Reading-Related Brain Changes in Audiovisual Processing: Cross-Sectional and Longitudinal MEG Evidence

The ability to establish associations between visual objects and speech sounds is essential for human reading. Understanding the neural adjustments required for acquisition of these arbitrary audiovisual associations can shed light on fundamental reading mechanisms and help reveal how literacy builds on pre-existing brain circuits. To address these questions, the present longitudinal and cross-sectional MEG studies characterize the temporal and spatial neural correlates of audiovisual syllable congruency in children (age range, 4–9 years; 22 males and 20 females) learning to read. Both studies showed that during the first years of reading instruction children gradually set up audiovisual correspondences between letters and speech sounds, which can be detected within the first 400 ms of a bimodal presentation and recruit the superior portions of the left temporal cortex. These findings suggest that children progressively change the way they treat audiovisual syllables as a function of their reading experience. This reading-specific brain plasticity implies (partial) recruitment of pre-existing brain circuits for audiovisual analysis.SIGNIFICANCE STATEMENT Linking visual and auditory linguistic representations is the basis for the development of efficient reading, while dysfunctional audiovisual letter processing predicts future reading disorders. Our developmental MEG project included a longitudinal and a cross-sectional study; both studies showed that children''s audiovisual brain circuits progressively change as a function of reading experience. They also revealed an exceptional degree of neuroplasticity in audiovisual neural networks, showing that as children develop literacy, the brain progressively adapts so as to better detect new correspondences between letters and speech sounds.     

Disrupted neural responses to phonological and orthographic processing in dyslexic children: an fMRI study

Developmental dyslexia, characterized by difficulty in reading, has been associated with phonological and orthographic processing deficits. fMRI was performed on dyslexic and normal-reading children (8-12 years old) during phonological and orthographic tasks of rhyming and matching visually presented letter pairs. During letter rhyming, both normal and dyslexic reading children had activity in left frontal brain regions, whereas only normal-reading children had activity in left temporo-parietal cortex. During letter matching, normal-reading children showed activity throughout extrastriate cortex, especially in occipito-parietal regions, whereas dyslexic children had little activity in extrastriate cortex during this task. These results indicate dyslexia may be characterized in childhood by disruptions in the neural bases of both phonological and orthographic processes important for reading.     

Neurodevelopmental trajectories of letter and speech sound processing from preschool to the end of elementary school

Learning to read alphabetic languages starts with learning letter–speech-sound associations. How this process changes brain function during development is still largely unknown. We followed 102 children with varying reading skills in a mixed-longitudinal/cross-sectional design from the prereading stage to the end of elementary school over five time points (n = 46 with two and more time points, of which n = 16 fully-longitudinal) to investigate the neural trajectories of letter and speech sound processing using fMRI. Children were presented with letters and speech sounds visually, auditorily, and audiovisually in kindergarten (6.7yo), at the middle (7.3yo) and end of first grade (7.6yo), and in second (8.4yo) and fifth grades (11.5yo). Activation of the ventral occipitotemporal cortex for visual and audiovisual processing followed a complex trajectory, with two peaks in first and fifth grades. The superior temporal gyrus (STG) showed an inverted U-shaped trajectory for audiovisual letter processing, a development that in poor readers was attenuated in middle STG and absent in posterior STG. Finally, the trajectories for letter-speech-sound integration were modulated by reading skills and showed differing directionality in the congruency effect depending on the time point. This unprecedented study captures the development of letter processing across elementary school and its neural trajectories in children with varying reading skills.     

Emergence of the neural network underlying phonological processing from the prereading to the emergent reading stage: A longitudinal study

Numerous studies have shown that phonological skills are critical for successful reading acquisition. However, how the brain network supporting phonological processing evolves and how it supports the initial course of learning to read is largely unknown. Here, for the first time, we characterized the emergence of the phonological network in 28 children over three stages (prereading, beginning reading, and emergent reading) longitudinally. Across these three time points, decreases in neural activation in the left inferior parietal cortex (LIPC) were observed during an audiovisual phonological processing task, suggesting a specialization process in response to reading instruction/experience. Furthermore, using the LIPC as the seed, a functional network consisting of the left inferior frontal, left posterior occipitotemporal, and right angular gyri was identified. The connection strength in this network co‐developed with the growth of phonological skills. Moreover, children with above‐average gains in phonological processing showed a significant developmental increase in connection strength in this network longitudinally, while children with below‐average gains in phonological processing exhibited the opposite trajectory. Finally, the connection strength between the LIPC and the left posterior occipitotemporal cortex at the prereading level significantly predicted reading performance at the emergent reading stage. Our findings highlight the importance of the early emerging phonological network for reading development, providing direct evidence for the Interactive Specialization Theory and neurodevelopmental models of reading.     

Clinical neurophysiology of visual and auditory processing in dyslexia: A review

Neurophysiological studies on children and adults with dyslexia provide a deeper understanding of how visual and auditory processing in dyslexia might relate to reading deficits. The goal of this review is to provide an overview of research findings in the last two decades on motion related and contrast sensitivity visual evoked potentials and on auditory event related potentials to basic tone and speech sound processing in dyslexia. These results are particularly relevant for three important theories about causality in dyslexia: the magnocellular deficit hypothesis, the temporal processing deficit hypothesis and the phonological deficit hypothesis. Support for magnocellular deficits in dyslexia are primarily provided from evidence for altered visual evoked potentials to rapidly moving stimuli presented at low contrasts. Consistently ERP findings revealed altered neurophysiological processes in individuals with dyslexia to speech stimuli, but evidence for deficits processing certain general acoustic information relevant for speech perception, such as frequency changes and temporal patterns, are also apparent.     

Preschool impairments in auditory processing and speech perception uniquely predict future reading problems

Developmental dyslexia is characterized by severe reading and spelling difficulties that are persistent and resistant to the usual didactic measures and remedial efforts. It is well established that a major cause of these problems lies in poorly specified phonological representations. Many individuals with dyslexia also present impairments in auditory temporal processing and speech perception, but it remains debated whether these more basic perceptual impairments play a role in causing the reading problem. Longitudinal studies may help clarifying this issue by assessing preschool children before they receive reading instruction and by following them up through literacy development. The current longitudinal study shows impairments in auditory frequency modulation (FM) detection, speech perception and phonological awareness in kindergarten and in grade 1 in children who receive a dyslexia diagnosis in grade 3. FM sensitivity and speech-in-noise perception in kindergarten uniquely contribute to growth in reading ability, even after controlling for letter knowledge and phonological awareness. These findings indicate that impairments in auditory processing and speech perception are not merely an epiphenomenon of reading failure. Although no specific directional relations were observed between auditory processing, speech perception and phonological awareness, the highly significant concurrent and predictive correlations between all these variables suggest a reciprocal association and corroborate the evidence for the auditory deficit theory of dyslexia.     

Neural intersections of the phonological,visual magnocellular and motor/cerebellar systems in normal readers: Implications for imaging studies on dyslexia

We used fMRI to explore the extent of the anatomical overlap of three neural systems that the literature on developmental dyslexia associates with reading: the auditory phonological, the visual magnocellular, and the motor/cerebellar systems. Twenty‐eight normal subjects performed four tasks during fMRI scans: word and pseudoword reading, auditory rhyming for letter names, visual motion perception, and a motor sequence learning task. We found that the left occipitotemporal cortex (OTC), which previous studies reported to be dysfunctional in dyslexia, can be fractionated into different functional areas: an anterior and lateral area that was activated by both reading and auditory rhyming tasks; a posterior area that was commonly activated by both the reading and the motion perception task and a medial/intermediate area, including the so‐called Visual Word Form Area, which was specifically activated by the reading task. These results show that the left OTC is an area of segregated convergence of different functional systems. We compared our results with the hypoactivation pattern reported for reading in a previous cross‐cultural PET study on 36 dyslexic subjects from three countries. The region of decreased activation in dyslexia overlapped with regions that are specific for reading and those activated during both the auditory rhyming task and the single word and pseudoword reading task described in the present fMRI study. No overlap was found with the activation patterns for the visual motion perception task or for the motor sequence learning task. These observations challenge current theories of dyslexia. Hum Brain Mapp 34:2669–2687, 2013. © 2012 Wiley Periodicals, Inc.     

fMRI during word processing in dyslexic and normal reading children

The present study addresses phonological processing in children with developmental dyslexia. Following the hypothesis of a core deficit of assembled phonology in dyslexia a set of hierarchically structured tasks was applied that specifically control for different kinds of phonological coding (assembled versus addressed phonological strategies). Seventeen developmental dyslexics and 17 normal reading children were scanned during four different tasks: (1) passive viewing of letter strings (control condition), (2) passive reading of non-words, (3) passive reading of legal words, and (4) a task requiring phonological transformation. Statistical analysis of the data was performed using statistical parametric mapping (SPM96). Comparison of patterns of activation in dyslexic and normal reading children revealed significant differences in Broca's area and the left inferior temporal region for both, non-word reading and the phonological transformation task. The present data provide new evidence for alteration of the phonological system in dyslexic children, and in particular, the system that mediates assembled phonological coding.     

Training pseudoword reading in acquired dyslexia: a phonological complexity approach

Background: Individuals with acquired phonological dyslexia experience difficulty associating written letters with corresponding sounds, especially in pseudowords. Previous studies have shown that reading can be improved in these individuals by training letter–sound correspondence, practicing phonological skills, or using combined approaches. However, generalisation to untrained items is typically limited.

Aims: We investigated whether principles of phonological complexity can be applied to training letter–sound correspondence reading in acquired phonological dyslexia to improve generalisation to untrained words. Based on previous work in other linguistic domains, we hypothesised that training phonologically “more complex” material (i.e., consonant clusters with small sonority differences) would result in generalisation to phonologically “less complex” material (i.e., consonant clusters with larger sonority differences), but this generalisation pattern would not be demonstrated when training the “less complex” material.

Methods & Procedures: We used a single-participant, multiple baseline design across participants and behaviours to examine phonological complexity as a training variable in five individuals. Based on participants’ error data from a previous experiment, a “more complex” onset and a “less complex” onset were selected for training for each participant. Training order assignment was pseudo-randomised and counterbalanced across participants. Three participants were trained in the “more complex” condition and two in the “less complex” condition while tracking oral reading accuracy of both onsets.

Outcomes & Results: As predicted, participants trained in the “more complex” condition demonstrated improved pseudoword reading of the trained cluster and generalisation to pseudowords with the untrained, “simple” onset, but not vice versa.

Conclusions: These findings suggest phonological complexity can be used to improve generalisation to untrained phonologically related words in acquired phonological dyslexia. These findings also provide preliminary support for using phonological complexity theory as a tool for designing more effective and efficient reading treatments for acquired dyslexia.     

The long road to automation: neurocognitive development of letter-speech sound processing

In transparent alphabetic languages, the expected standard for complete acquisition of letter-speech sound associations is within one year of reading instruction. The neural mechanisms underlying the acquisition of letter-speech sound associations have, however, hardly been investigated. The present article describes an ERP study with beginner and advanced readers in which the influence of letters on speech sound processing is investigated by comparing the MMN to speech sounds presented in isolation with the MMN to speech sounds accompanied by letters. Furthermore, SOA between letter and speech sound presentation was manipulated in order to investigate the development of the temporal window of integration for letter-speech sound processing. Beginner readers, despite one year of reading instruction, showed no early letter-speech sound integration, that is, no influence of the letter on the evocation of the MMN to the speech sound. Only later in the difference wave, at 650 msec, was an influence of the letter on speech sound processing revealed. Advanced readers, with 4 years of reading instruction, showed early and automatic letter-speech sound processing as revealed by an enhancement of the MMN amplitude, however, at a different temporal window of integration in comparison with experienced adult readers. The present results indicate a transition from mere association in beginner readers to more automatic, but still not "adult-like," integration in advanced readers. In contrast to general assumptions, the present study provides evidence for an extended development of letter-speech sound integration.     

N300 indexes deficient integration of orthographic and phonological representations in children with dyslexia

In transparent orthographies, like German, children with developmental dyslexia (DD) are mainly characterized by a reading fluency deficit. The reading fluency deficit might be traced back to a scarce integration of orthographic and phonological representations. In order to address this question, the present study used EEG to investigate the N300, an ERP component which has been associated with the integration of orthographic and phonological representations. Twenty children without DD and 18 children with DD performed a phonological (P)-orthographic (O) matching task (P-O condition), which tapped the integration of orthographic and phonological representations. A control task was applied which did not require the integration of orthographic and phonological representations and consisted only of orthographic information (O-O condition). The O-O condition revealed a similar N300 distribution between groups with a bilateral activity over fronto-temporal electrodes. However, in the P-O condition N300 differentiated the 2 groups of children. The control group revealed greater activity over left fronto-temporal electrodes, whereas the N300 was distributed bilaterally in the group of children with DD suggesting deficient integration of orthographic and phonological representations. These findings might be related to the reading fluency deficit as it was also observed that better reading fluency was correlated with higher (r=-.36) and earlier peaking (r=-.33) N300 amplitudes in the left hemisphere and attenuated N300 amplitudes (r=.45) in the right hemisphere. Standardized low-resolution electromagnetic tomography analysis (sLORETA) revealed that children with DD rely more on right temporo-parietal brain areas compared to children without DD. Furthermore, in order to rule out that earlier deficient processes might influence the group differences found in the N300, we analyzed the N170 for group differences. We did not find significant differences between children without DD and children with DD. In conclusion the results suggest deficient integration of orthographic and phonological representations in dyslexia, as indexed by the N300, and further highlight how this activity is relevant for fluent reading.     

Auditory evoked potentials to speech and nonspeech stimuli are associated with verbal skills in preschoolers

Children’s obligatory auditory event-related potentials (ERPs) to speech and nonspeech sounds have been shown to associate with reading performance in children at risk or with dyslexia and their controls. However, very little is known of the cognitive processes these responses reflect. To investigate this question, we recorded ERPs to semisynthetic syllables and their acoustically matched nonspeech counterparts in 63 typically developed preschoolers, and assessed their verbal skills with an extensive set of neurocognitive tests. P1 and N2 amplitudes were larger for nonspeech than speech stimuli, whereas the opposite was true for N4. Furthermore, left-lateralized P1s were associated with better phonological and prereading skills, and larger P1s to nonspeech than speech stimuli with poorer verbal reasoning performance. Moreover, left-lateralized N2s, and equal-sized N4s to both speech and nonspeech stimuli were associated with slower naming. In contrast, children with equal-sized N2 amplitudes at left and right scalp locations, and larger N4s for speech than nonspeech stimuli, performed fastest. We discuss the possibility that children’s ERPs reflect not only neural encoding of sounds, but also sound quality processing, memory-trace construction, and lexical access. The results also corroborate previous findings that speech and nonspeech sounds are processed by at least partially distinct neural substrates.     

Perceptual organization of speech signals by children with and without dyslexia

Developmental dyslexia is a condition in which children encounter difficulty learning to read in spite of adequate instruction. Although considerable effort has been expended trying to identify the source of the problem, no single solution has been agreed upon. The current study explored a new hypothesis, that developmental dyslexia may be due to faulty perceptual organization of linguistically relevant sensory input. To test that idea, sentence-length speech signals were processed to create either sine-wave or noise-vocoded analogs. Seventy children between 8 and 11 years of age, with and without dyslexia participated. Children with dyslexia were selected to have phonological awareness deficits, although those without such deficits were retained in the study. The processed sentences were presented for recognition, and measures of reading, phonological awareness, and expressive vocabulary were collected. Results showed that children with dyslexia, regardless of phonological subtype, had poorer recognition scores than children without dyslexia for both kinds of degraded sentences. Older children with dyslexia recognized the sine-wave sentences better than younger children with dyslexia, but no such effect of age was found for the vocoded materials. Recognition scores were used as predictor variables in regression analyses with reading, phonological awareness, and vocabulary measures used as dependent variables. Scores for both sorts of sentence materials were strong predictors of performance on all three dependent measures when all children were included, but only performance for the sine-wave materials explained significant proportions of variance when only children with dyslexia were included. Finally, matching young, typical readers with older children with dyslexia on reading abilities did not mitigate the group difference in recognition of vocoded sentences. Conclusions were that children with dyslexia have difficulty organizing linguistically relevant sensory input, but learn to do so for the structure preserved by sine-wave signals before they do so for other sorts of signal structure. These perceptual organization deficits could account for difficulties acquiring refined linguistic representations, including those of a phonological nature, although ramifications are different across affected children.     

Brain activation profiles during the early stages of reading acquisition

In the present study, we demonstrate for the first time the presence of an aberrant brain mechanism for reading in children who have just started acquiring reading skills. Children who, at the end of kindergarten, are found to be at risk for developing reading problems display markedly different activation profiles than children who have, at this stage, already mastered important prereading skills. This aberrant profile is characterized by the lack of engagement of the left-hemisphere superior temporal region, an area normally involved in converting print into sound, and an increase in activation in the corresponding right-hemisphere region. This finding is consistent with current cognitive models of reading acquisition and dyslexia, pointing to the critical role of phonologic awareness skills in learning to read.     

Deviant neurophysiological responses to phonological regularities in speech in dyslexic children

Developmental dyslexia is strongly associated with a phonological deficit. Yet, implicit phonological processing (in)capacities in dyslexia remain relatively unexplored. Here we use a neurophysiological response sensitive to experience-dependent auditory memory traces, the mismatch negativity (MMN), to investigate implicit phonological processing of natural speech in dyslexic and normally reading children. In a modified passive oddball design that minimizes the contribution of acoustic processes, we presented non-words that differed by the degree of phonotactic probability, i.e. the distributional frequency of phoneme combinations in a given language. Overall morphology of ERP responses to the non-words indicated comparable processing of acoustic-phonetic stimulus differences in both children groups. Consistent with previous findings in adults, normally reading children showed a significantly stronger MMN response to the non-word with high phonotactic probability (notsel) as compared to the non-word with low phonotactic probability (notkel), suggesting auditory cortical tuning to statistical regularities of phoneme combinations. In contrast, dyslexic children did not show this sensitivity to phonotactic probability. These findings indicate that the phonological problems often reported in dyslexia relate to a subtle deficit in the implicit phonetic-phonological processing of natural speech.     

Out‐of‐synchrony speech entrainment in developmental dyslexia

Developmental dyslexia is a reading disorder often characterized by reduced awareness of speech units. Whether the neural source of this phonological disorder in dyslexic readers results from the malfunctioning of the primary auditory system or damaged feedback communication between higher‐order phonological regions (i.e., left inferior frontal regions) and the auditory cortex is still under dispute. Here we recorded magnetoencephalographic (MEG) signals from 20 dyslexic readers and 20 age‐matched controls while they were listening to ～10‐s‐long spoken sentences. Compared to controls, dyslexic readers had (1) an impaired neural entrainment to speech in the delta band (0.5–1 Hz); (2) a reduced delta synchronization in both the right auditory cortex and the left inferior frontal gyrus; and (3) an impaired feedforward functional coupling between neural oscillations in the right auditory cortex and the left inferior frontal regions. This shows that during speech listening, individuals with developmental dyslexia present reduced neural synchrony to low‐frequency speech oscillations in primary auditory regions that hinders higher‐order speech processing steps. The present findings, thus, strengthen proposals assuming that improper low‐frequency acoustic entrainment affects speech sampling. This low speech‐brain synchronization has the strong potential to cause severe consequences for both phonological and reading skills. Interestingly, the reduced speech‐brain synchronization in dyslexic readers compared to normal readers (and its higher‐order consequences across the speech processing network) appears preserved through the development from childhood to adulthood. Thus, the evaluation of speech‐brain synchronization could possibly serve as a diagnostic tool for early detection of children at risk of dyslexia. Hum Brain Mapp 37:2767–2783, 2016. © 2016 Wiley Periodicals, Inc .     

Newborn brain event-related potentials revealing atypical processing of sound frequency and the subsequent association with later literacy skills in children with familial dyslexia

The role played by an auditory-processing deficit in dyslexia has been debated for several decades. In a longitudinal study using brain event-related potentials (ERPs) we investigated 1) whether dyslexic children with familial risk background would show atypical pitch processing from birth and 2) how these newborn ERPs later relate to these same children’s pre-reading cognitive skills and literacy outcomes. Auditory ERPs were measured at birth for tones varying in pitch and presented in an oddball paradigm (1100 Hz, 12%, and 1000 Hz, 88%). The brain responses of the typically reading control group children (TRC group, N = 25) showed clear differentiation between the frequencies, while those of the group of reading disability with familial risk (RDFR, 8 children) and the group of typical readers with familial risk (TRFR, 14 children) did not differentiate between the tones. The ERPs of the latter two groups differed from those of the TRC group. However, the two risk groups also showed a differential hemispheric ERP pattern. Furthermore, newborn ERPs reflecting passive change detection were associated with phonological skills and letter knowledge prior to school age and with phoneme duration perception, reading speed (RS) and spelling accuracy in the 2nd grade of school. The early obligatory response was associated with more general pre-school language skills, as well as with RS and reading accuracy (RA). Results suggest that a proportion of dyslexic readers with familial risk background are affected by atypical auditory processing. This is already present at birth and also relates to pre-reading phonological processing and speech perception. These early differences in auditory processing could later affect phonological representations and reading development. However, atypical auditory processing is unlikely to suffice as a sole explanation for dyslexia but rather as one risk factor, dependent on the genetic profile of the child.     

The neurophysiological basis of the integration of written and heard syllables in dyslexic adults

ObjectiveLetter-speech sound integration in fluent readers takes place automatically and is dependent on temporal synchrony between letters and sounds. In developmental dyslexia, however, letter-speech sound associations are hard to learn, compromising accurate and fluent reading. We studied the effect of printed text on processing speech sounds in dyslexic and fluent adult readers.MethodsVisual stimuli were presented with sequences of spoken syllables including vowel or consonant changes, or changes in syllable intensity, frequency, or vowel duration. As visual material, written syllables or their scrambled images were used. The auditory stimuli were presented either synchronously with the visual stimuli or time delayed. The mismatch negativity (MMN), an index of automatic neural change detection, was recorded.ResultsMMN amplitudes were larger to syllable changes in combination with written syllables than with scrambled images in fluent readers. However, dyslexic readers showed no difference between syllables vs. scrambled image condition. Furthermore, MMNs to consonant and frequency changes peaked later in dyslexic than fluent readers.ConclusionsOur results suggest deficient and sluggish audiovisual integration in dyslexic individuals, which is not dependent on the phonological relevance of the deviant type.SignificanceUnlike previous studies, our study included several different types of syllable changes presented with concurrent print, enabling us to determine in more detail the nature of the audiovisual deficit in dyslexia.     

Examining the neural and cognitive processes that underlie reading through naming speed tasks

We combined fMRI with eye tracking and speech recording to examine the neural and cognitive mechanisms that underlie reading. To simplify the study of the complex processes involved during reading, we used naming speed (NS) tasks (also known as rapid automatized naming or RAN) as a focus for this study, in which average reading right‐handed adults named sets of stimuli (letters or objects) as quickly and accurately as possible. Due to the possibility of spoken output during fMRI studies creating motion artifacts, we employed both an overt session and a covert session. When comparing the two sessions, there were no significant differences in behavioral performance, sensorimotor activation (except for regions involved in the motor aspects of speech production) or activation in regions within the left‐hemisphere‐dominant neural reading network. This established that differences found between the tasks within the reading network were not attributed to speech production motion artifacts or sensorimotor processes. Both behavioral and neuroimaging measures showed that letter naming was a more automatic and efficient task than object naming. Furthermore, specific manipulations to the NS tasks to make the stimuli more visually and/or phonologically similar differentially activated the reading network in the left hemisphere associated with phonological, orthographic and orthographic‐to‐phonological processing, but not articulatory/motor processing related to speech production. These findings further our understanding of the underlying neural processes that support reading by examining how activation within the reading network differs with both task performance and task characteristics.     

Survey of fMRI results regarding a phonological deficit in children and adults with dyslexia: fundamental deficit or indication of compensation?

Early neuroimaging studies exploring the neurobiological correlates of the phonological deficit in dyslexia were restricted to adult probands with dyslexia due to the exposure to radioactivity in the course of PET measurements. The differences in activation between normal adult readers and adults with dyslexia recorded in these studies left open the issue of whether or not these are indeed fundamental activation deficits or only a reflection of lifelong experience with poor reading and writing skills and thus should be interpreted as a sign of compensation. Development of fMRI in recent years has enabled the investigation of children with dyslexia in order to explore the neurobiological activation patterns that underlie dyslexia. On the whole, the imaging findings in children and adults with dyslexia indicate that the left-hemisphere inferior frontal differences in activation, as well as the dorsal and ventral temporal differences in activation observed in all age groups during the processing of phonological language tasks are to be regarded as a fundamental biological deficit in dyslexia. Right-hemisphere differences in activation, which in German-speaking areas are observable in adults but only rarely in children with dyslexia, might, on the other hand, be regarded as a sign of developmental compensation.