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<title>Parcellation of the language cortex</title>
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<h1>The role of Broca’s area in language</h1>
<p>Broca’s area, has long been known to support language production (Broca, 1861; Sahin et al., 2009) and comprehension processes (Zurif, 1980). For the localization of Broca’s area, defined as consisting of BA 44 and BA 45, see Fig. 1. Its function in language comprehension is still a matter of considerable debate (Grodzinsky & Santi, 2008; Hagoort, 2005; 2008; Rogalsky & Hickok, 2011). Although the different views agree upon the involvement of Broca’s area in language comprehension, they debate its particular role in this process. This discussion takes place on multiple levels. At the most general level, the claim is made that Broca’s region supports action observation and execution and that its part in language is related to motor-based speech production and comprehension processes (Pulvermüller & Fadiga, 2010; Rizzolatti & Arbib, 1998). At the next level, the claim is that Broca’s region supports verbal working memory (Smith & Jonides, 1999) and that this is why this region shows activation when processing syntactically complex sentences (Caplan & Waters, 1999; Rogalsky, Matchin & Hickok, 2008). At a linguistic level, subregions of Broca’s area have been allocated to different aspects of language processing, either seeing BA 44 as supporting syntactic structure building, BA 44/45 as supporting thematic role assignment and BA 45/47 supporting semantic processes (Friederici, 2002), or specifying Broca’s area (BA 44/45) as the region supporting the computation of syntactic movement (Grodzinsky, 2000), or defining Broca’s region (BA 44/45/47) as the space for the unification of different aspects in language (Hagoort, 2005). This debate was and is based on a large number of neuroimaging studies as well as neurophysiological and behavioral studies with healthy individuals and with patients suffering from circumscribed brain lesions in the IFG. The majority of these are described in different review articles published over the past decade (Bookheimer, 2002; Friederici, 2002; Grodzinsky, 2000; Grodzinsky & Friederici, 2006; Hagoort, 2005; Rogalsky & Hickok, 2011). The present review will not reiterate each of these studies, but will discuss recent studies that have contributed possible solutions to the open issues at the linguistic level and the related verbal working memory processes.
A large number of studies in different indo-european languages have investigated the neural substrate of syntactic processes by varying syntactic complexity. In these languages the canonical word order is subject-first either with a subject-verb-object or a subject-object-verb structure. Studies in these languages often compare brain activation for the processing of noncanonical object-first to canonical subject-first sentences using different sentence types in which the object-noun phrase is moved to a position in front of the subject-noun phase, called Movement in linguistics (for studies in different languages, see Table 4 and Fig. 5). In linguistic terms, this means that the object-noun phrase (now antecedent) leaves an empty position in the original structure (gap) of the sentence. What is analyzed in the imaging studies is the difference in the brain activation between sentences containing movement or not, or the difference between sentences varying the distance of the antecedent-gap relation (short/long). The studies listed in Table 4 show an activation increase in Broca’s area (BA 44 and/or BA 45) for Movement operations across different languages with the exceptions of three studies. These are: Caplan et al. (2002), who presented the critical sentences together with semantically implausible sentences and employed a plausibility judgement task, and two studies which only found IFG activation for a long, but not for a short antecedent-gap relation, suggesting an interaction between syntactic structure and distance as such (Cooke et al., 2001; Fiebach et al., 2005). However, the finding that these two studies only observed an effect for the long conditions could be explained by the fact that their short conditions differed from the long conditions in the number of intervening noun phrases.
To summarize, the processing of sentences in which the canonical word order is changed requires the involvement of Broca’s area.</p>
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<h1>Figure 5: Syntax-related activations in the left hemisphere</h1>
<div class="document">
<p>Maxima of activations for different types of syntactic complexity are color coded: studies investigating Movement (red), Scrambling (yellow) and Nesting (green). The studies reporting these maxima are listed in Table 4.</p>
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