Functional specificity for high-level linguistic processing in the human brain

“Note that these tasks were not designed to be directly comparable to one another, even though they have all been shown to produce behavioral difficulty effects with the hard condition, leading to lower accuracies and longer reaction times (2, 27).”

“(This data set partially overlaps with the data set reported in [13].”

“and bodies (Kanwisher 2010), and high-level regions specialized for understanding the meaning of sentences (Binder et al. 1997; Fedorenko et al. 2011) and the contents of other people’s thoughts (Saxe and Kanwisher 2003; Saxe and Powell 2006).”

“However, music selectivity was weak in raw voxel responses, and its detection required a decomposition method.”

“…from phonological/sub-lexical, to word level, to phrase/sentence level (Bautista and Wilson, 2016; Blank and Fedorenko, 2020; Blank et al., 2016; Fedorenko et al., 2011, 2012, 2020; Regev et al., 2021); and supports linguistic operations that are related to both the processing of word meanings…”

“Similarly, the so-called languageselective regions in the left posterior temporal lobe show a significant, albeit weaker, response to music [26].”

“…stimulus 91,100–102 — and show little or no response when participants perform demanding cognitive tasks, including solving arithmetic problems 22,91,103,104 and logic puzzles 105,106 , understanding computer code 107,108 and carrying out working memory and cognitive control tasks…”

“These tasks include music perception [16], arithmetic processing [16,60,61], executive function tasks [16,49], thinking about others’ mental states (A.”

“The location of these regions midway between higher visual areas, auditory areas and language and semantic areas (Fedorenko et al., 2011; Pobric et al., 2007; Visser et al., 2010) suggests a genuine MD region, situated to integrate higher visual, auditory and semantic/language processing.”

“In Fedorenko et al. (2011) we tried to ameliorate this concern by performing a whole-brain search—at liberal thresholds—for overlap between the responses to the language contrast (sentences > pseudoword lists) and the hard > easy contrast in each of the non-linguistic demanding tasks.”

“Indeed, we have recently shown for the case of high-level language processing that fROI-based methods reveal much greater functional specificity than was apparent in prior work using traditional methods (Fedorenko et al., 2011).”

“2 in Fedorenko et al. 2011; Fedorenko et al. 2010; 2012).”

“, Smith & Levy, 2013) – is primarily evident during naturalistic sentence processing in (1) the domain-specific, fronto-temporal language (LANG) network (Fedorenko et al., 2011), or (2) the domain-general, fronto-parietal multiple demand (MD) network (Duncan, 2010).”

“…could be symbolic but non-linguistic (for example, ‘9’), and the use of symbolic non-linguistic representations does not engage linguistic resources (for example, mathematical reasoning elicits no response in the language brain areas and is preserved in individuals with severe aphasia 51,77–79 ).”

“As expected, and replicating prior work (e.g., Fedorenko et al., 2010; Fedorenko et al., 2011; Blank et al., 2016; Mahowald & Fedorenko, 2016), the language fROIs showed a robust sentences > nonwords effect (ts(48)>8.”

“Specifically, language regions respond at or below a lowlevel baseline to tasks that have general demands similar to those of the sentence–picture matching task (see Fedorenko et al., 2011, 2012a).”

“High-level language processing recruits an extended set of cortical regions in the human brain (Blank et al. 2014; Tie et al. 2014) that are distinct from those recruited for other cognitive functions (Fedorenko et al. 2011; Blank et al. 2014).”

“…unit of sentence processing across varying experimental designs and task demands (Caplan et al. 2008) that can be spatially disentangled from surrounding domain-general areas serving working memory, cognitive control and music processing (Makuuchi et al. 2009; Fedorenko et al. 2011, 2012).”

“This localizer has been successfully used in a number of studies (Axelrod, Bar, Rees, & Yovel, 2015; Blank, Kanwisher, & Fedorenko, 2014; Fedorenko et al., 2011, 2012; Koster-Hale & Saxe, 2011; Maruyama, Pallier, Jobert, Sigman, & Dehaene, 2012; Xu et al., 2015).”

“Therefore these regions might not always activate during sentence comprehension, and damage will not necessarily result in sentence comprehension deficits.”

“Previous studies (e.g. Fedorenko et al., 2011) claimed that no brain area was more sensitive to syntactic information than to lexical information, therefore we also tested whether part of Broca's area responds more to the jabberwocky condition (JP) than to the scrambled real words (RWO).”

“…tasks, like arithmetic calculations, executive function tasks, music perception, action/gesture observation, and non-verbal social information (Fedorenko et al., 2011; Monti et al., 2012; Pritchett et al., 2018; Jouravlev et al., 2019; Paunov, 2019; see Fedorenko & Varley, 2016, for a review).”

“Unpredictable linguistic stimuli are known to evoke their largest responses in the language network – a set of frontal and temporal brain regions which respondmost strongly and selectively during language comprehension [127,128].”

“Some initial attempts have been made to address some of these challenges (Fedorenko et al., 2011; Xu et al., 2009; Yarkoni et al., 2011), but much remains to be done.”

“…such as phonological, semantic, and syntactic processing (Indefrey and Levelt, 2004; Hickok and Poeppel, 2007; Rauschecker and Scott, 2009; Fedorenko et al., 2011; Price, 2012), but also on widely distributed and often overlapping brain regions that make broader, or ‘domain-general’,…”

“…Thompson-Schill, 2014), which is more strongly activated by language tasks contrasted with math, working memory, cognitive control, or music tasks (Fedorenko et al., 2011) and is distinguished from the peripheral language-related sensorimotor systems such as the visual word form area (Dehaene and…”

“…functions are not strongly localized: many other areas outside the classical “language areas” are active during language processing; and, conversely, the language areas may also be activated during non-linguistic processing (Stowe et al., 2005; Anderson, 2010; see, however, Fedorenko et al., 2011).”

“Within the language network [51], NL-MVPD individuated four subnetworks reliably across two different tasks performed by the participants.”

“Domain generality does not, however, preclude the concomitant existence of domain-specific substrates for language, either due to ontogenetic (developmental) or phylogenetic (evolutionary) specialization within these systems, or to additional specialized circuitry (1, 5, 7, 9, 13, 37, 59)—although the actual demonstra…”

“, 1997), for judging others’ actions (Saxe and Kanwisher, 2003), for linguistic functions (Fedorenko et al., 2011) and so on.”

“, 2016; Van de Cavey and Hartsuiker, 2016; Hepner and Nozari, 2019), and interest in how distinct brain networks must coordinate to accomplish language comprehension and other complex cognitive processes (Fedorenko et al., 2011; Fedorenko, 2014).”

“[42,51] found different activated brain regions when comparing a music-localizer based on a scrambling manipulation to a languagelocalizer based on the reading of sentences versus lists of non-words.”

“However, lmSTC is by no means the only region consistently implicated in higher-order semantic processing, as research has reliably documented the involvement of the anterior regions of the temporal lobe (20, 22), left inferior parietal lobe (12, 20), and left inferior frontal cortices (13, 14).”

“In addition, each participant completed a spatial working memory task (Fedorenko et al., 2011), used in some control analyses to characterize brain regions sensitive to sentence scrambling, as described below.”

“Previous studies have shown that a subset of areas within prefrontal and lateral temporal cortex are sensitive to linguistic con- tent, but not to difficulty of working memory tasks (Fedorenko et al., 2011).”

“1244 0022-3077/19 Copyright © 2019 the American Physiological Society www.jn.org Downloaded from www.physiology.org/journal/jn at (174.063.084.192) on April 5, 2019. memory, inhibitory control, and music (Fedorenko et al. 2011, 2012; Monti et al. 2012; see Fedorenko and Varley 2016 for a review).”

“Replicating prior work (Fedorenko et al., 2011), we find no response to the spatial WM task in the language fROIs.”

“, 2016) but, critically, shows virtually no engagement in non-linguistic tasks and is therefore language-selective (Fedorenko et al., 2011; for a review, see: Fedorenko and Varley, 2016).”

“As the left middle frontal gyrus has been implicated in verbal working memory (Fedorenko et al., 2011), this reduced recruitment in TLE may also reflect some insufficiency in verbal working memory.”

“Furthermore, language responses in the frontal lobe are known to vary in their anatomical location across individuals (Fedorenko et al., 2011).”

“To examine responses to sentences in the inferior frontal gyrus (IFG), we defined individual ROIs as top 5% of vertices responsive in the BWs TCs or VWs FF strings contrasts within a published IFG mask (Fedorenko et al., 2011).”

“The contrast between these conditions has been shown to reveal DMN (easy > hard) and MDN (hard > easy) (Fedorenko et al., 2011).”

“nonwords that might engage such non-linguistic processes, the identified regions exhibit robust language selectivity in their responses, showing little or no response to nonlinguistic tasks (Fedorenko et al., 2011; Fedorenko et al., 2012a; Pritchett et al., 2018; Ivanova et al., 2019; Jouravlev et al., 2019; For a rev…”

“Taken together, all these neuroimaging studies contribute to a better understanding of the neurological bases of language development across the life span [105], particularly the development of word recall as measured by verbal fluency and confrontation naming tasks.”

“2010) and has been found to show functional specificity for high-level linguistic processing, but not for a variety of non-linguistic functions including arithmetic, cognitive control, music and working memory (Fedorenko et al. 2011).”

“46 Discussion We examined neural responses to cognitive demands associated with high-level language production—lexical access and phrase-structure building—in the fronto-temporal languageselective network (Fedorenko et al., 2011) using a robust precision fMRI approach.”

“…approach, using half of the data of each subject (blocks 1–3) for fROI definition and the other half (blocks 4–6) for response estimation (cf. Fedorenko et al. 2011, 2013). fROI definition followed the GSS procedure (Julian et al. 2012): For each fROI type, subject-specific activation maps…”

“…paragraph reading rather than single-word reading, the words have typically been presented one at a time in serial visual presentation (SVP, e.g., Martin-Loeches et al., 2008; Fedorenko et al., 2011; Pallier et al., 2011), with participants often asked to do a secondary task such as probe matching.”

“26; see Fedorenko and Blank 9 for a review).”

“…et al., 2006) and in music (Koelsch et al., 2009; Schulze et al., 2011; Williamson et al., 2010), and is associated with the inferior frontal regions that are implicated in both domains (Koelsch et al., 2009; Schulze, Zysset, Mueller, Friederici, & Koelsch, 2011; but see Fedorenko et al., 2011).”

“Studies of math and logic, domains that also make extensive use of symbols, show that those domains do not rely on the language system (Amalric and Dehaene, 2019; Cohen et al., 2000; Fedorenko et al., 2011; Monti et al., 2009; Monti et al., 2012; Pinel and Dehaene, 2010; Varley et al., 2005), a conclusion consistent w…”

“Fiebach and Schubotz, 2006; Gelfand and Bookheimer, 2003; Koelsch, 2005; Patel, 2003; Tettamanti and Weniger, 2006) though to date this is not unequivocally proven (Fedorenko et al., 2011; Rogalsky et al., 2011).”

“Recent research also suggests that cortical regions sensitive to language can be functionally defined in individual adults (Fedorenko et al., 2010, 2011).”

“While these domains appear to be neurobiologically distinct in adults (Fedorenko et al., 2011), it remains an open question whether language and EF rely on distinct neural processes in childhood, or if these become differentiated throughout development.”

“…comprehension, we examine two candidate brain networks, each functionally localized in individual participants: the language-selective network (Fedorenko et al., 2011) and the domain-general multiple-demand (MD) network implicated in executive functions, including working memory (Duncan et…”

“…they together fit well with the language network ( Figure 2B , middle) (Friederici, 2011), which shows selective activation by sentences in contrast to nonword lists and not by multiple non-linguistic tasks, such as, arithmetic, working memory, cognitive control, or music (Fedorenko et al., 2011).”

“…math and language were compared to each other, the networks observed for each domain are similar to those previously identified with other control conditions (e.g. lists of non-words for language and hard vs. easy contrast in a logic task) (e.g. Coetzee and Monti, 2018 ; Fedorenko et al., 2011 ).”

“In the case of MFG, this is consistent with prior work implicating MFG in both language comprehension and more general cognitive demand (e.g. working memory; Fedorenko et al., 2011; Mineroff et al., 2018).”

“The word-level semantic model predicts cortical voxels across regions in the frontal, parietal, and temporal lobes beyond core language-specific regions(Fedorenko et al., 2011, 2013).”

“Diverse nonlinguistic inputs and tasks elicit little or no response in the language areas in spite of strongly activating other brain areas (e.g., see Monti et al. 2009, 2012; Fedorenko et al. 2011; Ivanova et al. 2020; Chen et al. 2023).”

“…but not to mental arithmetic, music perception, executive function tasks, action/gesture perception, or computer programming (Amalric & Dehaene, 2019; Fedorenko et al., 2011; Ivanova et al., 2020; Jouravlev et al., 2019; Liu et al., 2020; Monti et al., 2009, 2012; Pritchett et al., 2018).”

“processing, aware of the fact that multiple (more than 10) brain areas may contribute to language processing in adults, including the left frontal lobe, left temporal/parietal lobes, right temporal lobe, cerebellum hippocampus basal ganglia (Fedorenko et al. 2011; Deniz Can et al. 2013), and especially the prefrontal…”