributional, visuotactile, body action, affective and social interaction. All these factors explained unique variance in performance on lexical and semantic tasks, demonstrating that they make important contributions to the representation of word meaning. An important and novel finding was that the socialness dimension clustered with the auditory modality and with mouth and head actions. We suggest this reflects experiential learning from verbal interpersonal interactions. Moreover, formally modelling the network structure of semantic space revealed pairwise partial correlations between most dimensions and highlighted the centrality of the interoception dimension. Altogether, these findings provide new insights into the architecture of conceptual space, including the importance of inner and social experience, and highlight promising avenues for future research.

Overlapping neural correlates underpin theory of mind and semantic cognition: Evidence from a meta-analysis of 344 functional neuroimaging studies.

Key unanswered questions for cognitive neuroscience include whether social cognition is underpinned by specialised brain regions and to what extent it simultaneously depends on more domain-general systems. Until we glean a better understanding of the full set of contributions made by various systems, theories of social cognition will remain fundamentally limited. In the present study, we evaluate a recent proposal that semantic cognition plays a crucial role in supporting social cognition. While previous brain-based investigations have focused on dissociating these two systems, our primary aim was to assess the degree to which the neural correlates are overlapping, particularly within two key regions, the anterior temporal lobe (ATL) and the temporoparietal junction (TPJ). We focus on activation associated with theory of mind (ToM) and adopt a meta-analytic activation likelihood approach to synthesise a large set of functional neuroimaging studies and compare their results with studies of semantic cognition. As a key consideration, we sought to account for methodological differences across the two sets of studies, including the fact that ToM studies tend to use nonverbal stimuli while the semantics literature is dominated by language-based tasks. Overall, we observed consistent overlap between the two sets of brain regions, especially in the ATL and TPJ. This supports the claim that tasks involving ToM draw upon more general semantic retrieval processes. We also identified activation specific to ToM in the right TPJ, bilateral anterior mPFC, and right precuneus. This is consistent with the view that, nested amongst more domain-general systems, there is specialised circuitry that is tuned to social processes.

Socialness effects in lexical-semantic processing.

Contemporary theories of semantic representation posit that social experience is an important source of information for deriving meaning. However, there is a lack of behavioral evidence in support of this proposal. The aim of the present work was to test whether words' degree of social relevance, or socialness, influences lexical-semantic processing. In Study 1, across a series of item-level regression analyses, we found that (a) socialness can facilitate responses in lexical, semantic, and memory tasks, and (b) limited evidence for an interaction of socialness with concreteness. In Studies 2-3, we tested the preregistered hypothesis that social words, compared to nonsocial words, will be associated with faster and more accurate responses during a syntactic classification task. We found that socialness has a facilitatory effect on noun decisions (Study 3), but not verb decisions (Study 2). Overall, our results suggest that the socialness of a word affects lexical-semantic processing but also that this is task-dependent. These findings constitute novel evidence in support of proposals that social information is an important dimension of semantic representation. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Investigating the role of working memory resources across aesthetic and non-aesthetic judgements.

Aesthetic judgements dominate much of daily life by guiding how we evaluate objects, people, and experiences in our environment. One key question that remains unanswered is the extent to which more specialised or largely general cognitive resources support aesthetic judgements. To investigate this question in the context of working memory, we examined the extent to which a working memory load produces similar or different response time interference on aesthetic compared with non-aesthetic judgements. Across three pre-registered experiments that used Bayesian multi-level modelling approaches (N > 100 per experiment), we found clear evidence that a working memory load produces similar response time interference on aesthetic judgements relative to non-aesthetic (motion) judgements. We also showed that this similarity in processing across aesthetic versus non-aesthetic judgements holds across variations in the form of art (people vs. landscape; Experiments 1-3), medium type (artwork vs. photographs; Experiment 2), and load content (art images vs. letters; Experiments 1-3). These findings suggest that across a range of experimental contexts, as well as different processing streams in working memory (e.g., visual vs. verbal), aesthetic and motion judgements commonly rely on a domain-general cognitive system, rather than a system that is more specifically tied to aesthetic judgements. In doing so, these findings shine new light on the working memory resources that support aesthetic judgements, as well as on how domain-general cognitive systems operate more generally in cognition.

Social semantics: the organization and grounding of abstract concepts.

concepts, like justice and friendship, are central features of our daily lives. Traditionally, abstract concepts are distinguished from other concepts in that they cannot be directly experienced through the senses. As such, they pose a challenge for strongly embodied models of semantic representation that assume a central role for sensorimotor information. There is growing recognition, however, that it is possible for meaning to be 'grounded' via cognitive systems, including those involved in processing language and emotion. In this article, we focus on the specific proposal that social significance is a key feature in the representation of some concepts. We begin by reviewing recent evidence in favour of this proposal from the fields of psycholinguistics and neuroimaging. We then discuss the limited extent to which there is consensus about the definition of 'socialness' and propose essential next steps for research in this domain. Taking one such step, we describe preliminary data from an unprecedented large-scale rating study that can help determine how socialness is distinct from other facets of word meaning. We provide a backdrop of contemporary theories regarding semantic representation and social cognition and highlight important predictions for both brain and behaviour. This article is part of the theme issue 'Concepts in interaction: social engagement and inner experiences'.

Graded functional organisation in the left inferior frontal gyrus: evidence from task-free and task-based functional connectivity.

The left inferior frontal gyrus (LIFG) has been ascribed key roles in numerous cognitive domains, including language, executive function and social cognition. However, its functional organisation, and how the specific areas implicated in these cognitive domains relate to each other, is unclear. Possibilities include that the LIFG underpins a domain-general function or, alternatively, that it is characterized by functional differentiation, which might occur in either a discrete or a graded pattern. The aim of the present study was to explore the topographical organisation of the LIFG using a bimodal data-driven approach. To this end, we extracted functional connectivity (FC) gradients from 1) the resting-state fMRI time-series of 150 participants (77 female), and 2) patterns of co-activation derived meta-analytically from task data across a diverse set of cognitive domains. We then sought to characterize the FC differences driving these gradients with seed-based resting-state FC and meta-analytic co-activation modelling analyses. Both analytic approaches converged on an FC profile that shifted in a graded fashion along two main organisational axes. An anterior-posterior gradient shifted from being preferentially associated with high-level control networks (anterior LIFG) to being more tightly coupled with perceptually-driven networks (posterior). A second dorsal-ventral axis was characterized by higher connectivity with domain-general control networks on one hand (dorsal LIFG), and with the semantic network, on the other (ventral). These results provide novel insights into a graded functional organisation of the LIFG underpinning both task-free and task-constrained mental states, and suggest that the LIFG is an interface between distinct large-scale functional networks.

A network-level test of the role of the co-activated default mode network in episodic recall and social cognition.

Resting-state network research is extremely influential, yet the functions of many networks remain unknown. In part, this is due to typical (e.g., univariate) analyses independently testing the function of individual regions and not examining the full set of regions that form a network whilst co-activated. Connectivity is dynamic and the function of a region may change based on its current connections. Therefore, determining the function of a network requires assessment at this network-level. Yet popular theories implicating the default mode network (DMN) in episodic memory and social cognition, rest principally upon analyses performed at the level of individual brain regions. Here we use independent component analysis to formally test the role of the DMN in episodic and social processing at the network level. As well as an episodic retrieval task, two independent datasets were employed to assess DMN function across the breadth of social cognition; a person knowledge judgement and a theory of mind task. Each task dataset was separated into networks of co-activated regions. In each, the co-activated DMN, was identified through comparison to an a priori template and its relation to the task model assessed. This co-activated DMN did not show greater activity in episodic or social tasks than high-level baseline conditions. Thus, no evidence was found to support hypotheses that the co-activated DMN is involved in explicit episodic or social tasks at a network-level. The networks associated with these processes are described. Implications for prior univariate findings and the functional significance of the co-activated DMN are considered.

Convergent connectivity and graded specialization in the rostral human temporal lobe as revealed by diffusion-weighted imaging probabilistic tractography.

In recent years, multiple independent neuroscience investigations have implicated critical roles for the rostral temporal lobe in auditory and visual perception, language, and semantic memory. Although arising in the context of different cognitive functions, most of these suggest that there is a gradual convergence of sensory information in the temporal lobe that culminates in modality- and perceptually invariant representations at the most rostral aspect. Currently, however, too little is known regarding connectivity within the human temporal lobe to be sure of exactly how and where convergence occurs; existing hypotheses are primarily derived on the basis of cross-species generalizations from invasive nonhuman primate studies, the validity of which is unclear, especially where language function is concerned. In this study, we map the connectivity of the human rostral temporal lobe in vivo for the first time using diffusion-weighted imaging probabilistic tractography. The results indicate that convergence of sensory information in the temporal lobe is in fact a graded process that occurs along both its longitudinal and lateral axes and culminates in the most rostral limits. We highlight the consistency of our results with those of prior functional neuroimaging, computational modeling, and patient studies. By going beyond simple fasciculus reconstruction, we systematically explored the connectivity of specific temporal lobe areas to frontal and parietal language regions. In contrast to the graded within-temporal lobe connectivity, this intertemporal connectivity was found to dissociate across caudal, mid, and rostral subregions. Furthermore, we identified a basal rostral temporal region with very limited connectivity to areas outside the temporal lobe, which aligns with recent evidence that this subregion underpins the extraction of modality- and context-invariant semantic representations.

The variation of function across the human insula mirrors its patterns of structural connectivity: evidence from in vivo probabilistic tractography.

The human insula is a functionally complex yet poorly understood region of the cortex, implicated in a wide range of cognitive, motor, emotion and somatosensory activity. To elucidate the functional role of the insula, the current study used in vivo probabilistic tractography to map the structural connectivity of seven anatomically-defined insular subregions. The connectivity patterns identified reveal two complementary insular networks connected via a dual route architecture, and provide key insights about the neural basis of the numerous functions ascribed to this area. Specifically, anterior-most insular regions were associated with a ventrally-based network involving orbital/inferior frontal and anterior/polar temporal regions, forming part of a key emotional salience and cognitive control network associated with the implementation of goal-directed behavior. The posterior and dorsal-middle insular regions were associated with a network focused on posterior and (to a lesser extent) anterior temporal regions via both dorsal and ventral pathways. This is consistent with the involvement of the insula in sound-to-speech transformations, with an implicated role in the temporal resolution, sequencing, and feedback processes crucial for auditory and motor processing, and the monitoring and adjustment of expressive performance.