Hemispheric roles in the perception of speech prosody

“Dorsolateral prefrontal cortex, including BA 46 and BA 9, is involved in controlling attentional demands of tasks and maintaining information in working memory (Corbetta and Shulman, 2002; Knight et al., 1999; MacDonald et al., 2000; Mesulam, 1981).”

“This languagespecific effect can be understood from the conceptualization of the IPL as part of an auditory-motor integration circuit in speech perception (Hickok and Poeppel, 2000; Wise et al., 2001).”

“In this context, understanding the functional anatomy of language cannot be deduced from a single experiment; rather, it requires the integration of results from multiple experiments using multiple techniques.”

“Together, the studies suggest an involvement of the RH for the processing of intonational (pitch) information during sentence processing, but, in addition, indicate that the actual lateralization partly depends on task demands (90, 205) and on the presence of concurrent segmental information (30, 68).”

“In adults, functional MRI (fMRI) revealed that suprasegmental information is processed predominantly in the right hemisphere, again involving inferior frontal and temporal regions (21, 22).”

“In the presence of a left-hemisphere dysfunction, the right hemisphere assumes left-hemisphere tasks at which it is intrinsically less competent [ Gandour et al., 2003, 2004 ], resulting in overactivation.”

“A cross-language (Chinese and English) fMRI study ( Gandour et al. 2004 ) was conducted to examine brain activity elicited by selective attention to Mandarin tone and intonation (statement versus question), as presented in three- and one-syllable utterance pairs.”

“It is particularly in agreement with the concept of spectral fine-tuning of the right auditory cortex and its importance in tracking and processing pitch contours in speech (Kotz et al., 2003; Gandour et al., 2004) or the specific role for pitch in tone languages (for review, see Zatorre and Gandour, 2008).”

“Although syntactic and semantic processes are supported primarily by the left-hemisphere, the processing of prosody is primarily a function of the right-hemisphere (Gandour et al., 2004; Pell et al., 2006; Friederici et al., 2002).”

“Keywords: asymmetry, fMRI, lateralization, spectral, temporal”

“…in lexical tone perception; specifically, the right hemisphere is involved in auditory/acoustic processing, whereas the left hemisphere is responsive to linguisic/phonological functions (Gandour et al., 2000, 2002, 2004; Gandour, 2006; Hsieh et al., 2001; Wang et al., 2003; Wong et al., 2004).”

“…emotional prosody, most researchers regard right-hemisphere structures as part of a distributed and bilateral neural system engaged during the processing of emotionally laden speech (Adolphs, Damasio, & Tranel, 2002; Gandour et al., 2004; Kotz et al., 2003; Sidtis & Van Lancker Sidtis, 2003).”

“The left-dominant responses to listening sentences or words are only observed for native listeners in contrast to the bilateral or inconsistent activations for non-native listeners (Bottini et al., 1994; Perani et al., 1996; Dehaene et al., 1997; Gandour et al., 2004).”

“…facilitate and disambiguate speech comprehension, especially under acoustically compromised conditions, amongst them talker familiarity (e.g., Eisner and McQueen, 2005); prior experience and knowledge (e.g., Hannemann et al., 2007); and prosodic and emotional cues (e.g., Gandour et al., 2004).”

“The neural implementation of prosodic information, however, is less straightforward (Friederici and Alter, 2004; Gandour et al., 2004).”

“A right hemispheric bias for prosody processing has been observed in adults (Friederici & Alter, 2004; Gandour et al., 2004; Meyer, Alter, Friederici, Lohmann, & Von Cramon, 2002; Poeppel, 2003; Zatorre & Belin, 2001), children (Wartenburger et al.”

“Left dominance may be observed if the prosodic cue used is lexically or morphosyntactically relevant such as lexical tone in adults who speak a tonal language (Gandour et al., 2004; Kreitewolf, Friederici, & von Kriegstein, 2014; Sato, Sogabe, & Mazuka, 2007, 2010).”

“…bilaterally [Friederici et al., 2003; Kotz et al., 2002; Rossell et al., 2001], prosodic processes mainly rely on the right superior temporal gyrus (STG) [Gandour et al., 2004; Meyer et al., 2002; Zatorre et al., 1992], and syntactic processes involve different regions in the left temporal cortex.”

“Activation in the right IFG has been found during pitch processing in both nonlinguistic (Liu et al., 2006; Hsieh et al., 2001; Humphries, Willard, Buchsbaum, & Hickok, 2001) and speech contexts (Wildgruber, Ackermann, Kreifelts, & Ethofer, 2006; Gandour et al., 2004; Meyer et al., 2002).”

“…(Berckmoes & Vingerhoets, 2004; Eisenmajer et al., 1996), while others have argued that prosody that serves a linguistic function (e.g., lexical stress, phrasing, speech acts) involves the left hemispheric structures (Gandour et al., 2004; Wildgruber, Pihan, Ackerman, Erb, & Grodd, 2002).”

“The right temporal lobe activation has been widely observed during the phonological processing for Chinese subjects (29, 30), whereas the right anterior temporal pole was anatomically related to speakers of Chinese (23).”

“Steinhauer, K., Alter, K., Friederici, A.D., 1999.”

“These considerations are supported by findings from neuropsychology that if segmental information is included in the signal, the left hemisphere is additionally recruited (Friederici & Alter, 2004; Gandour et al., 2004; Poeppel, 2003; Perkins, Baran, & Gandour, 1996).”

“This suggests that the actual lateralization of prosodic processing partly depends on task demands (Plante et al., 2002; Gandour et al., 2004; Kotz et al., 2006) and on the presence of concurrent segmental information (Van Lancker and Sidtis, 1992; Baum and Pell, 1999; Friederici and Alter, 2004; Pannekamp et al.”

“two hemispheres via the corpus callosum [Gandour et al., 2004].”

“For example, Gandour et al. (2004) found that pitch in tonal languages was processed in left-lateralized areas when associated with semantic meaning (in native tonal language speakers) and rightlateralized areas when analyzed by lower-level acoustic/auditory processes (in English speakers that were…”

“Thereby the present observations extend previous fMRI findings which have associated these areas with the prelexical processing of phonetic cues and features of phonological significance in the perceiver’s native language (Jäncke et al., 2002; Jacquemot et al., 2003; Gandour et al., 2004).”

“Our data show that compared with acoustic variations, phonological variations elicited less activation in the core auditory region, exactly the right lateral HG, which has been suggested to function as a ‘‘pitch center’’ [30,31] that is specifically responsive to pitch sequences in speech or non-speech context [21,24,…”

“All rights reserved. doi:10.1016/j.bandl.2006.10.003 cues in speech is strong (Baum & Pell, 1999; Gandour et al., 2004).”

“…ECD solutions) without losing the subject’s data entirely: other valid observations from the subject will be used in the model nevertheless and will contribute to the enhanced power of a larger subject sample (Wolfinger, 1997; Carrier et al., 2001; Gandour et al., 2004; Frost et al., 2004).]”

“…and fMRI evidence has shown that processing lexical tones is left-lateralized in native Mandarin speakers, especially for pitch contour information, while processing acoustic tones remains right-lateralized regardless of listeners' language background (Gandour et al., 2004; Tong et al., 2005).”

“Processing of linguistic prosody, as it possibly occurred in the German group, in contrast has usually been attributed to right-hemispheric brain areas (Gandour et al., 2003, 2004; see also Friederici and Alter, 2004).”

“Neuroimaging studies have also shown differences in Chinese compared with English speakers in the oral language network when processing tonal information [Gandour et al., 2003, 2004; Hsieh et al., 2001; Klein et al., 2001].”

“According to Gandour et al. (2004), hemispheric lateralization for speech prosody processing is based on both the high-level linguistic representations and the low-level encoding of acoustic cues.”

“By using pseudowords (Gandour et al., 2002), short unstructured lists (Gandour et al., 2003b; Hsieh et al., 2001), and 3-syllable meaningless utterances (Gandour et al., 2003a, 2004; cf. syntactic speech, Meyer et al., 2002), we eliminated or minimized conceptual – semantic processing.”

“In fMRI studies on auditory degraded speech perception, the rSTS has shown to be sensitive to spectral fine-tuning (Scott, 2000; Zatorre, Belin, & Penhune, 2002) and pitch contours (Gandour et al., 2004; Kotz et al., 2003).”

“At the same time, studies of linguistic processing in tonal languages (Gandour et al., 1998, 2004) and left hemisphere processing of visual sign languages by individuals with profound hearing impairment (Petitto et al., 2000; Finney et al., 2001) suggest that the left hemisphere may have a…”

“…lateralization patterns of lexical tone processing (e.g., Van Lancker and Fromkin, 1973; Baudoin-Chial, 1986; Hsieh et al., 2001; Wang et al., 2001; Gandour et al., 2002, 2004; Tervaniemia and Hugdahl, 2003; Luo et al., 2006; Zatorre and Gandour, 2008; Li et al., 2010; Krishnan et al., 2011; Jia…”

“But neural correlates of prosodic pitch have only been observed to be right lateralized within the context of certain tasks, and typically when lexical content is no longer identifiable (Baum & Pell, 1999; Gandour et al., 2004; Kotz et al., 2003; Kreitewolf et al., 2014; Meyer, Alter, & Friederici, 2003; Meyer et al.,…”

“The right pSTG and f-pSTS (Gandour et al., 2004; Wiethoff et al., 2008), together with the regions in the right PTe (Griffiths et al., 1999), might be involved in this suprasegmental and pitch decoding from auditory cues related to angry prosody.”

“This is consistent with previous work suggesting that linguistic prosody processing is supported by the functions of neither one specific area nor a single hemisphere (e.g., Gandour et al., 2004; Tong et al., 2005; Witteman et al., 2011).”

“This is in line with previous fMRI results indicating a left-localization of lexical pitch only when language processing beyond the auditory analysis is required [Gandour et al., 2004].”

“Right STG activation has also been found in Chinese speech processing (Xiao et al, 2005 ; Gandour et al, 2004 ; Wang, Sereno, Jongman, & Hirsch, 2003 ), and researchers have suggested its role in tonal pitch processing of the Chinese language (Zatorre & Gandour, 2008 ; Gandour et al, 2004 ).”

“Accordingly, if the speaker has no knowledge or experience with the language, then one does not expect to see a left laterality effect in this region when performing speech prosody tasks [Gandour et al., 2004; Tong et al., 2005].”

“…with naïve listeners of a language tending to show right-hemisphere lateralization when processing novel speech sounds (Hsieh et al., 2001; Gandour et al., 2002, 2004; Wong et al., 2004; Luo et al., 2006; Xu et al., 2006; Zatorre and Gandour, 2008; Myers et al., 2009; Friederici, 2011).”

“…of lexical tones, native Mandarin speakers exhibit a right-ear advantage for identifying lexical tones (Wang, Behne, Jongman, & Sereno, 2004; Wang, Jongman, & Sereno, 2001), a result consistent with the neuroimaging literature (Gandour et al., 2004; Wong, Parsons, Martinez, & Diehl, 2004).”

“Research has shown that different types of prosody are processed differently in the brain (e.g., Friederici & Alter, 2004; Gandour et al., 2004), so the identification of discernible aspects of prosodic performance may be useful in discovering brain-behaviours connections.”

“…left hemisphere (Binder et al., 1997; Belin et al., 2000; Binder et al., 2000; Belin and Zatorre, 2003; Obleser et al., 2006), and pitch, prosody, and emotional qualities in human vocals more heavily involving the right hemisphere (Zatorre et al., 1992; Buchanan et al., 2000; Gandour et al., 2004).”

“Moreover, task-related neural activity strongly overlaps during intonation and tone processing in fronto-parietal brain areas, albeit intonation processing induces bilateral activity and tone processing is more left-lateralised (Gandour, Dzemidzic, et al., 2003; Gandour, Wong, et al., 2003; Gandour et al., 2004).”

“Brain imaging experiments reported a right hemispheric activation for the processing of suprasegmental information (Gandour et al., 2004; Meyer, Alter, Friederici, Lohmann, & von Cramon, 2002).”

“…& Guion, 2004), native speakers (e.g., Wang et al., 2017; Xi et al., 2010; K. Yu et al., 2014; L. Yu et al., 2015), and nonnative listeners (e.g., Gandour et al., 2004; Klein, Zatorre, Milner, & Zhao, 2001; Wong, Parsons, Martinez, & Diehl, 2004), and we argue that L2 speech acquisition can be…”

“These additional areas of neural activation are consistent with findings in the literature regarding their role in the processing of intonation (Brown et al., 2004; Gandour et al., 2003, 2004; Hesling et al., 2005; Meyer et al., 2002, 2004; Plante et al., 2002).”

“…specifics of the task (Plante et al. 2002; Tong et al. 2005), the type of prosody involved (e.g., affective vs. linguistic, Wildgruber et al. 2004), and the degree of propositional information contained in the stimulus materials (Gandour et al. 2002, 2004; Hesling et al. 2005; Tong et al. 2005).”

“While the general functional differences between the two hemispheres have been established [Gandour et al., 2000, 2002, 2004; Tong et al., 2005; Wong et al., 2004], the precise mechanism underlying the functional asymmetry at the acoustic level remains a matter of debate.”

“Brain imaging studies have found activation in the left frontal lobe and inferior parietal lobe for tasks involving active discrimination between lexical word tones (Gandour et al., 2000, 2003, 2004; Klein, Zatorre, Milner, & Zhao, 2001).”

“Both types of information are processed simultaneously (Buchanan et al. 2000 ; Schirmer et al. 2002 ; Gandour et al. 2004; Tong et al. 2005), and may be incongruent in affective meaning, as in the case of irony, sarcasm, or indeed by just not saying what is meant.”

“, 1994; Ross and Monnot, 2008) dimensions, others have linked such domains predominantly to the LH (Van Lancker, 1980; Emmorey, 1987; Brådvik et al., 1991; Baum et al., 1997; Gandour et al., 2004).”

“These areas have been reported to be related to prosodic processes (Fujimaki et al., 2004; Gandour et al., 2004).”

“…et al., 2000), acoustic cue (pitch/duration; e.g. Van Lancker & Sidtis, 1992), stimulus type (syllable/word/sentence; e.g. Meyer et al., 2002; Gandour et al., 2004), as well as methodological factors including design (event-related/blocked; c.f. Kotz, Meyer, Paulmann, 2006) and…”

“…(e.g., Mandarin) developed this finding, observing an overall preference for prosodic cues in the right hemisphere for native and nonnative speakers, but a left hemisphere sensitivity to linguistically relevant tonal information in the native group only (Tong et al., 2005; Gandour et al., 2004).”

“In light of these views on brain lateralization, Gandour et al. have put forward a comprehensive hypothesis on this issue (Gandour et al., 2000, 2004; Gandour, 2006).”

“…lie in music-selective neural circuitry in the right IFG as in non-tonal language speakers, or overlap with the neural circuitries of lexical tone processing (cf. Tong et al. 2005; Luo et al. 2006; Gandour et al. 2002; 2004; Wong et al. 2004; Li et al. 2010; Gu et al. 2013; Zhang et al. 2016).”

“language comprehension tasks evoke more semantic processing (Gandour, Dzemidzic, et al., 2003; Gandour, Wong, et al., 2003; Gandour, Wong, & Hutchins, 1998; Gandour et al., 2004).”

“…and Linguistics Compass ª 2011 Blackwell Publishing Ltd speech sounds at the level of the brainstem as well as differences in hemispheric lateralisation (e.g. Chandrasekaran et al. 2007; Gandour et al. 1998, 2000, 2002, 2004; Krishnan and Gandour 2009; Krishnan et al. 2005; Xu et al. 2006).”

“In this context our observed right hemispheric lateralization of EFRs in children is of interest because it has long been held that prosodic information is preferentially processed in the right hemisphere (Friederici and Alter, 2004; Gandour et al., 2004; Homae et al., 2006; Plante et al., 2006; Wartenburger et al., 2…”