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Connectivity of BA46 involvement in the executive control of language/ Estudio de la conectividad del AB46 en el control ejecutivo del lenguaje.

Although executive functions depend on extended dynamic networks including different brain areas (Koziol & Budding, 2009; Stuss & Alexander, 2007; Tonkonogy & Puente, 2009), it is well known that the prefrontal cortex plays a major controlling and monitoring role (Jurado & Rosselli, 2007; Stuss & Knight, 2002). The prefrontal areas of the frontal lobes (granular frontal cortex) can be regarded as association areas or intrinsic cortical areas (Mesulam, 2002). Luria (1980) considers that the prefrontal regions correspond to tertiary areas (which participate in processing information of various types) of the cerebral cortex. As a matter of fact, the prefrontal lobes maintain extensive connections, particularly with other cortical areas, the limbic system, the cortical and subcortical motor areas, and the sensory cortex (Damasio & Anderson, 2003).

Frequently, three different prefrontal syndromes associated with specific disturbances in executive functions are separated (e.g., Chayer & Freedman, 2001): Dorsolateral, medial and orbitofrontal. It has been proposed that these three prefrontal subsystems can be reduced to only two: dorsolateral and medial/ orbital (Ardila, 2008). The first one (dorsolateral) relates to cognition control (metacognition). Dysfunction in this region disrupts essential component cognitive processes, including impairments in working memory, abstracting difficulties, and inability to organize a behavioral response to novel or complex stimuli (Fuster, 2008; Stuss & Alexander, 2000). Various researchers, including Luria (1980), have noted perseveration, stimulus bound behavior, echopraxia, and echolalia. According to Fuster (2001, 2002, 2008), the most general executive function of the lateral prefrontal cortex is temporal organization of goal-directed actions in the domains of behavior, cognition, and language.

Lateral differences are observed: whereas left prefrontal damage is more directly associated with cognitive processes, right damage is associated with both restriction of affect and emotional dyscontrol and defects in the perception or comprehension of emotional information (Goldberg, 2001; Grafman, 2006). Anosognosia, impaired empathy, and defects in the appreciation of humor (Shammi & Stuss, 1999) are also found. Following lesion to the right dorsolateral area, a transcortical motor aprosodia is expected (Ross, 1981), whereas a left-sided dorsolateral lesion will produce a decline in verbal fluency on word generation tasks and so-called extrasylvian (transcortical) motor aphasia (Ardila, 2014; Berthier, 1999).

Extrasylvian (transcortical) motor aphasia due to lesions affecting the left dorsolateral prefrontal cortex is characterized by non-fluent language, good comprehension, and good repetition. Therefore, prosody, articulation, and grammar are preserved. The patient presents long latencies in language when beginning to speak or when answering questions. Open questions are slow and incomplete, and the patient tends to repeat the words included in the question. Expressive language is limited with some tendency to echolalia and perseveration; occasionally verbal paraphasias are observed. This type of aphasia has been interpreted as a language disturbance at the pragmatic level (use of the language according to the specific social context) (Benson & Ardila, 1996; Berthier, 1999).

The core Brodmann area (BA) in the dorsolateral prefrontal cortex is BA46. BA46 is known as anterior middle frontal gyrus. Actually, B A46 roughly corresponds with the dorsolateral prefrontal cortex. In the human brain it occupies approximately the middle third of the middle frontal gyrus and the most rostral portion of the inferior frontal gyrus (Mesulam, 1986, 2002). Interestingly, BA46 is regarded as one of the most recently evolved parts of the human brain that undergoes a prolonged period of maturation that lasts until adulthood (Collins, 2001).

Contemporary neuroimaging technique studies have supported the hypothesis regarding an active involvement of BA46 in linguistic processes (see Brodmann's Interactive Atlas), including verbal fluency (Abrahams et al., 2003), phonological processing (Heim, Opitz, Muller, & Friederici, 2003), semantic processing (Wang, 2008), and language comprehension (Turken & Dronkers, 2011). Taken together, all these findings support the conclusion that BA46 significantly participates in language. Furthermore, they suggest that it is not involved in a single linguistic process, but simultaneously in several verbal abilities.

It is noteworthy that the BA46 possesses extensive intracortical as well as fronto-subcortical connections (Cummings, 1983; Damasio & Anderson, 2003). Advancing the analysis of the functional connectivity of BA46 becomes most important in understanding its real contribution to the language brain system.

Currently, there are several techniques that can potentially demonstrate brain circuitries or networks. These techniques are grouped under the term "brain connectivity". Recently, a new alternative to study brain connectivity has been proposed by Robinson et al. (2010) known as meta-analytic connectivity modeling or MACM. MACM is based in automatic meta-analysis done by pooling co-activation patterns. The technique takes advantage of the Brainmap.org's repository of functional MRI studies, and of a special software (Sleuth) provided by the same group, to find, filter, organize, plot, and export the peaks coordinates for further statistical analysis of its results. Sleuth provides a list of foci, in Talairach or MNI coordinates, each one representing the center of mass of a cluster of activation. The method takes the region of interest (for instance, BA46), makes it the independent variable, and interrogates the database for studies showing activation of the chosen target. The query is easily filtered with different conditions (such as age, normal vs. patients, type of paradigm, domain of cognition, etc.). By pooling the data with these conditions the tool provides a universe of co-activations that can be statistically analyzed for significant commonality. As a final step, Activation Likelihood Estimation (ALE) (Laird et al., 2005; Turkeltaub et al., 2002) that can be performed utilizing GingerALE, another software also provided by Brainmap, assesses the probability of an event to occur at voxel level across the studies. Areas of coactivation will show a network related to the function and domains selected as filter criteria.

Considering the complex role of BA46 in language, a meta-analytic connectivity analysis utilizing MACM on the participation of BA46 in language was developed. The objective of this study as to analyze the left BA46 participation in the brain language circuits associated with different language functions.

Methods

Procedure

The DataBase of Brainmap (brainmap.org) was accessed utilizing Sleuth 2.2 on January 2, 2014. Sleuth is the software provided by Brainmap to query its database. The meta-analysis was intended to assess the network of coactivations in which BA46 is involved.

The search conditions were: (1) studies reporting BA46 activation; (2) studies using fMRI ; (3) context: normal subjects; (4) activations: activation only; (5) handedness: right-handed subjects; (6) age 18-60 years; (7) domain: cognition, subtype: language; (8) Language: English.

Data analysis

(ALE) meta-analysis was then performed utilizing GingerALE. ALE maps were thresholded at p<0.01 corrected for multiple comparisons and false discovery rate. Only clusters of 200 or more cubic mm where accepted as valid clusters. ALE results were overlaid onto an anatomical template suitable for MNI coordinates, also provided by BrainMap.org. For this purpose we utilized the Multi-Image Analysis GUI (Mango) (http://ric.uthscsa. edu/mango/). Mosaics of 5 x 6 insets of transversal fusion images were generated utilizing a plugin of the same tool, selecting every other image, starting on image No. 10, and exported to a 2D-jpg image. A 3D rendition of the brain was also obtained. The left hemisphere lateral view has been chosen for display.

Results

Nineteen papers corresponding to 60 experimental conditions with a total of 245 subjects were selected (subjects participating in two different experiments were counted as two subjects) (Table 1).

Table 2 presents the main loci of brain connectivity of BA46 by Meta-analytic Connectivity Modeling (MACM). Eleven different clusters of activation were found, mostly related to the left hemisphere (Figure 1 and Figure 2).

The first cluster includes the frontal areas 6, 44, 45, 46, and 47 in the left hemisphere. That is, the whole frontal system involved in language production. Noteworthy, this as an extensive cluster with a volume about eight times larger than Cluster #2 and about 11 times larger than Cluster #3. Indeed, the rest of the activation clusters are relatively small.

The second cluster includes the right insula. Cluster #3 includes the left fusiforme gyrus (most likely activation of the culmen of the cerebellum is explained by the smoothing effect of the adjacent activation of the left fusiform gyrus). Cluster #4 includes the left fusiforme gyrus as well as its anterior extension (BA20). Cluster #5 and Cluster #10 refer to the left occipital lobe; whereas Cluster #7 corresponds to the right BA46. Cluster #8 is located in the superior parietal lobe. Cluster #9 corresponds to the Wernicke's area. And finally, Cluster #11 refers to the left basal ganglia.

[FIGURE 1 OMITTED]

Discussion

The main connectivity revealed by the extent and intensity of the principal clusters with expressive areas makes evident that BA46 basically participates in a language production system, which also includes BA44, BA45, BA47, and BA6 in the left frontal lobe. This system could be referred as the "frontal language production system", or simply, the "Broca's system" and in the current study it presents very small co activation with other brain areas. Noteworthy, functional studies have demonstrated that BA6 is involved in diverse language functions, including speech motor programming (Fox et al., 2000; Shuster & Lemieux, 2005), phonological processing (McDermott et al., 2003), language switching (Price, Green, & Von Studnitz, 1999), and even syntactic processing (Inui et al., 1998). The medial extension of BA6 corresponds to the supplementary motor area, a brain area clearly involved in language processing (De Carli et al., 2007; Basho et al., 2007). The involvement of BA44, BA45, and BA47 in language production, on the other hand, is quite obvious (Hickok & Poeppel, 2004; Grodzinsky & Amunts, 2006; Price, 2010).

The connections with the insula (Cluster #2) are understandable. Functional studies have demonstrated that the insula represents a core area in language processing, extensively connected with anterior and posterior language areas (see Ardila, Bernal, & Rosselli, 2014). Insular damage has been associated with aphasia since the 19th century (Wernicke, 1874). Thus, it is quite evident the role of the insula in diverse language functions (Ardila, 1999). However, it was quite surprising to find the activation in the right insula. It does not seem easy to find an explanation to the right-lateralized activation.

[FIGURE 2 OMITTED]

BA46 turned out to have some connections with posterior language areas involved in phonological, lexical, and semantic language processing (BA20, BA21, BA22 and BA37); these connections, however, are notoriously weaker than the connections with the rest of the frontal language production system (BA44, BA45, BA47, and BA6). The volume of Cluster #4 (BA37 and BA20) is some 14 times smaller than the volume of Cluster #1 ("language production system": BA44, BA45, BA47, and BA6). The volume of Cluster #9 (Wernicke's area: BA21 and BA22) is over 60 times smaller than the volume of Cluster #1. Evidently, the primary role of BA46 in language is related with language production control, not with phonological, lexical or semantic understanding. This is a conclusion easy to draw just taking a look of Figure 1 (right).

BA46 significant--albeit weak--connection with left BA37 (fusiform gyrus) is particularly interesting. It has been pointed out that left BA37 is a common node of two distinct networks visual recognition (perception) and semantic language functions (Ardila, Bernal, & Rosselli, 2015). Many of the tasks included in the current analyses involved semantic decisions using visual information. Therefore, results suggest certain involvement of BA46 in visual/semantic associations of words and language understanding. Noteworthy, two clusters (Cluster #5 and Cluster #10) point to some connections of BA46 with the occipital lobe. The co-activation of the occipital lobe is not totally unexpected considering the existence of a fasciculus going between the occipital lobe and the prefrontal cortex: the inferior occipitofrontal fasciculus. It has been suggested that this fasciculus is involved in language processing (Duffau et al., 2009; Mandonnet et al., 2007). Furthermore, it has been observed that this fasciculus has two branches: (1) a superficial and dorsal branch, which connects the frontal lobe with the superior parietal lobe and the posterior portion of the superior and middle occipital gyri; and (2) a deep and ventral branch, which connects the frontal lobe with the posterior portion of the inferior occipital gyrus and the posterior temporo-basal area. This observation is congruent with the role of this fasciculus in the semantic system, by showing that it is mainly connected with two areas involved in semantics: the occipital associative extrastriate cortex and the temporo-basal region (Martino et al., 2010).

The weakest cluster observed in this analysis was Cluster #11 (left lenticular nucleus). Indeed, the frontal cortex, including the dorsolateral prefrontal area, has extensive connections with subcortical areas, in particular with the striatum (Damasio & Anderson, 2003; Mesulam, 2002). Complex behavior has been frequently related with fronto-subcortical circuits (Bonelli & Cummings, 2007; Cummings, 1993).

Regardless of the diverse limitations that can be pointed to the present study (specific characteristics of the sample, implicit limitations of the method that was used, inclusion of language as a whole without distinguishing different language abilities, etc.) it can be concluded that BA46 is involved together with BA44, BA45, BA47 and BA6 in kind of "frontal language production system" (or "Broca's system"). A notoriously smaller albeit significant participation is also observed in language semantics and language understanding. Furthermore, considering that BA46 is the core dorsolateral prefrontal area involved in cognition control (metacognition), it can be suggested that BA46 plays the executive control in this frontal language production system; as a matter of fact, when BA46 is damaged, no active language production is observed (extrasylvian or transcortical motor aphasia; Benson & Ardila, 1996; Berthier, 1999).

doi: 10.7334/psicothema2015.174

References

References marked with an asterisk indicate studies included in the meta-analysis.

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Alfredo Ardila (1), Byron Bernal (2) and Monica Rosselli (3)

(1) Florida International University, (2) Miami Children's Hospital and (3) Florida Atlantic University

Received: June 29, 2015 * Accepted: November 18, 2015

Corresponding author: Alfredo Ardila

Department of Communication Sciences and Disorders

Florida International University

33199 Miami (E.E.U.U.)

e-mail: ardilaa@fiu.edu
Table 1

Studies of language paradigms included in the meta-analysis

Publication              Paradigm                            n    Foci

Booth et al., 2002       Visual Rhyming--Control             13   11
                         Visual Meaning--Rhyming             13   3
                         Meaning--Rhyming                    13   8
Dapretto & Bookheimer,   Semantic vs. Rest                   8    8
  1999
Devlin et al., 2003      Semantic + Phonological--Rest       12   26
                         Phonological > Semantic             12   34
Kang et al., 1999        Syntactic VPs--Fixation             14   7
                         Semantic VPs--Fixation              14   7
Schlosser et al., 1998   Verbal Fluency > Baseline           6    9
Jackson & Schacter,      Associative Encoding--Fixation      12   61
  2004
Poldrack et al., 2001    Rhyme Judgment                      8    2
                         Convex Compression Response         8    5
Peck et al., 2004        Sentences vs. Viewing Objects       10   13
Rowan et al., 2004       Verb Generation                     10   13
                         Action Tool Word Retrieval          20   7
Damasio et al., 2001     Action Word Retrieval               20   9
                         Concrete Entities--Picture          20   5
                           Control
Simmons et al., 2008     Word Assoc > Situation Generation   10   32
                         Early > Late Property Generation    10   26
                         Early > Late, Word Assoc > Sit      10   3
                           Gen
Sharp et al., 2010       Semantic Low Percept High Dif vs.   12   2
                         Semantic Low Perceptual Low Dif
                         Semantic High Perceptual Low Dif    12   5
                           vs.
                         Semantic Low Perceptual High
                           Diff,
Davis et al, 2008        All Words vs. Letter Strings        12   9
Desai et al., 2006       Generate Regular Verbs vs. Read     25   21
                         Regular Present Tense Verbs
                         Generate Irregular Verbs vs Read    25   25
                         Irregular Present Tense Verbs
                         Areas Correlated with Response      25   31
                           Time
Longe et al., 2007       Inflections (Nouns + Verbs) vs.     12   14
                         Baseline
Sabsevitz et al.,        Concrete > Abstract                 28   26
  2005
Bedny et al., 2006       Words (Nouns + Verbs) > Non-words   13   6
Rosen et al., 2000       Word Stem Completion--Fixation      8    7
Tan et al., 2003         Rhyme Decision English              12   4

Table 2

Main loci of brain connectivity of BA46 in language tasks by Meta-
analytic Connectivity Modeling (MACM)

Region (BA)                       x     y     z      ALE

Cluster #1
L Middle frontal gyrus (46)      -46   34     8    0.043545
L Precental frontal gyrus (6)    -50    6    24    0.031531
L Inferior frontal gyrus (47)    -36   28    -10   0.031294
L Inferior frontal gyrus (45)    -48   22    -6    0.030777
L Inferior frontal gyrus (44)    -54   14    40    0.030316
Cluster #2
R Insula (13)                    48    16    -4    0.028763
R Insula (13)                    40    24    -4    0.024963
Cluster #3
L Fusiform gyrus (37)            -42   -50   -20   0.024459
L Cerebellum culmen              -36   -46   -28   0.018386
Cluster #4
L Fusiform gyrus (37)            -52   -48   -2    0.022463
L Inferior temporal gyrus (20)   -54   -54   -12   0.019619
Cluster #5
L Occipital (18)                 -24   -94   -4    0.019987
Cluster #6
L Medial frontal lobe (32)       -8    18    44    0.020837
Cluster #7
R Middle frontal gyrus (46)      50    30    18    0.022843
Cluster #8
L Superior parietal (7)          -26   -66   50    0.020237
Cluster #9
L Superior temporal lobe (22)    -46   -24    0    0.016397
L Middle temporal lobe (21)      -48   -30   -4    0.016394
Cluster #10
L Middle occipital (19)          -32   -76   26    0.017609
Cluster #11
L Lenticular                     -26   14     2    0.01659

Region (BA)                      Volume ([mm.sup.3])

Cluster #1
L Middle frontal gyrus (46)            18,904
L Precental frontal gyrus (6)
L Inferior frontal gyrus (47)
L Inferior frontal gyrus (45)
L Inferior frontal gyrus (44)
Cluster #2
R Insula (13)                           2,424
R Insula (13)
Cluster #3
L Fusiform gyrus (37)                   1,728
L Cerebellum culmen
Cluster #4
L Fusiform gyrus (37)                   1,288
L Inferior temporal gyrus (20)
Cluster #5
L Occipital (18)                         568
Cluster #6
L Medial frontal lobe (32)               512
Cluster #7
R Middle frontal gyrus (46)              488
Cluster #8
L Superior parietal (7)                  432
Cluster #9
L Superior temporal lobe (22)            296
L Middle temporal lobe (21)
Cluster #10
L Middle occipital (19)                  288
Cluster #11
L Lenticular                             256
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Title Annotation:texto en ingles; BA46; circunvolucion frontal media anterior
Author:Ardila, Alfredo; Bernal, Byron; Rosselli, Monica
Publication:Psicothema
Date:Jan 1, 2016
Words:4602
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