Reduced benefit from long-term item frequency contributes to short-term memory deficits in dyslexia.

Dyslexia, a specific difficulty in acquiring proficient reading, is also characterized by reduced short-term memory (STM) capacity. Extensive research indicates that individuals with developmental dyslexia (IDDs) benefit less from exposure, and this hampers their long-term knowledge accumulation. It is well established that long-term knowledge has a great effect on performance in STM tasks, and thus IDDs' reduced benefit of exposure could potentially reduce their relative performance in such tasks, especially when frequent items, such as digit-words, are used. In this study we used a standard, widely used, STM assessment: the Digit Span subtest from the Wechsler Adult Intelligence Scale. The task was conducted twice: in native language and in second language. As exposure to native language is greater than exposure to second language, we predicted that IDDs' performance in the task administered in native language will reveal a larger group difference as compared to second language, due to IDDs' reduced benefit of item frequency. The prediction was confirmed, in line with the hypothesis that reduced STM in dyslexia to a large extent reflects reduced benefits from long-term item frequency and not a reduced STM per se.

Reduced categorical learning of faces in dyslexia.

The perception of phonological categories in dyslexia is less refined than in typically developing (TD) individuals. Traditionally, this characteristic was considered unique to phonology, yet many studies showed non-phonological perceptual difficulties. Importantly, measuring the dynamics of cortical adaptation, associated with category acquisition, revealed a broadly distributed faster decay of cortical adaptation. Taken together, these observations suggest that the acquisition of perceptual categories in dyslexia may be slower across modalities. To test this, we tested adult individuals with developmental dyslexia (IDDs) and TDs on learning of two unknown faces, yielding face-specific categorization. Initial accuracy was similar in the two groups, yet practice-induced increase in accuracy was significantly larger in TDs. Modeling the learning process (using Drift Diffusion Model) revealed that TDs' steeper learning results from a larger increase in their effective face-specific signal. We propose that IDDs' slower item-specific categorical learning of unknown faces indicates that slower categorical learning in dyslexia is a core, domain-general difficulty.

Frequency-specific contributions to auditory perceptual priors: Testing the predictive-coding hypothesis.

Perceptual priors formed by recent stimuli bias our immediate percept. These priors, expressing our implicit expectations, affect both high- and low-level processing stages. Yet, the nature of the inter-level interaction is unknown. Do priors operate top-down and bias low-level features toward recently experienced objects (predictive-coding hypothesis), or are low-level biases bottom-up driven and formed by local memory circuits? To decipher between these options in auditory perception, we used the "missing fundamental illusion", enabling the dissociation of low-level components from the high-level pitch. Surprisingly, in contrast to predictive coding, when the fundamental frequency was missing, pitch contraction across timbre categories was not found to the previously perceived high-level pitch, but to the physically present frequency. This bottom-up contribution of low-level memory components to perceptual priors, operating independently of recent high-level percepts, may stabilize the perceptual organization and underlie continuity between similar low-level features belonging to different object categories in the auditory modality.

Neural correlates of statistical learning in developmental dyslexia: An electroencephalography study.

The human brain extracts statistical regularities from the surrounding environment in a process called statistical learning. Behavioural evidence suggests that developmental dyslexia affects statistical learning. However, surprisingly few studies have assessed how developmental dyslexia affects the neural processing underlying this type of learning. We used electroencephalography to explore the neural correlates of an important aspect of statistical learning - sensitivity to transitional probabilities - in individuals with developmental dyslexia. Adults diagnosed with developmental dyslexia (n = 17) and controls (n = 19) were exposed to a continuous stream of sound triplets. Every so often, a triplet ending had a low transitional probability given the triplet's first two sounds ("statistical deviants"). Furthermore, every so often a triplet ending was presented from a deviant location ("acoustic deviants"). We examined mismatch negativity elicited by statistical deviants (sMMN), and MMN elicited by location deviants (i.e., acoustic changes). Acoustic deviants elicited a MMN which was larger in the control group than in the developmental dyslexia group. Statistical deviants elicited a small, yet significant, sMMN in the control group, but not in the developmental dyslexia group. However, the difference between the groups was not significant. Our findings indicate that the neural mechanisms underlying pre-attentive acoustic change detection and implicit statistical auditory learning are both affected in developmental dyslexia.

Poor synchronization yet adequate tempo-keeping in adults with autism.

Sensorimotor synchronization to external events is fundamental to social interactions. Adults with autism spectrum condition (ASC) have difficulty with synchronization, manifested in both social and non-social situations, such as paced finger-tapping tasks, where participants synchronize their taps to metronome beats. What limits ASC's synchronization is a matter of debate, especially whether it stems from reduced online correction of synchronization error (the "slow update" account) or from noisy internal representations (the "elevated internal noise" account). To test these opposing theories, we administered a synchronization-continuation tapping task, with and without tempo changes. Participants were asked to synchronize with the metronome and continue the tempo when it stopped. Since continuation is based only on internal representations, the slow update hypothesis predicts no difficulty, whereas the elevated noise hypothesis predicts similar or enhanced difficulties. Additionally, tempo changes were introduced, to assess whether adequate updating of internal representations to external changes is possible when given a longer temporal window for updating. We found that the ability to keep the metronome's tempo after it stopped did not differ between ASC and typically developing (TD) individuals. Importantly, when given a longer period to adapt to external changes, keeping a modified tempo was also similar in ASC. These results suggest that synchronization difficulties in ASC stem from slow update rather than elevated internal noise.

Perceptual history biases in serial ensemble representation.

Ensemble perception refers to the visual system's ability to efficiently represent groups of similar objects as a unified percept using their summary statistical information. Most studies focused on extraction of current trial averages, giving little attention to prior experience effects, although a few recent studies found that ensemble mean estimations contract toward previously presented stimuli, with most of these focusing on explicit perceptual averaging of simultaneously presented item ensembles. Yet, the time element is crucial in real dynamic environments, where we encounter ensemble items over time, aggregating information until reaching summary representations. Moreover, statistical information of objects and scenes is learned over time and often implicitly and then used for predictions that shape perception, promoting environmental stability. Therefore, we now focus on temporal aspects of ensemble statistics and test whether prior information, beyond the current trial, biases implicit perceptual decisions. We designed methods to separate current trial biases from those of previously seen trial ensembles. In each trial, six circles of different sizes were presented serially, followed by two test items. Participants were asked to choose which was present in the sequence. Participants unconsciously rely on ensemble statistics, choosing stimuli closer to the ensemble mean. To isolate the influence of earlier trials, the two test items were sometimes equidistant from the current trial mean. Results showed membership judgment biases toward current trial mean, when informative (largest effect). On equidistant trials, judgments were biased toward previously experienced stimulus statistics. Comparison of similar conditions with a shifted stimulus distribution ruled out a bias toward an earlier, presession, prototypical diameter. We conclude that ensemble perception, even for temporally experienced ensembles, is influenced not only by current trial mean but also by means of recently seen ensembles and that these influences are somewhat correlated on a participant-by-participant basis.

Training allows switching from limited-capacity manipulations to large-capacity perceptual processing.

In contrast to perceptual tasks, which enable concurrent processing of many stimuli, working memory (WM) tasks have a very small capacity, limiting cognitive skills. Training on WM tasks often yields substantial improvement, suggesting that training might increase the general WM capacity. To understand the underlying processes, we trained a test group with a newly designed tone manipulation WM task and a control group with a challenging perceptual task of pitch pattern discrimination. Functional magnetic resonance imaging (fMRI) scans confirmed that pretraining, manipulation was associated with a dorsal fronto-parietal WM network, while pitch comparison was associated with activation of ventral auditory regions. Training induced improvement in each group, which was limited to the trained task. Analyzing the behavior of the group trained with tone manipulation revealed that participants learned to replace active manipulation with a perceptual verification of the position of a single salient tone in the sequence presented as a tentative reply. Posttraining fMRI scans revealed modifications in ventral activation of both groups. Successful WMtrained participants learned to utilize auditory regions for the trained task. These observations suggest that the huge task-specific enhancement of WM capacity stems from a task-specific switch to perceptual routines, implemented in perceptual regions.

Repeated series learning revisited with a novel prediction on the reduced effect of item frequency in dyslexia.

Developmental dyslexia, a difficulty with acquiring fluent reading, has also been characterized by reduced short-term memory (STM) capacity, which is often operationalized with span tasks. The low performance of individuals with dyslexia (IDDs) in such tasks is commonly attributed to poor phonological memory. However, we suggest an alternative explanation based on the observation that many times the items that are used in spans tasks are high-frequency items (e.g., digit words). We suggest that IDDs do not enjoy the benefit of item frequency to the same extent as controls, and thus their performance in span tasks is especially hampered. On the contrary, learning of repeated sequences was shown to be largely independent of item frequency, and therefore this type of learning may be unimpaired in dyslexia. To test both predictions, we used the Hebb-learning paradigm. We found that IDDs' performance is especially poor compared to controls' when high-frequency items are used, and that their repeated series learning does not differ from that of controls. Taken together with existing literature, our findings suggest that impaired learning of repeated series is not a core characteristic of dyslexia, and that the reports on reduced STM in dyslexia may to a large extent be explained by reduced benefit of item frequency.

Reduced Learning of Sound Categories in Dyslexia Is Associated with Reduced Regularity-Induced Auditory Cortex Adaptation.

A main characteristic of dyslexia is poor use of sound categories. We now studied within-session learning of new sound categories in dyslexia, behaviorally and neurally, using fMRI. Human participants (males and females) with and without dyslexia were asked to discriminate which of two serially-presented tones had a higher pitch. The task was administered in two protocols, with and without a repeated reference frequency. The reference condition introduces regularity, and enhances frequency sensitivity in typically developing (TD) individuals. Enhanced sensitivity facilitates the formation of "high" and "low" pitch categories above and below this reference, respectively. We found that in TDs, learning was paralleled by a gradual decrease in activation of the primary auditory cortex (PAC), and reduced activation of the superior temporal gyrus (STG) and left posterior parietal cortex (PPC), which are important for using sensory history. No such sensitivity was found among individuals with dyslexia (IDDs). Rather, IDDs showed reduced behavioral learning of stimulus regularities and no regularity-associated adaptation in the auditory cortex or in higher-level regions. We propose that IDDs' reduced cortical adaptation, associated with reduced behavioral learning of sound regularities, underlies their impoverished use of stimulus history, and consequently impedes their formation of rich sound categories.SIGNIFICANCE STATEMENT Reading difficulties in dyslexia are often attributed to poor use of phonological categories. To test whether poor category use could result from poor learning of new sound categories in general, we administered an auditory discrimination task that examined the learning of new pitch categories above and below a repeated reference sound. Individuals with dyslexia (IDDs) learned categories slower than typically developing (TD) individuals. TD individuals showed adaptation to the repeated sounds that paralleled the category learning in their primary auditory cortex (PAC) and other higher-level regions. In dyslexia, no brain region showed such adaptation. We suggest that poor learning of sound statistics in sensory regions may underlie the poor representations of both speech and nonspeech categories in dyslexia.

Slow update of internal representations impedes synchronization in autism.

Autism is a neurodevelopmental disorder characterized by impaired social skills, motor and perceptual atypicalities. These difficulties were explained within the Bayesian framework as either reflecting oversensitivity to prediction errors or - just the opposite - slow updating of such errors. To test these opposing theories, we administer paced finger-tapping, a synchronization task that requires use of recent sensory information for fast error-correction. We use computational modelling to disentangle the contributions of error-correction from that of noise in keeping temporal intervals, and in executing motor responses. To assess the specificity of tapping characteristics to autism, we compare performance to both neurotypical individuals and individuals with dyslexia. Only the autism group shows poor sensorimotor synchronization. Trial-by-trial modelling reveals typical noise levels in interval representations and motor responses. However, rate of error correction is reduced in autism, impeding synchronization ability. These results provide evidence for slow updating of internal representations in autism.

Dissecting the Roles of Supervised and Unsupervised Learning in Perceptual Discrimination Judgments.

Our ability to compare sensory stimuli is a fundamental cognitive function, which is known to be affected by two biases: choice bias, which reflects a preference for a given response, and contraction bias, which reflects a tendency to perceive stimuli as similar to previous ones. To test whether both reflect supervised processes, we designed feedback protocols aimed to modify them and tested them in human participants. Choice bias was readily modifiable. However, contraction bias was not. To compare these results to those predicted from an optimal supervised process, we studied a noise-matched optimal linear discriminator (Perceptron). In this model, both biases were substantially modified, indicating that the "resilience" of contraction bias to feedback does not maximize performance. These results suggest that perceptual discrimination is a hierarchical, two-stage process. In the first, stimulus statistics are learned and integrated with representations in an unsupervised process that is impenetrable to external feedback. In the second, a binary judgment, learned in a supervised way, is applied to the combined percept.SIGNIFICANCE STATEMENT The seemingly effortless process of inferring physical reality from the sensory input is highly influenced by previous knowledge, leading to perceptual biases. Two common ones are contraction bias (the tendency to perceive stimuli as similar to previous ones) and choice bias (the tendency to prefer a specific response). Combining human psychophysical experiments with computational modeling we show that they reflect two different learning processes. Contraction bias reflects unsupervised learning of stimuli statistics, whereas choice bias results from supervised or reinforcement learning. This dissociation reveals a hierarchical, two-stage process. The first, where stimuli statistics are learned and integrated with representations, is unsupervised. The second, where a binary judgment is applied to the combined percept, is learned in a supervised way.

Training-induced improvement in working memory tasks results from switching to efficient strategies.

It is debated whether training with a working memory (WM) task, particularly n-back, can improve general WM and reasoning skills. Most training studies found substantial improvement in the trained task, with little to no transfer to untrained tasks. We hypothesized that training does not increase WM capacity, but instead provides opportunities to develop an efficient task-specific strategy. We derived a strategy for the task that optimizes WM resources and taught it to participants. In two sessions, 14 participants who were taught this strategy performed as well as fourteen participants who trained for 40 sessions without strategy instructions. To understand the mechanisms underlying the no-instruction group's improvement, participants answered questionnaires during their training period. Their replies indicate that successful learners discovered the same strategy and their improvement was associated with this discovery. We conclude that n-back training allows the discovery of strategies that enable better performance with the same WM resources.

Short-term memory capacity and sensitivity to language statistics in dyslexia and among musicians.

Poor short-term memory (STM) capacity in individuals with dyslexia (IDDs) and enhanced STM capacity in musicians are well documented, yet their causes are disputed. Previous studies also found poor use of stimuli statistics by IDDs and enhanced use by musicians. We hypothesized that these observations are functionally related, as follows: Enhanced sensitivity to statistics facilitates musicians' benefit from each exposure, and reduced sensitivity to statistics hinders IDDs' benefit. Thus, larger group differences are expected for larger exposure: STM capacity, which is sensitive to item familiarity, will thus be larger among musicians, and smaller among IDDS, particularly for high-frequency items. Testing this hypothesis using syllable span, we found that musicians' advantage and IDDs' difficulty were indeed larger for high-frequency syllables than for low-frequency ones. By contrast, benefits from sequence repetition did not differ between musicians, controls and IDDs, suggesting that online sequence learning is based on a different mechanism. To test this dissociation we recruited, in addition to native Hebrew speakers, native English speakers, matched to the Hebrew-speaking controls. Their spans for high-frequency syllables in Hebrew, which do not have high frequency in English, were small - as expected from reduced exposure to these syllables. Yet, their benefit from sequence repetition was similar to that of the three Hebrew-speaking groups. Taken together, these experiments suggest that different sensitivities to item frequency explain some of the population-related variability in STM tasks. By contrast, benefits from sequence repetition do not depend on item familiarity, and do not differ between groups.

Benefits from morphological regularities in dyslexia are task dependent.

Are difficulties of individuals with dyslexia (IDDs) reduced or enhanced in tasks where linguistic regularities typically facilitate performance, such as vocabulary acquisition and reading? If impaired short-term memory and poor phonological decoding pose the main impediments to IDDs, then they are expected to compensate for these difficulties with a greater reliance on linguistic regularities, to reduce online load. However, if reduced benefits from regularities pose the main bottleneck, IDDs might benefit less than good readers from regularities in spite of their online difficulties. To test that, we administered two experiments. In a novel paradigm of auditory vocabulary acquisition in Hebrew, novel words were presented either with or without familiar morphological structure. Participants with dyslexia showed a reduced recall benefit from familiar structure as compared with controls. However, their recognition was facilitated by morphological structure and did not significantly differ from controls'. In the second experiment, participants read novel words with and without familiar structure. Benefit from structure familiarity for IDDs was significantly smaller than for controls, in spite of IDDs' greater potential benefit from familiar structure due to their reduced overall accuracy. However, when asked to emphasize speed in reading, structure familiarity was found to be beneficial for IDDs, without compromising accuracy. These results imply that accumulative acquisition of sublexical regularities is less efficient in dyslexia, though in some tasks this knowledge is accessible and beneficial. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Auditory frequency discrimination is correlated with linguistic skills, but its training does not improve them or other pitch discrimination tasks.

Brain-training, aimed at advancing and improving cognitive and perceptual abilities, is vastly studied because of its immense promise. Yet, there are major controversies regarding its main claim that intensive weeks' training on a single challenging task could improve performance in related untrained tasks. Ample training studies showing transfer were criticized for flawed design. We now explored the impact of perceptual training (auditory frequency discrimination), applying a carefully controlled intensive training experiment. First, we administered a battery of perceptual, linguistic, and cognitive tasks to a large population to determine "near" to "far" tasks according to (pretraining) correlations in performance. This assessment revealed significant correlations between simple pitch discrimination and complex linguistic tasks, including reading and syntactic reasoning. Second, we administered a broad test battery before (and after) training, which included several tasks assessing pitch discrimination, and the linguistic tasks that showed pretraining correlation with auditory frequency discrimination. The test group trained with 2 tone frequency discrimination for 40 sessions. An active control group trained with a working memory (n-back) task for the same duration, and a passive control group was only tested before and after training. Pretraining performance levels were similar in the three groups. Our results were straightforward. No transfer was found to untrained tasks that rely on pith discrimination, or to linguistic tasks that showed pretraining correlation. Mild to marginal transfer was found only to pitch discrimination tasks using almost exactly the trained protocol. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Perceptual bias reveals slow-updating in autism and fast-forgetting in dyslexia.

Individuals with autism and individuals with dyslexia both show reduced use of previous sensory information (stimuli statistics) in perceptual tasks, even though these are very different neurodevelopmental disorders. To better understand how past sensory information influences the perceptual experience in these disorders, we first investigated the trial-by-trial performance of neurotypical participants in a serial discrimination task. Neurotypical participants overweighted recent stimuli, revealing fast updating of internal sensory models, which is adaptive in changing environments. They also weighted the detailed stimuli distribution inferred by longer-term accumulation of stimuli statistics, which is adaptive in stable environments. Compared to neurotypical participants, individuals with dyslexia weighted earlier stimuli less heavily, whereas individuals with autism spectrum disorder weighted recent stimuli less heavily. Investigating the dynamics of perceptual inference reveals that individuals with dyslexia rely more on information about the immediate past, whereas perception in individuals with autism is dominated by longer-term statistics.

Shorter cortical adaptation in dyslexia is broadly distributed in the superior temporal lobe and includes the primary auditory cortex.

Studies of the performance of individuals with dyslexia in perceptual tasks suggest that their implicit inference of sound statistics is impaired. Previously, using two-tone frequency discrimination, we found that the effect of previous trials' frequencies on the judgments of individuals with dyslexia decays faster than the effect on controls' judgments, and that the adaptation (decrease of neural response to repeated stimuli) of their ERP responses to tones is shorter (Jaffe-Dax et al., 2017). Here, we show the cortical distribution of these abnormal dynamics of adaptation using fast-acquisition fMRI. We find that faster decay of adaptation in dyslexia is widespread, although the most significant effects are found in the left superior temporal lobe, including the auditory cortex. This broad distribution suggests that the faster decay of implicit memory of individuals with dyslexia is a general characteristic of their cortical dynamics, which also affects sensory cortices.

Auditory Stimulus Processing and Task Learning Are Adequate in Dyslexia, but Benefits From Regularities Are Reduced.

Purpose: The reading deficit for people with dyslexia is typically associated with linguistic, memory, and perceptual-discrimination difficulties, whose relation to reading impairment is disputed. We proposed that automatic detection and usage of serial sound regularities for individuals with dyslexia is impaired (anchoring deficit hypothesis), leading to the formation of less reliable sound predictions. Agus, Carrión-Castillo, Pressnitzer, and Ramus, (2014) reported seemingly contradictory evidence by showing similar performance by participants with and without dyslexia in a demanding auditory task that contained task-relevant regularities. To carefully assess the sensitivity of participants with dyslexia to regularities of this task, we replicated their study. Method: Thirty participants with and 24 without dyslexia performed the replicated task. On each trial, a 1-s noise stimulus was presented. Participants had to decide whether the stimulus contained repetitions (was constructed from a 0.5-s noise segment repeated twice) or not. It is implicit in this structure that some of the stimuli with repetitions were themselves repeated across trials. We measured the ability to detect within-noise repetitions and the sensitivity to cross-trial repetitions of the same noise stimuli. Results: We replicated the finding of similar mean performance. However, individuals with dyslexia were less sensitive to the cross-trial repetition of noise stimuli and tended to be more sensitive to repetitions in novel noise stimuli. Conclusion: These findings indicate that online auditory processing for individuals with dyslexia is adequate but their implicit retention and usage of sound regularities is indeed impaired.

Dyslexics' faster decay of implicit memory for sounds and words is manifested in their shorter neural adaptation.

Dyslexia is a prevalent reading disability whose underlying mechanisms are still disputed. We studied the neural mechanisms underlying dyslexia using a simple frequency-discrimination task. Though participants were asked to compare the two tones in each trial, implicit memory of previous trials affected their responses. We hypothesized that implicit memory decays faster among dyslexics. We tested this by increasing the temporal intervals between consecutive trials, and by measuring the behavioral impact and ERP responses from the auditory cortex. Dyslexics showed a faster decay of implicit memory effects on both measures, with similar time constants. Finally, faster decay of implicit memory also characterized the impact of sound regularities in benefitting dyslexics' oral reading rate. Their benefit decreased faster as a function of the time interval from the previous reading of the same non-word. We propose that dyslexics' shorter neural adaptation paradoxically accounts for their longer reading times, since it reduces their temporal window of integration of past stimuli, resulting in noisier and less reliable predictions for both simple and complex stimuli. Less reliable predictions limit their acquisition of reading expertise.

Neurophysiological mechanisms of cortical plasticity impairments in schizophrenia and modulation by the NMDA receptor agonist D-serine.

Schizophrenia is associated with deficits in cortical plasticity that affect sensory brain regions and lead to impaired cognitive performance. Here we examined underlying neural mechanisms of auditory plasticity deficits using combined behavioural and neurophysiological assessment, along with neuropharmacological manipulation targeted at the N-methyl-D-aspartate type glutamate receptor (NMDAR). Cortical plasticity was assessed in a cohort of 40 schizophrenia/schizoaffective patients relative to 42 healthy control subjects using a fixed reference tone auditory plasticity task. In a second cohort (n = 21 schizophrenia/schizoaffective patients, n = 13 healthy controls), event-related potential and event-related time-frequency measures of auditory dysfunction were assessed during administration of the NMDAR agonist d-serine. Mismatch negativity was used as a functional read-out of auditory-level function. Clinical trials registration numbers were NCT01474395/NCT02156908 Schizophrenia/schizoaffective patients showed significantly reduced auditory plasticity versus healthy controls (P = 0.001) that correlated with measures of cognitive, occupational and social dysfunction. In event-related potential/time-frequency analyses, patients showed highly significant reductions in sensory N1 that reflected underlying impairments in θ responses (P < 0.001), along with reduced θ and ß-power modulation during retention and motor-preparation intervals. Repeated administration of d-serine led to intercorrelated improvements in (i) auditory plasticity (P < 0.001); (ii) θ-frequency response (P < 0.05); and (iii) mismatch negativity generation to trained versus untrained tones (P = 0.02). Schizophrenia/schizoaffective patients show highly significant deficits in auditory plasticity that contribute to cognitive, occupational and social dysfunction. d-serine studies suggest first that NMDAR dysfunction may contribute to underlying cortical plasticity deficits and, second, that repeated NMDAR agonist administration may enhance cortical plasticity in schizophrenia.