signature=5a5aedb582a14169a2045badbad186e2,Neurocognitive Signatures of Naturalistic Reading of Scie...

Participants

Sixty-two right-handed native English speakers were recruited. Seven participants did not finish the first session due to eye-tracker or MR scanner technical issues. One participant was excluded due to very low accuracy (50%) for an in-scanner comprehension test and poor behavioural testing results outside the scanner. One participant was found to be left-handed after the behavioural session, leaving 51 participants aged between 18 and 40 years in our analysis. Eye-tracking data were missing for one participant during one run containing one text, leading to its exclusion for the analysis for KS. Forty-nine out of the 51 participants completed the behavioural testing session, of which only 46 correctly performed the Letter Number Sequencing task. Therefore, behavioural data analysis included 49 participants (23 males, mean age ± SD = 22.69 ± 4.57). fMRI data for neural correlates of Reading and Integrative Processing included 51 participants (24 males, mean age ± SD = 22.67 ± 4.52). Forty-six participants (21 males, mean age ± SD = 22.84 ± 4.63) were included in the fMRI multiple regression models for neural correlates of individual differences in Integrative Processing. Forty-five participants (21 males, mean age ± SD = 22.47 ± 3.88) were included in fMRI regression models for neural correlates of individual differences in sentential processing of texts with different KS optimality.

All participants had normal or corrected to normal vision, and had no history of mental or neurological disorder. The study was approved by the Pennsylvania State University Institutional Review Board (IRB) and was performed in accordance with the ethical standards described in the IRB. Written informed consent was obtained from all participants before they took part in the study.

Materials

Prior to the experiment, five expository texts of STEM contents were modified from previous research stimuli (see FollmerF = 0.05, p = 0.99), lexical decision time (LDT, F = 1.07, p = 0.38), log frequency (F = 0.25, p = 0.91), naming response time (NRT, F = 1.41, p = 0.23), orthographic neighbourhood density (OLD, F = 0.04, p = 0.99), phonological neighbourhood density (PLD, F = 0.34, p = 0.85), concreteness (F = 0.24, p = 0.91), and number of phonemes (NPhon, F = 0.02, p = 0.99). However, one-way ANOVAs for average word length and AoA were significant at p 

KS quantified as maximal betweenness centrality (MBC, Graph Centrality)

Fifteen key terms were selected as nodes from each of the five textsAnalysis of Lexical Aggregates Reader (ALA-Reader)k in a network, its partial betweenness with respect to the other two nodes i and j is defined as the probability that node k falls on a randomly selected path linking nodes i and j. The betweenness centrality value of node k is the sum of the partial betweenness values in respect to all pairs of nodes in the network except for ktextual KS, the centrality values were normalised and quadratic terms were calculated. Higher quadratic centrality values (further away from zero, which is the average in the normalised distribution) indicate sub-optimal KS, while lower quadratic centrality values (closer to zero) indicate more optimal KS.

Procedure

After providing consent, participants underwent a structural MRI scan, followed by a practice session for self-paced reading in the scanner. They were instructed to click a button to advance from one sentence to the next. Each sentence was presented for up to 8 seconds after which the next sentence automatically appeared on the screen. At the end of each text they answered 10 comprehension questions. Once the practice session ended, the participants completed five self-paced reading sessions, during which time simultaneous fMRI and eye-tracking data were collected. On a second visit, which was usually one week after the in-scanner reading session, participants completed a battery of behavioural tests.

Behavioural data collection and processing

In the behavioural session, the Gray Silent Reading Test, Raven’s Progressive Matrices, Letter Number Sequencing and Attention Network tests were presented to participants via E-Prime 2.0, and the Reading Background Questionnaire was completed on an internet browser. Detailed information of each test is as below.

Gray Silent Reading Test (GSRT)

The GSRT test measures reading comprehension competence

Raven’s progressive matrices

The Raven’s test measures analogical reasoning

Letter number sequencing (LNS)

The LNS task measures working memory. The task was adapted from the Wechsler Adult Intelligence Scale (WAIS-III)

Attention network test (ANT)

The ANT tests measure the alerting and orienting skills of attention and the inhibitory control ability of executive functionet al.

Reading Background Questionnaire (RBQ)

Participants were administered 20 questions constructed based on previous research

Correlational analyses showed significant correlations between E-device reading and non-reading time, and pair-wise correlations among reading time, reading preference and reading attitude/ability (see also Follmer et al.’s analyses

Behavioural data analyses

To test what cognitive measures contribute to participants’ reading comprehension behaviourally, we performed non-parametric correlation tests checking correlations between GSRT or question-answering accuracy with the Raven’s scores, LNS scores, the ANT Alerting, Orienting, and Conflict scores, the RBQ E-device reading and reading preference indices. Because the mean accuracy of the performance assessment scores and the GSRT scores violated the assumption of normality (Shapiro-Wilk W test, both ps 

Eye-tracking data acquisition and processing

The basic idea of fixation-related fMRI paradigm, as first explored by Marsman et al.

Data adjustment was later performed to address drifting issues caused by the calibration accuracy decline over time. For fixations falling outside (above or below) the range of predefined target regions, manual adjustment was performed using the Data Viewer software. Instead of using auto-adjustment which brings all fixations onto one horizontal line, we performed trial-by-trial correction adjusting all of the fixations in a single try only along the y axis (vertical adjustment) so as to maintain readers’ original eye fixation patterns.

MRI data acquisition

Data were acquired using a 3 T Siemens Magnetom Prisma Fit scanner with a 64-channel phased array coil. We acquired a MPRAGE scan with T1 weighted contrast [176 ascending sagittal slices with A/P phase encoding direction; voxel size = 1 mm isotropic; FOV = 256 mm; TR = 1540 ms; TE = 2.34 ms; acquisition time = 216 s; flip angle = 9°; GRAPPA in-plane acceleration factor = 2; brain coverage is complete for cerebrum, cerebellum and brain stem]. After the T1, we acquired five functional runs of T2* weighted echo planar sequence images [30 interleaved axial slices with A/P phase encoding direction; voxel size = 3 × 3 × 4 mm; FOV = 240 mm; TR = 400 ms; TE = 30 ms; acquisition time varied on the speed of self-paced reading, maximal 306 s; multiband acceleration factor for parallel slice acquisition = 6; flip angle = 35°; brain coverage misses the top of the parietal lobe and the lower end of the cerebellum]. Additionally, we collected a pair of spin echo sequence images with A/P and P/A phase encoding direction [30 axial interleaved slices; voxel size = 3 × 3 × 4 mm; FOV = 240 mm; TR = 3000 ms; TE = 51.2 ms; flip angle = 90°] to calculate distortion correction for the multiband sequences

fMRI data preprocessing and analyses

Data preprocessing and analysis were performed in SPM12 v6906 (http://www.fil.ion.ucl.ac.uk/spm). Functional imaging preprocessing consisted of correction of field inhomogeneity artefacts with the HySCO toolbox (Hyperelastic Susceptibility Artifact Correction)th degree B-Spline Interpolation algorithm and further smoothed with a Gaussian kernel of 8 mm full-width-at-half-maximum (FWHM).

In the GLM analysis, the design matrix contained one psychological regressor of interest, the “Content Word” condition, specifying the onsets and gaze durations of first pass fixations and regressions for content words (informed by eye-tracking data). The index of word position in sentences (starting from 1) was incorporated as a parametric modulator of the “Content Word” condition. We also included two psychological regressors of non-interest: “Non-Content Word” and “Instructions”: the “Non-Content Word” condition modelled fixations on non-content (function) words and ocular regressions, and the “Instructions” condition modelled two seconds of instructions presented at the beginning of each run. Because of the self-paced reading, all psychological regressors at the first level were subject-specific. Finally, we included six motion parameters and three physiological regressors (white matter, ventricular, and non-ventricular CSF space signal). We then applied a high pass filter with a cut off period of 128 s, and the temporal autocorrelation was accounted for with the FAST option in SPM12

At the group-level, the beta maps of the Integrative Processing obtained at the subject-level were entered into one multiple regression model as the dependent variable (N = 46). The following eight independent variables were included to checked the effect of individual differences: (1) GSRT, (2) Raven’s, (3) span-weighted LNS, the (4) Alerting, (5) Orienting, and (6) Conflict effects of the ANT, (7) the RBQ E-device reading index and (8) the RBQ reading preference index. At the whole brain level, we applied cluster-level FWE-correction p 

To further investigate Integrative Processing due to the effects of textual KS (measured as MBC, see Materials in the Methods), the beta maps of Integrative Processing of each text were entered into a subject-level regression model including the linear and quadratic terms of MBC as the independent variable. At the group level, the beta maps of quadratic MBC correlates of the Integrative processes were entered into an one-sample t-test (N = 50) for the main effect and a multiple regression model (N = 45) with the same eight independent variables for individual differences as mentioned before. We applied cluster-level FWE-correction p 

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