Listening and Reading Part of Brain That Controls

Psychol Neurosci. Author manuscript; bachelor in PMC 2011 Apr 25.

Published in final edited class as:

Psychol Neurosci. 2009; 2(ii): 111–123.

PMCID: PMC3081613

NIHMSID: NIHMS181522

Brain activation for reading and listening comprehension: An fMRI study of modality effects and individual differences in language comprehension

Abstract

The study compared the brain activation patterns associated with the comprehension of written and spoken Portuguese sentences. An fMRI written report measured brain activity while participants read and listened to sentences virtually general world cognition. Participants had to decide if the sentences were true or simulated. To mirror the transient nature of spoken sentences, visual input was presented in rapid serial visual presentation format. The results showed a common core of amodal left inferior frontal and middle temporal gyri activation, as well as modality specific brain activation associated with listening and reading comprehension. Reading comprehension was associated with more left-lateralized activation and with left inferior occipital cortex (including fusiform gyrus) activation. Listening comprehension was associated with extensive bilateral temporal cortex activation and more overall activation of the whole cortex. Results besides showed individual differences in encephalon activation for reading comprehension. Readers with lower working retentiveness capacity showed more activation of correct-hemisphere areas (spillover of activation) and more activation in the prefrontal cortex, potentially associated with more demand placed on executive command processes. Readers with college working memory capacity showed more activation in a frontal-posterior network of areas (left angular and precentral gyri, and right inferior frontal gyrus). The activation of this network may be associated with phonological rehearsal of linguistic information when reading text presented in rapid serial visual format. The study demonstrates the modality fingerprints for language comprehension and indicates how low- and high working retentivity chapters readers deal with reading text presented in serial format.

Keywords: fMRI, linguistic communication comprehension, reading span

Introduction

Linguistic information tin be conveyed in the form of speech and written text, but information technology is the content of the message that is ultimately essential for the higher-level processes in language comprehension, such as making inferences and associations between text information and knowledge near the world. The goal of this report was to investigate the brain activation for listening and reading comprehension processes and the effects of modality of language input on brain activation for linguistic communication comprehension. The report also aimed to investigate individual differences in encephalon activation for individuals with high or low linguistic communication processing chapters, every bit indexed past the Daneman and Carpenter (1980) reading span test.

Practiced readers tend to exist adept listeners, and good listeners tend to exist good readers. Behavioral studies have shown that listening and reading comprehension are 2 closely-related skills. Every bit schooling increases, and so does the strength of the correlation between reading and listening comprehension performance (Just & Carpenter, 1987). Skilled readers retrieve phonological data faster and more automatically than less skilled readers (Berth, Perfetti, & MacWhinney, 1999; Berth, Perfetti, MacWhinney, & Chase, 2000). Successful reading relies on an interaction between decoding linguistic visual input and accessing phonological data.

Models of language comprehension draw higher-level cognitive processes of text comprehension (inference-making and semantic access, for instance) every bit amodal processes. This ways that higher-order cerebral processes draw on the manipulation of text data in abstruse form (Stonemason & Just, 2006). Kintsch (1998) proposed that discourse processing is based on the semantic construction of text, which is broken down into amodal units of meaning called propositions. According to the model, comprehension thrives on the integration of this amodal, propositional information. Booth et al.'s (2002a; 2002b) model of language comprehension, which is based on evidence from brain imaging studies, postulates that although auditory and visual word form processing involves distinct cortical areas, semantic processing is independent of input modality. Semantic processes are associated with the aforementioned network of encephalon activation whether the input is visual or auditory.

Encephalon imaging studies of language comprehension have demonstrated that there is a comparable network of areas of the encephalon activated in higher-society cerebral processes of reading and listening comprehension. There is a high similarity in cortical areas recruited for listening and reading comprehension processes at the word, sentence, and soapbox level (Jobard, Vigneau, Mazoyer, & Tzourio-Mazoyer, 2007). Encephalon imaging studies convincingly point the same direction of behavioral studies, indicating that the higher-order processes of listening and reading comprehension are intertwined, rather than divide. On the one hand, there are differential cortical areas recruited past modality-specific processes, such every bit the processing of word course by the visual give-and-take form surface area (Cohen et al., 2002). Still there are comparable areas recruited by amodal processes such as inference-making and other higher-level cognitive processes (Berth et al., 2002a; Carpentier et al., 2001; Constable et al., 2004; Jobard, Crivello, & Tzourio-Mazoyer, 2003; Michael, Keller, Carpenter, & Just, 2001;). Encephalon imaging studies of complexity furnishings on encephalon activation take also shown comparable modulation of brain activity by job complication in both listening and reading comprehension tasks (Carpentier et al., 2001; Constable et al., 2004; Merely, Carpenter, Keller, Boil, & Thulborn, 1996; Keller, Carpenter, & Merely, 2001; Michael et al., 2001).

The amodal network of cortical areas involved in language processing, called the language network, centralizes higher-order language comprehension processes. The modality-specific, or dedicated, primary areas are involved in lower-level processing (Jobard et al., 2007; Mesulam, 1998). Ane of the amodal centers of language processing in the encephalon, or amodal computation node, is the left inferior frontal gyrus (LIFG). In plough, the master areas, or modality-specific areas, include the primary auditory and visual cortices. Booth and colleagues (2002a) reported modality-independent activation in semantic processing over the LIFG and left middle temporal gyrus, but modality-specific activation of the fusiform gyrus (written words) and of the superior temporal gyrus (spoken words). Brain imaging studies accept mapped specific linguistic integrative and input-to-meaning functions onto these known cortical structures: (1) the LIFG, for amodal processes, integration of textual information, and response to language chore complexity; (2) the auditory cortex, for the primary processing of speech and the decoding of acoustic features in speech (prosody, for example); and (3) the visual cortex and the fusiform gyrus, for the processing of visual input to meaning.

The processing of abstract, amodal information in language

The well-known left-hemisphere linguistic communication network implicated in the processing of language and soapbox includes the LIFG, the superior and centre temporal gyri, the inferior temporal gyrus, and the angular gyrus (Bookheimer, 2002). The LIFG is implicated in a variety of linguistic communication tasks, which include production and comprehension (Schlosser, Aoyagi, Fullbright, Gore, & McCarthy, 1998) and retrieval of meaning (Wise & Price, 2006). Lesions to the portion of the LIFG known as Broca's Expanse (the LIFG pars opercularis, or BA 44, co-ordinate to Broca's original definition, but currently including a larger portion of the IFG (Bookheimer, 2002)) are associated with language production disorders, such equally articulatory and spoken communication aphasias and naming deficits (Bookheimer, 2002; Obler & Gjerlow, 1999).

The LIFG is implicated in several specialized language tasks that require some level of manipulation and integration of amodal data. LIFG activation has been associated with syntactic and semantic processing (Keller et al., 2001), increasing comprehension workload (Constable et al., 2004; Just et al., 1996; Michael et al., 2001), increasing complexity of language units (word, judgement, text) (Jobard et al., 2007), and reading tongue-twister sentences (Keller, Carpenter, & Simply, 2003). The LIFG has been described as part of a network of cortical areas activated in the integration of information into the reader's understanding of text. This is similar to what models of comprehension call the mental (state of affairs) model (Mason & Merely, 2006).

The decoding of speech

The primary auditory cortex corresponds anatomically to BAs 22, 21, and 20. Heschl's gyrus (BA 22) is well-known for its association with general processing of auditory stimuli; likewise, the posterior portion of the superior temporal gyrus, or STG (BA 42), and the superior temporal sulcus (BAs 22 and 42) are also associated with general processing of auditory input (Cabeza & Nyberg, 2000; Jobard et al., 2003; Schlosser et al., 1998). A number of functional imaging studies of comprehension have reported differential activation of STG in listening comprehension relative to reading comprehension (Carpentier et al., 2001; Lawman et al., 2004; Jobard et al., 2007; Michael et al., 2001).

The decoding of written text

The primary visual cortex decodes visual information from printed words, sentences, and text. Neuroimaging studies of lexical tasks take also revealed the importance of the fusiform gyrus for lexical processing in writing systems of several languages, including alphabets and logographs (Bolger, Perfetti, & Schneider, 2005; Cohen et al., 2000; 2002; Jobard et al., 2007; Tan, Laird, Li, & Fox, 2005). In terms of subspecialization of cortical function, in that location is a portion of the fusiform gyrus referred to as the visual word form area (VWFA) that is paramount for the lexical processes that bridge the gap betwixt linguistic visual input and speech representations.

The VWFA is largely implicated in mapping visual information to meaning or retrieval of significant (Cohen et al., 2000; 2002; Cohen & Dehaene, 2004), specially in reading ideograms (Bolger et al., 2005). It processes information of fine-grained visual grade such equally (but non exclusively) information required for discriminating betwixt words and for combining visual and verbal linguistic information (Devlin, Jamison, Gonnerman, & Mathews, 2006; Vigneau, Jobard, Mazoer, & Tzourio-Mazoer, 2005). Lesions to the VWFA are associated with impairments in oral reading and oral naming tasks (Hills et al., 2005).

Nonetheless, based on a metanalysis of 35 neuroimaging studies, Jobard and colleagues (2003) argued that the concept of a VWFA invites more refined investigation and that neuroimaging studies have withal to consistently corroborate the concept of a written discussion lexicon in the brain. It is possible that VWFA activation is more than specific to reading words in deep orthographies.

The unlike levels of complexity of the mapping between the printed word and the sounding out of that word may be associated with unlike cognitive processes, and may be underpinned by different networks of cortical activation. For example, differences in encephalon activation have been reported for Italian (transparent orthography) and English language (nontransparent, deep orthography) reading comprehension. For Italian reading comprehension at that place was a language-specific left superior temporal activation, whereas for English reading there was a language-specific left inferior posterior temporal activation (Paulesu et al., 2000). Portuguese, in a putative continuum of orthographic transparency, is less transparent than Italian and Spanish, only it is more transparent than French and, specially, English (Seymour, Aro, & Erskine, 2003).

For shallow orthographies, the development of reading skill may be more than strongly associated with phonological processing abilities. For deep orthographies, the development of reading skill may be more strongly associated with the ability to combine visual and phonological processing abilities. Paulesu and colleagues (2001) found that dyslexic readers in Italian (transparent orthography) perform better in reading tasks than dyslexics who read English language and French. The written report shows an event of cultural variety on language processing. Of course, independently of the linguistic communication, dyslexic readers always performed worse than normal, control readers. A cross-linguistic comparing among dissimilar orthographies showed that children learning shallow orthographies go accurate and fluent in reading words sooner than children who must learn to read French, Danish, and particularly English (Seymour et al., 2003). An investigation of children learning to read in Portuguese showed a strong correlation of higher-span reading skills with phonological processing abilities and a weak correlation with visual processing abilities (Capovilla, Capovilla, & Suiter, 2004a; Capovilla & Capovilla, 2004b).

Individual differences in reading comprehension skill, right-hemisphere spillover of activation, and reading text in rapid serial visual presentation format

Reading comprehension is a complex, higher-level cerebral process in which there are systematic individual differences in skill and performance. For instance, expert readers are faster and more than accurate at comprehension of syntactically complex sentences than are poor readers (Just & Carpenter, 1987). One of the questions that brain imaging studies have attempted to reply is which individual differences in brain functioning underpin private differences in reading skill.

Individual differences in reading comprehension are likely to be associated with a quantifiable measure of consumption of encephalon resources during task performance. Resources consumption can be measured by the amount of brain activation in unlike areas of the brain. In a study of judgement comprehension with varying lexical and syntactic difficulty, Prat, Keller and But (2007) constitute greater correct-hemisphere activation in less-skilled participants than in skilled participants. In other words, to perform the more difficult tasks, less-skilled participants required boosted recruitment of encephalon activation in correct-hemisphere homologues of the areas normally activated during linguistic communication comprehension (see Prat & Only, 2008 for a word of the brain bases of private differences in language comprehension).

In the present study the reading task was a relatively easy one for higher-level students. We did not modulate sentence syntactic or lexical difficulty. However, because the reading chore was presented in a novel, unconventional class (serial presentation), we expected to discover prove of unlike brain activation between loftier and depression capacity readers for dealing with the reading task. The visual stimuli were presented in rapid serial visual presentation format (RSVP), where words appear on the screen 1 at a fourth dimension. RSVP is an anarchistic form of rapid reading developed by Forster (1970). RSVP differs from normal reading because the duration of gaze on each give-and-take is non under the command of the reader, words cannot be skipped, and the words that have already been read cannot be read once again (no backtracking).

Method

Participants

Twelve right-handed speakers of Portuguese as a first language (eight males), mean age 29.9 years (SD = five.74; range 20-xl years), were recruited for the study. Participants were highly educated: 11 were enrolled in graduate schoolhouse programs and 1 was in the senior twelvemonth of college. All participants were financially compensated for the practice session and the fMRI data collection. Each participant gave signed informed consent canonical by the University of Pittsburgh and Carnegie Mellon University Institutional Review Boards.

Stimulus

The stimuli consisted of 24 statements (12 in print, 12 spoken, all in Portuguese) about full general world knowledge, for case: O Everest é conhecido como a montanha mais alta do mundo east está localizado no Nepal ("The Everest is known every bit the highest mount in the world and is located in Nepal") (for complete transcription of the stimuli used in the study, run into Appendix A). The sentences were controlled for topic, length, and duration of play. The auditory and visual sentences were constrained to 12 to 16 words in length. Both auditory and visual stimuli were presented for a total of six seconds. Visual stimuli were farther constrained to 65 to 85 characters, to maintain consistency in give-and-take presentation speed beyond sentences. Several other precautionary steps were taken: Auditory sentences were digitally recorded and (1) spoken with little or no prosody; (2) digitally edited for comparable volume (Goldwave v5.06, Goldwave inc.); and (three) edited for onset at fourth dimension nix and completion within milliseconds of fourth dimension = 6 sec. Visual sentences had no punctuation other than the final period.

Visual sentences were presented ane word at a time to control for visual processing rate, which would be somewhat up to the participant if sentences were presented in their entirety on the screen. Both auditory and visual sentences had to exist candy sequentially and inside the aforementioned amount of fourth dimension (six sec). Hence, the reading stimuli did not let for backtracking.

fMRI scanner and acquisition parameters

Imaging was done on a Siemens Allegra 3.0 Tesla scanner used in conjunction with a commercial birdcage, quadrature-drive radio-frequency whole-head coil. Sixteen oblique-axial images were selected to maximize coverage of the unabridged cortex. The images were collected using an EPI conquering sequence, with TR = 1000 ms, TE = 30 ms, flip angle = 60°, and a 64 × 64 acquisition matrix with a voxel size of 3.135-mm × 3.125-mm × 5-mm with a one-mm gap. The book scan was constructed from 160 3DMPRAGE oblique-axial images that were collected with TR = 2000 ms, TE = 3.34 ms, 7° flip-angle, and a 256 × 256 FOV, resulting in ane-mm × 1-mm × one-mm voxels.

Process

Participants were positioned in the scanner bed as comfortably as possible, with their head strapped to the caput whorl to help avoid excessive move. An angled mirror was adjusted in front of the eyes to reflect the visual stimuli projected onto a rear-projection screen in the bore of the magnet. Audio stimuli were transmitted via special headphones designed to fit the ears snugly and reduce scanner noise interference. Visual stimuli were displayed word-past-discussion on the center of the screen using rapid serial visual presentation (RSVP). RSVP word presentation charge per unit for each word for each sentence was calculated separately for each stimulus sentence. Word presentation charge per unit for each sentence was calculated with a formula that incorporated each word length times fifty milliseconds, then the sum of all these multiplications was subtracted from half dozen seconds, and the result then divided by the number of words in the sentence. The result of this subtraction-and-partition is a variable intercept, which was added to the number obtained in the multiplication of word length times fifty milliseconds. This addition gave the total fourth dimension a word was to be presented on the screen.

Sentences were displayed as a unmarried status (Portuguese listening solitary or Portuguese reading alone) in two blocks of six sentences for each condition. In between each cake of vi sentences, an "X" was presented on the centre of the screen (fixation condition). During fixation, participants were instructed to clear their mind. To ensure that participants were reading and listening to the sentences, there were True or Fake (T/F) questions after each sentence. Xx-five per centum of sentences were false.

Behavioral information assay

Participants responded to sentences presented visually or auditorily using mice buttons. Anatomical mapping of response fingers was used such that the index finger on the left hand was used to reply to true statements, and the index finger on the right hand to respond to imitation statements. The response options were rear projected onto a screen 20 cm from the participant. True or false responses and response times were nerveless in the scanner as participants pressed the mice buttons. Response times were recorded by the Coglab experimental software.

fMRI information assay

Distribution of activation

Information were analyzed using SPM2 (Wellcome Section of Cerebral Neurology, Academy College, London). Images were corrected for slice acquisition timing, move-corrected, normalized to the Montreal Neurological Constitute (MNI) template, resampled to 2-mm × 2-mm × ii-mm voxels, and smoothed with an 8-mm Gaussian kernel to decrease spatial noise. Statistical analysis was performed on individual and grouping data past using the general linear model and Gaussian random field theory as implemented in SPM2 (Friston et al., 1995). Group analyses were performed using a random-furnishings model. Statistical maps were superimposed on normalized T1-weighted images. Automated anatomical labeling (AAL) (Tzourio-Mazoyer et al., 2002), as implemented in the SPM2 software, was employed to name activation cluster centroids and adjacent areas of activation.

Correlation with reading span scores

The reading bridge scores for each participant were correlated with the individual images for each participant'southward contrast betwixt reading comprehension and fixation, as implemented in SPM2. The reading span scores were also correlated with a measure of recruitment of right-hemisphere areas: the ratio of total voxels recruited in left hemisphere in relation to voxels recruited in the right hemisphere for reading comprehension. The participants' reading span scores ranged from two to 5 (M = 2.79; SD = 0.86).

Results

Behavioral results

There were no significant differences between the results for Portuguese listening and reading comprehension accurateness (listening comprehension 1000 = 0.90, SD = 0.ten; reading comprehension M = 0.90, SD = 0.12) or response times (listening comprehension Grand = 907 ms, SD = 329; reading comprehension M = 1030 ms, SD = 323). Every bit expected, input modality did not take an upshot on speed of processing or accuracy of sentence comprehension.

fMRI results

Mutual network of activation for listening and reading comprehension

Listening and reading comprehension activated a common left-lateralized network of areas in left inferior frontal gyrus (LIFG) and left middle temporal gyrus (LMTG). These areas are part of a modality-independent network of activation associated with language comprehension (Berth el al., 2002a; Constable et al., 2004). Right junior frontal gyrus was too activated for both reading and listening comprehension. The mutual areas of activation are shown in white in Figure one. Listening comprehension showed big clusters of activation in bilateral (posterior and inductive) superior temporal and centre temporal gyri. Bilateral posterior superior temporal lobe activation has been consistently associated with auditory comprehension in brain imaging studies of language comprehension (Lawman et al., 2004; Jobard et al., 2007; Michael et al., 2001). In turn, reading comprehension showed activation of bilateral fusiform gyri. The left fusiform gyrus is an expanse associated with the processing of written words.

Cortical areas activated for listening and reading comprehension (p < 0.001, uncorrected; T = 4.02; extent threshold = twenty voxels; (a) illustrates the overlap of common subsets of cortical areas of activation for listening comprehension and/or reading comprehension contrasted with fixation (reddish = listening only; light-green = reading merely; white = listening and reading), and shows the areas of activation (b) only in listening comprehension and (c) only in reading comprehension)

Modality fingerprints in brain activation: listening > reading and more than overall activation of the brain for listening comprehension

The dissimilarity between listening and reading comprehension showed more than activation for listening comprehension in bilateral superior temporal gyri. There was also more activation for listening comprehension in bilateral center temporal gyri, correct angular gyrus, and right insula. Listening comprehension was also associated with more than overall activation of the whole brain when compared to reading comprehension (listening > fixation: 22,682 total voxels activated; reading > fixation: 3,579 total voxels for the contrasts beyond all participants, run into Tables one and ​ two).

Table 1

Activation for listening comprehension contrasted with fixation

Location	Voxels	T-value	MNI
			ten	y	z
Temporal lobe
L mid temp gyrus	20,497	25.92	-62	-22	-six
R sup temp gyrus
L sup temp gyrus
L lingual gyrus
R mid temp gyrus
Fifty + R calcarine
50 + R thalamus
Frontal lobe
Fifty precentral gyrus	244	ix.97	-36	-two	58
L mid frontal gyrus
L supp motor area	749	7.09	-2	4	62
R supp motor expanse
L sup frontal gyrus
L inf frontal gyrus	534	6.86	-44	iv	26
Fifty precentral gyrus
R inf frontal gyrus	260	v.69	46	18	26
R inf frontal gyrus
R mid frontal gyrus
50 sup frontal gyrus	123	5.66	-12	42	50
L med frontal gyrus
R precentral gyrus	21	4.88	46	-14	lx
Occipital lobe
R sup occipital gyrus	31	6.35	28	-76	xviii
Fifty fusiform gyrus	119	6.02	-48	-58	-20
L inf temp gyrus
R caudate	27	5.91	10	6	22
50 caudate	37	5.73	-eighteen	30	0
50 inf occipital gyrus	40	v.07	-40	-76	-12

Table 2

Activation for reading comprehension contrasted with fixation

Location	Voxels	T-value	MNI
			ten	y	z
Frontal lobe
L inf frontal gyrus	628	x.41	-46	4	26
50 precentral gyrus
Fifty inf frontal gyrus	534	9.25	-44	30	-10
R inf frontal gyrus	210	7.xx	46	10	24
Fifty supp motor area	292	vi.61	-two	8	60
R inf frontal gyrus	136	five.95	34	26	-10
R insula
Fifty mid frontal gyrus	25	5.26	-44	8	54
L precentral
50 insula	58	five.18	-30	26	-eight
L inf frontal gyrus
Occipital and temporal lobes
L caudate	thirty	vii.62	-8	6	12
L inf occipital gyrus	1,205	7.49	-34	-76	-x
50 fusiform gyrus
L mid temp gyrus
R inf occipital gyrus	212	half dozen.95	48	-74	-14
R fusiform gyrus
R inf temporal gyrus
R inf occipital gyrus	157	6.85	22	-96	-10
R lingual gyrus
L putamen	46	5.57	-22	-two	viii
Parietal lobe
50 sup parietal lobe	46	4.84	-28	-60	54
50 inf parietal lobe

Reading > listening and left-lateralized encephalon activation for reading comprehension

Reading comprehension showed more activation in the left inferior occipital lobe, including the left fusiform gyrus (Effigy 2, Table iii). Activation of the occipital lobe is associated with processing visual stimuli and is consistent with reading-specific activation establish in other studies (Lawman et al., 2004). In most participants, the brain activation for reading comprehension was left-lateralized. Eight participants had more activated voxels in the left hemisphere than in the right hemisphere for reading comprehension. The boilerplate ratio of left-to-right hemisphere voxels was two.5 (SE = 1.2) (boilerplate number of voxels activated in left hemisphere = 1864.5; SE = 680.5; average number of voxels activated in right hemisphere = 1718.ane, SE = 630.7).

Brain activation for the contrast between reading and listening comprehension (p < 0.001 uncorrected; T = 4.02; extent threshold = twenty voxels; blue ellipses highlight the bilateral center and superior temporal gyri activation for listening comprehension > reading comprehension; cherry-red ellipsis highlights the left inferior occipital lobe activation for reading comprehension > listening comprehension)

Table 3

Contrasts between listening and reading comprehension

	Voxels	T-value	MNI
Listen > read			10	y	z
Temporal lobe
R sup temp gyrus	14,129	fifteen.78	52	-14	2
Fifty calcarine
R lingual gyrus
R mid temp
L sup temp gyrus	3,397	14.67	-40	-34	8
L mid temp gyrus
L Heschl'southward gyrus
R mid temp gyrus	110	4.99	sixty	-52	14
R angular gyrus
Frontal lobe
R sup front end gyrus	24	7.55	36	-iv	62
R precentral gyrus	30	5.13	54	-10	46
R supp motor surface area	23	v.08	viii	0	62
Subcortical
Fifty hypothalamus	146	8.06	-two	0	-12
R thalamus	53	5.xix	18	-26	-4
Read > mind			x	y	z
50 inf occipital gyrus	65	5.36	-34	-90	-16
L lingual gyrus
L fusiform gyrus

The brain activation for reading comprehension was more left-lateralized than the encephalon activation for listening comprehension. Left-lateralized encephalon activeness for reading comprehension has been widely reported in studies of reading comprehension (Constable et al., 2004; Jobard et al., 2007; Simply et al., 1996; Michael et al., 2001). Reading comprehension had a significant number of active voxels in areas associated with visual processing. There was activation in clusters in the left junior occipital cortex and in the right inferior occipital lobe (Tabular array ii).

Private differences in encephalon activation for reading comprehension

The results evidence that the ratio of voxels recruited in the left hemisphere versus voxels recruited in the right hemisphere was positively correlated with reading span scores (r = .76; p < .01). This indicates that lower capacity readers had to recruit significantly more voxels in correct-hemisphere areas of the brain than higher capacity readers. This result corroborates previous findings of spillover of encephalon activation in less-skilled readers (Prat et al., 2007).

The correlation between reading span scores and brain activation for reading comprehension showed two characteristics of individual differences in brain activation for reading comprehension in rapid series format. Lower capacity readers (low spans) showed more than activation in the left middle frontal gyrus, an expanse associated with executive control and strategic processes. Higher capacity readers (high spans) showed more than activation in left angular, precentral and postcentral gyri, and right inferior frontal gyrus. The network of activation for improve readers may be associated with phonological rehearsal. The differences in correlation of reading ability and encephalon activation may be evidence of different strategies for reading comprehension in RSVP format betwixt higher and lower capacity readers (Effigy 3). At that place were no significant correlations between reading span scores and the brain activation for listening comprehension.

Correlation between encephalon activation for reading comprehension and reading span scores. (p < 0.001, extent threshold = 20 voxels; top: negative correlation with activation for reading comprehension: LMFG MNI: x = -34; y = 52; z = x, cluster size = 45 voxels (t = 7.01); bottom: positive correlation with activation for reading comprehension: L angular gyrus MNI: x = -34, y = -58, z = -18, cluster size = 21 voxels (t = 4.82); Fifty precentral + postcentral gyri MNI: 10 = -28, y = -32, z = 56, cluster size = 31 voxels (t = v.80); 50 postcentral gyrus + paracentral lobule MNI: x = -16, y = -34, z = sixty, cluster size = 41 voxels (t = vii.46); R middle + junior frontal gyri MNI: x = 42, y = 22, z = 32, cluster size = 23 voxels (t = 4.88))

Discussion

The results demonstrate the modality fingerprints for encephalon activation for listening and reading comprehension of Portuguese sentences and the brain activation associated with individual differences in reading text presented in serial format. The brain activation for listening and reading comprehension showed that processing of spoken communication and print resulted in differential activation in modality-sensitive areas. In addition, listening comprehension resulted in more overall activation over the whole brain. The extensive crimson-colored clusters of activation in Figure 1 illustrate the difference in overall brain activation for listening comprehension. The brain activation for reading comprehension was more left-lateralized. These results corroborate previous studies of listening and reading comprehension (Constable et al., 2004; Jobard et al., 2007; Michael et al., 2001).

It is interesting to compare the results from the nowadays study to studies of the influence of input modality in other languages. The present written report showed that the activation for reading comprehension was more left-lateralized, and listening comprehension, more than bilaterally distributed. The laterality difference between activation for comprehension of impress and speech corroborates studies that compared brain activation for listening and reading activation in other languages (Constable et al., 2004; Jobard et al., 2007; Michael et al., 2001). One explanation proposed for the departure in laterality of brain activity for reading and listening comprehension is developmental. Listening comprehension begins at a much earlier phase in life than reading comprehension. Consequently, an established left-laterality of language processing may influence a left-disposition of encephalon response to visual linguistic stimulus (Michael et al., 2001).

Other studies of modality furnishings on brain activation have reported differences in the location of LIFG activation for listening and reading comprehension. The difference in LIFG activation was modulated by job difficulty. The studies showed an increase in activation in pars triangularis (BA 45) associated with increasing difficulty of the comprehension tasks (Carpentier et al., 2001; Lawman et al., 2004; Michael et al., 2001). Still, the studies also showed more modality-specific (and non task-specific) activation in LIFG for reading relative to listening comprehension (Carpentier et al., 2001; Lawman et al., 2004). In the current report, LIFG activation for listening and for reading comprehension was identified in two overlapping clusters of activation. Our results did show a larger cluster of activation in LIFG for reading comprehension in the contrast with fixation, but the difference was non statistically significant in the group comparing between reading and listening comprehension. The interpretation of the differences in LIFG activation in other studies is that in that location may exist some degree of modal subspecialization in this surface area of the brain (Lawman et al., 2004). This subspecialization of LIFG activation was not found in the present study; however, task difficulty was not manipulated in the nowadays stimuli.

Modality fingerprints for listening comprehension

The modality-specific bilateral superior temporal cortex activation in listening comprehension, relative to reading comprehension, replicated previous findings of the bilaterality of activation for this process. Both the STG and MTG are well known for their association with early speech communication processing, and with spoken-discussion recognition tasks (Cabeza & Nyberg, 2000). Listening comprehension too showed more activation in right athwart gyrus. Left angular gyrus activation has been associated with phonology-to-orthography (and vice-versa) conversion in lexical tasks (Booth et. al, 2002b). Listening comprehension also showed more than activation than reading comprehension in the superior frontal gyrus (SFG). The activation of superior frontal areas of the brain has been associated with phonological processing without visual input (Katzir, Misra, & Poldrack, 2005).

Modality fingerprints for reading comprehension

The contrast betwixt reading comprehension and its auditory counterpart showed clear modality-specific activation. Reading comprehension activated more left junior occipital cortex, including the left fusiform gyrus. The fusiform gyrus is activated in word reading tasks in dissimilar writing systems (Bolger et al., 2005; Tan et al., 2005).

The activation for reading comprehension assorted with listening did not show more activation of the visual word form area (VWFA) for reading. The VWFA is a modality-specific expanse of the brain known for its response to written visual stimuli. The distance betwixt the summit of fusiform activation closest to the VWFA (located at MNI coordinates x = -42, y = -52, z = -22) was approximately 20.0 mm from the standard VWFA coordinates reported in a meta-assay of 16 brain imaging studies of discussion reading (MNI x = -44, y = -58, z = -fifteen) (Vigneau et al., 2005). This distance is to a higher place the ∼five.0 mm standard departure reported for the coordinates encompassing the visual word grade area (Cohen et al., 2000; 2002; Cohen & Dehaene, 2004). The left junior occipital activation found for reading comprehension extends slightly to posterior left fusiform gyrus, but it was more posterior than the activation usually reported for reading tasks (Bolger et al., 2005; Carpentier et al., 2001; Lawman et al., 2004; Tan et al., 2005).

Studies have shown that brain activation is sensitive to the consistency of the mapping between orthography and audio. Paulesu and colleagues (2000) showed that reading Italian (transparent, consistent spelling-to-audio rules) resulted in more activation of left superior temporal areas (phoneme processing), while reading English (deep, inconsistent spelling-to-audio rules) resulted in more than activation of left posterior junior temporal gyrus. Buchweitz, Mason, Hasegawa and Merely (2009) found that reading dissimilar writing systems inside the aforementioned language (Japanese) resulted in different brain activation for reading a logographic (kanji) and a syllabic (hiragana) writing system. Reading sentences in the logographic system (more visually complex ideograms) activated more occipitotemporal lobe areas, associated with visual processing. Reading sentences in the syllabic system activated more than areas associated with phonological processes. Information technology has been shown that the VWFA is more agile for existent words than for consonant strings, which indicates that it is an expanse that becomes attuned to the orthographic regularities that constrain alphabetic character combination (Cohen et al., 2002).

It is plausible that activation of the VWFA is more than prominent in deep orthographies such as English and French. More transparent orthographies, such equally Spanish, Italian, and Portuguese, accept a more compatible mapping of print to the sounds of the language. The same grouping of letters within these languages volition probable lead to the same pronunciation. In transparent orthographies reading can exist achieved by a regular grapheme-phoneme route (graphophonological route). However, in deep orthography languages, similar English and French, there are several letter combinations that volition read differently (for example, in English language, "int" in "pint" or "mint," or "eard" in "heard" or "beard"). To correctly read those words that do not follow the most usual spelling-to-sound rules, it is necessary to engage another road, called the direct or lexicosemantic route (Jobard et al., 2003). In a deep orthography, readers use the lexicosemantic route to build on the associations between arbitrary mappings of print to sound every bit they see new words (Jobard et al., 2003). The lexicosemantic route of reading is based on the associations betwixt word grade and meaning. Discussion forms, in deep languages, are similar images arbitrarily associated with meaning and audio. Hence, the VWFA may play a much more than important role in reading deep orthographies than in reading more than shallow orthographies, like Portuguese. Of grade, the present written report does non dominion out the part of the VWFA in shallow orthographies. It simply shows that reading sentences in Portuguese, relative to listening to sentences in Portuguese, did not show more activation that could be related to processing word form. The dissimilarity between reading and fixation in the present study did testify activation located at the VWFA coordinates specified above. Further investigation into the modulation of VWFA activation by dissimilar orthographies is warranted.

More overall brain activation for listening comprehension and left-laterality of activation for reading comprehension

Listening comprehension showed more overall activation of the encephalon than reading comprehension. Ane caption for the departure in full agile voxels is that the greater activation in listening could be a issue of the transient nature of auditory input. Auditory information is presented sequentially, while reading allows for backtracking, if necessary (Michael et al., 2001).

In the present study, however, reading stimuli were presented in sequential format, which did not allow for backtracking (RSVP format). In RSVP format, the visual input mirrors the transient feature of auditory input. The sequential presentation did not seem to event in an increase in overall reading activation, possibly ruling out the caption that more overall activation for listening comprehension is due to the transient nature of auditory input. The results showed that despite the RSVP presentation of visual sentences, the departure in overall brain activation for listening, compared to reading, remains. This difference has been reported in other studies that investigated how modality influences sentence comprehension (Lawman et al., 2004; Michael et al., 2001). Information technology seems that a different explanation for the greater overall activation in listening is warranted.

The developmental explanation for the bilateral distribution of brain activation in listening versus the left-lateralized activation in reading (Michael et al., 2001) may be a more suitable explanation for the difference in laterality of activation in listening and reading brain activation, and of greater overall activation in listening. This explanation posits that once humans learn to read, the brain response is grafted onto the existing left-lateralized, amodal brain network for linguistic communication comprehension. Later-learned reading comprehension is postulated every bit a second-order skill that is grafted onto the already-learned listening comprehension skill (Michael et al., 2001). Thus, as reading comprehension evolves into an automatic skill (like listening comprehension) and as humans larn to successfully operate higher-order cerebral processes in reading (inference-making, text integration, syntactic parsing), the cerebral responses to these processes are mapped onto the same areas of the brain that are associated with higher-level processes of listening comprehension. Every bit a kickoff-order language comprehension skill, listening comprehension retains near of the cognitive response that is intrinsic to its input class, whereas reading comprehension is mapped onto a preexisting brain response to language comprehension.

Individual differences in reading comprehension in RSVP format: Spillover, executive command, and phonological rehearsal

The study showed the brain activation for reading comprehension in rapid serial visual presentation. The transient nature of RSVP places additional load on working retention processes during reading possibly considering it removes the possibility of backtracking in reading comprehension. The results indicate that lower chapters readers recruited more cortical resources from right-hemisphere areas of the brain. Spillover of activation to the right-hemisphere is a known mechanism of recruitment of boosted cortical resources in lower chapters readers (Prat & Just, 2008). The results also bear witness that lower chapters readers had more than activation in the prefrontal cortex, specifically in left middle frontal gyrus. Activation of the prefrontal cortex has been consistently associated with executive control, and the left middle frontal gyrus is part of the executive network of cortical areas. The executive network consists of cortical areas activated in situations associated with controlled, strategic, and goal-oriented cerebral operations. It is a domain-full general network of the brain, held to be mediated by a frontal-parietal neural arrangement of processing centers (including dorsal and ventral left middle and inferior frontal cortex, medial frontal gyrus, and parietal cortices), and it is interpreted as a system engaged in dealing with novel cognitive tasks (D'Esposito et al., 1995).

The correlation of reading span scores and encephalon activity for higher-chapters readers showed significantly more activation in a frontal-posterior network of areas. The areas more activated for higher-bridge readers were left angular and precentral gyri, and RIFG. The activation of this network may be associated with phonological rehearsal of linguistic information.

The results indicated that higher and lower chapters readers may accept resorted to different strategies while reading sentences in RSVP format. Lower capacity readers, in add-on to recruiting more than right-hemisphere areas of the brain, may have resorted to more executive command processes to assistance keep rails of transient visual input and avoid comprehension processes from breaking down in a relatively novel type of reading task. Activation of the executive network of cortical areas is usually establish in tasks that demand increased attention to the stimulus and the ability to maintain previous data active in working retentivity, such as in dual tasks (D'Esposito et al., 1995; Jaeggi et al., 2003). More than activation in left middle frontal gyrus may indicate that reading in RSVP format may not take been a seamless process for lower capacity readers.

College-capacity readers, in plough, activated areas of the encephalon associated with phonological rehearsal of information. Rather than resorting to executive control processes, these readers may have been better able to adapt to the job of reading comprehension in transient, serial form. More activation in skilled readers may therefore reflect more successful subvocalization of the materials. More activation in areas associated with phonological rehearsal (left junior parietal lobe and LIFG) has been associated with the increased demand of reading natural language-twisters (Keller et al., 2003) and with keeping sound to meaning representations active in a second linguistic communication (Buchweitz et al., 2009). In the nowadays study, the high-span readers activated more left angular gyrus and RIFG, the right-hemisphere homologue of LIFG. The activation of RIFG may stand for spillover of activation from its left-hemisphere homologue.

The activation of this network of areas for high span readers may indicate that they were able to resort to phonological rehearsal of the words presented in serial format. Phonological rehearsal may have helped keep sound-to-meaning representations active after the written stimulus was no longer bachelor on the screen. Information technology is necessary to note that the study has a reduced number of participants and that there was no manipulation of task difficulty. Further studies with a larger population of participants, and that manipulate RSVP judgement difficulty, should be carried out to further ostend the present findings and evidence evidence of a correlation between reading bridge scores with brain activeness and with amend performance.

Determination

The study shows the modality fingerprints for the processing of spoken and written sentences. The differences in brain activation for reading and listening comprehension were found by and large in unimodal areas of the brain. The differences in brain activation between reading and listening comprehension approve other studies, even though the present study used a technique for serial presentation of written text. The study also shows individual differences in brain activation for reading comprehension in RSVP. The individual differences may be associated with different strategies for reading transient text. The results provide evidence to support ane of the premises underlying models of human comprehension: College-order language comprehension processes are amodal.

Acknowledgments

This inquiry was supported by the National Institute of Mental Wellness Grant MH029617. During the time of data collection Augusto Buchweitz was supported by the Brazilian Ministry of Education, CAPES BEX 3356-04-iii. Thanks to Jennifer Moore and Chantel Prat for comments on a previous version of this manuscript.

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Source: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3081613/

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