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Background and Objective: Modern neuroimaging developments have demonstrated that cognitive functions correlate with brain networks rather than specific areas. The purpose of this paper was to analyze the connectivity of Broca's area based on language tasks. Methods: A connectivity modeling study was performed by pooling data of Broca's activation in language tasks. Fifty-seven papers that included 883 subjects in 84 experiments were analyzed. Analysis of Likelihood Estimates of pooled data was utilized to generate the map; thresholds at p < 0.01 were corrected for multiple comparisons and false discovery rate. Resulting images were co-registered into MNI standard space. Results: A network consisting of 16 clusters of activation was obtained. Main clusters were located in the frontal operculum, left posterior temporal region, supplementary motor area, and the parietal lobe. Less common clusters were seen in the sub-cortical structures including the left thalamus, left putamen, secondary visual areas, and the right cerebellum. Conclusion: Broca's area-44-related networks involved in language processing were demonstrated utilizing a pooling-data connectivity study. Significance, interpretation, and limitations of the results are discussed. [ABSTRACT FROM AUTHOR]

This review scrutinizes several findings on human hierarchical processing within the prefrontal cortex (PFC) in diverse cognitive domains. Converging evidence from previous studies has shown that the PFC, specifically, BA44, may function as the essential region for hierarchical processing across the domains. In language fMRI studies, BA 44 was significantly activated for the hierarchical processing of center-embedded sentences and this pattern of activations was also observed in artificial grammar. The same pattern was observed in the visuo-spatial domain where BA44 was actively involved in the processing of hierarchy for the visual symbol. Musical syntax, which is the rule-based arrangement of musical sets, has also been construed as hierarchical processing as in the language domain such that the activation in BA44 was observed in a chord sequence paradigm. P600 ERP was also engendered during the processing of musical hierarchy. Along with a longstanding idea that a human's number faculty is developed as a "by-product of language faculty", BA44 was closely involved in hierarchical processing in mental arithmetic. This review extended its discussion of hierarchical "processing" to hierarchical "behavior", that is, human action which has been referred to as being hierarchically composed. Several lesion and TMS studies supported the involvement of BA44 for hierarchical processing in the action domain. Lastly, the hierarchical organization of cognitive controls was discussed within the PFC, forming a cascade of top-down hierarchical processes operating along a posterior-to-anterior axis of the lateral PFC including BA44 within the network. It is proposed that PFC is actively involved in different forms of hierarchical processing and specifically BA44 may play an integral role in the process. Taking levels of proficiency and subcortical areas into consideration may provide further insight into the functional role of BA44 for hierarchical processing. [ABSTRACT FROM AUTHOR]

Humans are unique in their ability to parse hierarchical structures of sentences. Previous studies demonstrated that syntactic processes at different hierarchies are subserved by distinct subregions in left inferior frontal gyrus (LIFG), in which BA45 is mainly involved in processing lower-level syntactic structures and BA44 is mainly involved in processing higher-level syntactic structures. However, little is known about whether semantic processes at different syntactic hierarchies show similar dissociations in LIFG. In the present fMRI experiment, participants read sentences with the structure "subject noun + verb + numeral + classifier + object noun", in which the object noun is constrained by the classifier at the lower-level and by the verb at the higher-level. The object noun was manipulated to be either semantically congruent or incongruent with the classifier at the lower-level and/or with the verb at the higher-level. Both the whole brain contrasts and the region of interest (ROI) analyses showed that, in LIFG, the semantic process of integrating the object noun with the classifier induced stronger activation in BA45 whereas the semantic process of integrating the object noun with the verb induced stronger activation in BA44. This dissociation demonstrates a neural segregation for semantic processes at different syntactic hierarchies, with the lower-level process relying more on neural substrates for general semantic processes and the higher-level process relying more on neural substrates for processing structural hierarchies. • In a minimal task demand, activations in LIFG enhanced for semantic congruence vs. incongruence. • The BA45 for the lower-level semantic process peaked around the locus related to semantic unification. • The BA44 for the higher-level semantic process peaked around the locus related to syntactic unification. [ABSTRACT FROM AUTHOR]

Among the many lines of research that have been exploring how embodiment contributes to cognition, one focuses on how the neural substrates of language may be shared, or at least closely coupled, with those of action. This paper revisits a particular proposal that has received considerable attention-namely, that the forms of hierarchical sequencing that characterize both linguistic syntax and goal-directed action are underpinned partly by common mechanisms in left Brodmann area (BA) 44, a cortical region that is not only classically regarded as part of Broca's area, but is also a core component of the human Mirror Neuron System. First, a recent multi-participant, multi-round debate about this proposal is summarized together with some other relevant findings. This review reveals that while the proposal is supported by a variety of theoretical arguments and empirical results, it still faces several challenges. Next, a narrower application of the proposal is discussed, specifically involving the basic word order of subject (S), object (O), and verb (V) in simple transitive clauses. Most languages are either SOV or SVO, and, building on prior work, it is argued that these strong syntactic tendencies derive from how left BA44 represents the sequential-hierarchical structure of goal-directed actions. Finally, with the aim of clarifying what it might mean for syntax and action to have "common" neural mechanisms in left BA44, two different versions of the main proposal are distinguished. Hypothesis 1 states that the very same neural mechanisms in left BA44 subserve some aspects of hierarchical sequencing for syntax and action, whereas Hypothesis 2 states that anatomically distinct but functionally parallel neural mechanisms in left BA44 subserve some aspects of hierarchical sequencing for syntax and action. Although these two hypotheses make different predictions, at this point neither one has significantly more explanatory power than the other, and further research is needed to elaborate and test them., Competing Interests: The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Kemmerer.)

Abstract: Verbs have two separate levels of meaning. One level reflects the uniqueness of every verb and is called the “root”. The other level consists of a more austere representation that is shared by all the verbs in a given class and is called the “event structure template”. We explore the following hypotheses about how, with specific reference to the motor features of action verbs, these two distinct levels of semantic representation might correspond to two distinct levels of the mirror neuron system. Hypothesis 1: Root-level motor features of verb meaning are partially subserved by somatotopically mapped mirror neurons in the left primary motor and/or premotor cortices. Hypothesis 2: Template-level motor features of verb meaning are partially subserved by representationally more schematic mirror neurons in Brodmann area 44 of the left inferior frontal gyrus. Evidence has been accumulating in support of the general neuroanatomical claims made by these two hypotheses—namely, that each level of verb meaning is associated with the designated cortical areas. However, as yet no studies have satisfied all the criteria necessary to support the more specific neurobiological claims made by the two hypotheses—namely, that each level of verb meaning is associated with mirror neurons in the pertinent brain regions. This would require demonstrating that within those regions the same neuronal populations are engaged during (a) the linguistic processing of particular motor features of verb meaning, (b) the execution of actions with the corresponding motor features, and (c) the observation of actions with the corresponding motor features. [Copyright &y& Elsevier]

This review scrutinizes several findings on human hierarchical processing within the prefrontal cortex (PFC) in diverse cognitive domains. Converging evidence from previous studies has shown that the PFC, specifically, BA44, may function as the essential region for hierarchical processing across the domains. In language fMRI studies, BA 44 was significantly activated for the hierarchical processing of center-embedded sentences and this pattern of activations was also observed in artificial grammar. The same pattern was observed in the visuo-spatial domain where BA44 was actively involved in the processing of hierarchy for the visual symbol. Musical syntax, which is the rule-based arrangement of musical sets, has also been construed as hierarchical processing as in the language domain such that the activation in BA44 was observed in a chord sequence paradigm. P600 ERP was also engendered during the processing of musical hierarchy. Along with a longstanding idea that a human's number faculty is developed as a "by-product of language faculty", BA44 was closely involved in hierarchical processing in mental arithmetic. This review extended its discussion of hierarchical "processing" to hierarchical "behavior", that is, human action which has been referred to as being hierarchically composed. Several lesion and TMS studies supported the involvement of BA44 for hierarchical processing in the action domain. Lastly, the hierarchical organization of cognitive controls was discussed within the PFC, forming a cascade of top-down hierarchical processes operating along a posterior-to-anterior axis of the lateral PFC including BA44 within the network. It is proposed that PFC is actively involved in different forms of hierarchical processing and specifically BA44 may play an integral role in the process. Taking levels of proficiency and subcortical areas into consideration may provide further insight into the functional role of BA44 for hierarchical processing.