Arithmetic in the signing brain: Differences and similarities in arithmetic processing between deaf signers and hearing non-signers.

Deaf signers and hearing non-signers have previously been shown to recruit partially different brain regions during simple arithmetic. In light of the triple code model, the differences were interpreted as relating to stronger recruitment of the verbal system of numerical processing, that is, left angular and inferior frontal gyrus, in hearing non-signers, and of the quantity system of numerical processing, that is, right horizontal intraparietal sulcus, for deaf signers. The main aim of the present study was to better understand similarities and differences in the neural correlates supporting arithmetic in deaf compared to hearing individuals. Twenty-nine adult deaf signers and 29 hearing non-signers were enrolled in an functional magnetic resonance imaging study of simple and difficult subtraction and multiplication. Brain imaging data were analyzed using whole-brain analysis, region of interest analysis, and functional connectivity analysis. Although the groups were matched on age, gender, and nonverbal intelligence, the deaf group performed generally poorer than the hearing group in arithmetic. Nevertheless, we found generally similar networks to be involved for both groups, the only exception being the involvement of the left inferior frontal gyrus. This region was activated significantly stronger for the hearing compared to the deaf group but showed stronger functional connectivity with the left superior temporal gyrus in the deaf, compared to the hearing, group. These results lend no support to increased recruitment of the quantity system in deaf signers. Perhaps the reason for performance differences is to be found in other brain regions not included in the original triple code model.

Working Memory for Signs with Poor Visual Resolution: fMRI Evidence of Reorganization of Auditory Cortex in Deaf Signers.

Stimulus degradation adds to working memory load during speech processing. We investigated whether this applies to sign processing and, if so, whether the mechanism implicates secondary auditory cortex. We conducted an fMRI experiment where 16 deaf early signers (DES) and 22 hearing non-signers performed a sign-based n-back task with three load levels and stimuli presented at high and low resolution. We found decreased behavioral performance with increasing load and decreasing visual resolution, but the neurobiological mechanisms involved differed between the two manipulations and did so for both groups. Importantly, while the load manipulation was, as predicted, accompanied by activation in the frontoparietal working memory network, the resolution manipulation resulted in temporal and occipital activation. Furthermore, we found evidence of cross-modal reorganization in the secondary auditory cortex: DES had stronger activation and stronger connectivity between this and several other regions. We conclude that load and stimulus resolution have different neural underpinnings in the visual-verbal domain, which has consequences for current working memory models, and that for DES the secondary auditory cortex is involved in the binding of representations when task demands are low.

Arithmetic in the adult deaf signing brain.

We have previously shown that deaf signers recruit partially different brain regions during simple arithmetic compared to a group of hearing non-signers, despite similar performance. Specifically, hearing individuals show more widespread activation in brain areas that have been related to the verbal system of numerical processing, i.e., the left angular and inferior frontal gyrus, whereas deaf individuals engaged brain areas that have been related to the quantity system of numerical processing, i.e., the right horizontal intraparietal sulcus. This indicates that compared to hearing non-signers, deaf signers can successfully make use of processes located in partially different brain areas during simple arithmetic. In this study, which is a conceptual replication and extension of the above-presented study, the main aim is to understand similarities and differences in neural correlates supporting arithmetic in deaf compared to hearing individuals. The primary objective is to investigate the role of the right horizontal intraparietal gyrus, the left inferior frontal gyrus, the hippocampus, and the left angular gyrus during simple and difficult arithmetic and how these regions are connected to each other. A second objective is to explore what other brain regions support arithmetic in deaf signers. Up to 34 adult deaf signers and the same amount of hearing non-signers will be enrolled in an functional magnetic resonance imaging study that will include simple and difficult subtraction and multiplication. Brain imaging data will be analyzed using whole-brain analysis, region of interest analysis and connectivity analysis. This is the first study to investigate neural underpinnings of arithmetic of different difficulties in deaf individuals.

The Organization of Working Memory Networks is Shaped by Early Sensory Experience.

Early deafness results in crossmodal reorganization of the superior temporal cortex (STC). Here, we investigated the effect of deafness on cognitive processing. Specifically, we studied the reorganization, due to deafness and sign language (SL) knowledge, of linguistic and nonlinguistic visual working memory (WM). We conducted an fMRI experiment in groups that differed in their hearing status and SL knowledge: deaf native signers, and hearing native signers, hearing nonsigners. Participants performed a 2-back WM task and a control task. Stimuli were signs from British Sign Language (BSL) or moving nonsense objects in the form of point-light displays. We found characteristic WM activations in fronto-parietal regions in all groups. However, deaf participants also recruited bilateral posterior STC during the WM task, independently of the linguistic content of the stimuli, and showed less activation in fronto-parietal regions. Resting-state connectivity analysis showed increased connectivity between frontal regions and STC in deaf compared to hearing individuals. WM for signs did not elicit differential activations, suggesting that SL WM does not rely on modality-specific linguistic processing. These findings suggest that WM networks are reorganized due to early deafness, and that the organization of cognitive networks is shaped by the nature of the sensory inputs available during development.

fMRI Evidence of Magnitude Manipulation during Numerical Order Processing in Congenitally Deaf Signers.

Congenital deafness is often compensated by early sign language use leading to typical language development with corresponding neural underpinnings. However, deaf individuals are frequently reported to have poorer numerical abilities than hearing individuals and it is not known whether the underlying neuronal networks differ between groups. In the present study, adult deaf signers and hearing nonsigners performed a digit and letter order tasks, during functional magnetic resonance imaging. We found the neuronal networks recruited in the two tasks to be generally similar across groups, with significant activation in the dorsal visual stream for the letter order task, suggesting letter identification and position encoding. For the digit order task, no significant activation was found for either of the two groups. Region of interest analyses on parietal numerical processing regions revealed different patterns of activation across groups. Importantly, deaf signers showed significant activation in the right horizontal portion of the intraparietal sulcus for the digit order task, suggesting engagement of magnitude manipulation during numerical order processing in this group.

Geometry in the brain optimized for sign language - A unique role of the anterior superior parietal lobule in deaf signers.

Geometry has been identified as a cognitive domain where deaf individuals exhibit relative strength, yet the neural mechanisms underlying geometry processing in this population remain poorly understood. This fMRI study aimed to investigate the neural correlates of geometry processing in deaf and hearing individuals. Twenty-two adult deaf signers and 25 hearing non-signers completed a geometry decision task. We found no group differences in performance, while there were some differences in parietal activation. As expected, the posterior superior parietal lobule (SPL) was recruited for both groups. The anterior SPL was significantly more activated in the deaf group, and the inferior parietal lobule was significantly more deactivated in the hearing group. In conclusion, despite similar performance across groups, there were differences in the recruitment of parietal regions. These differences may reflect inherent differences in brain organization due to different early sensory and linguistic experiences.

Reorganization of large-scale brain networks in deaf signing adults: The role of auditory cortex in functional reorganization following deafness.

If the brain is deprived of input from one or more senses during development, functional and structural reorganization of the deprived regions takes place. However, little is known about how sensory deprivation affects large-scale brain networks. In the present study, we use data-driven independent component analysis (ICA) to characterize large-scale brain networks in 15 deaf early signers and 24 hearing non-signers based on resting-state functional MRI data. We found differences between the groups in independent components representing the left lateralized control network, the default network, the ventral somatomotor network, and the attention network. In addition, we showed stronger functional connectivity for deaf compared to hearing individuals from the middle and superior temporal cortices to the cingulate cortex, insular cortex, cuneus and precuneus, supramarginal gyrus, supplementary motor area, and cerebellum crus 1, and stronger connectivity for hearing non-signers to hippocampus, middle and superior frontal gyri, pre- and postcentral gyri, and cerebellum crus 8. These results show that deafness induces large-scale network reorganization, with the middle/superior temporal cortex as a central node of plasticity. Cross-modal reorganization may be associated with behavioral adaptations to the environment, including superior ability in some visual functions such as visual working memory and visual attention, in deaf signers.

Associations Between Sign Language Skills and Resting-State Functional Connectivity in Deaf Early Signers.

The processing of a language involves a neural language network including temporal, parietal, and frontal cortical regions. This applies to spoken as well as signed languages. Previous research suggests that spoken language proficiency is associated with resting-state functional connectivity (rsFC) between language regions and other regions of the brain. Given the similarities in neural activation for spoken and signed languages, rsFC-behavior associations should also exist for sign language tasks. In this study, we explored the associations between rsFC and two types of linguistic skills in sign language: phonological processing skill and accuracy in elicited sentence production. Fifteen adult, deaf early signers were enrolled in a resting-state functional magnetic resonance imaging (fMRI) study. In addition to fMRI data, behavioral tests of sign language phonological processing and sentence reproduction were administered. Using seed-to-voxel connectivity analysis, we investigated associations between behavioral proficiency and rsFC from language-relevant nodes: bilateral inferior frontal gyrus (IFG) and posterior superior temporal gyrus (STG). Results showed that worse sentence processing skill was associated with stronger positive rsFC between the left IFG and left sensorimotor regions. Further, sign language phonological processing skill was associated with positive rsFC from right IFG to middle frontal gyrus/frontal pole although this association could possibly be explained by domain-general cognitive functions. Our findings suggest a possible connection between rsFC and developmental language outcomes in deaf individuals.

Effects of mild-to-moderate sensorineural hearing loss and signal amplification on vocal emotion recognition in middle-aged-older individuals.

Previous research has shown deficits in vocal emotion recognition in sub-populations of individuals with hearing loss, making this a high priority research topic. However, previous research has only examined vocal emotion recognition using verbal material, in which emotions are expressed through emotional prosody. There is evidence that older individuals with hearing loss suffer from deficits in general prosody recognition, not specific to emotional prosody. No study has examined the recognition of non-verbal vocalization, which constitutes another important source for the vocal communication of emotions. It might be the case that individuals with hearing loss have specific difficulties in recognizing emotions expressed through prosody in speech, but not non-verbal vocalizations. We aim to examine whether vocal emotion recognition difficulties in middle- aged-to older individuals with sensorineural mild-moderate hearing loss are better explained by deficits in vocal emotion recognition specifically, or deficits in prosody recognition generally by including both sentences and non-verbal expressions. Furthermore a, some of the studies which have concluded that individuals with mild-moderate hearing loss have deficits in vocal emotion recognition ability have also found that the use of hearing aids does not improve recognition accuracy in this group. We aim to examine the effects of linear amplification and audibility on the recognition of different emotions expressed both verbally and non-verbally. Besides examining accuracy for different emotions we will also look at patterns of confusion (which specific emotions are mistaken for other specific emotion and at which rates) during both amplified and non-amplified listening, and we will analyze all material acoustically and relate the acoustic content to performance. Together these analyses will provide clues to effects of amplification on the perception of different emotions. For these purposes, a total of 70 middle-aged-older individuals, half with mild-moderate hearing loss and half with normal hearing will perform a computerized forced-choice vocal emotion recognition task with and without amplification.

The cognitive hearing science perspective on perceiving, understanding, and remembering language: The ELU model.

The review gives an introductory description of the successive development of data patterns based on comparisons between hearing-impaired and normal hearing participants' speech understanding skills, later prompting the formulation of the Ease of Language Understanding (ELU) model. The model builds on the interaction between an input buffer (RAMBPHO, Rapid Automatic Multimodal Binding of PHOnology) and three memory systems: working memory (WM), semantic long-term memory (SLTM), and episodic long-term memory (ELTM). RAMBPHO input may either match or mismatch multimodal SLTM representations. Given a match, lexical access is accomplished rapidly and implicitly within approximately 100-400 ms. Given a mismatch, the prediction is that WM is engaged explicitly to repair the meaning of the input - in interaction with SLTM and ELTM - taking seconds rather than milliseconds. The multimodal and multilevel nature of representations held in WM and LTM are at the center of the review, being integral parts of the prediction and postdiction components of language understanding. Finally, some hypotheses based on a selective use-disuse of memory systems mechanism are described in relation to mild cognitive impairment and dementia. Alternative speech perception and WM models are evaluated, and recent developments and generalisations, ELU model tests, and boundaries are discussed.

Evidence of an Effect of Gaming Experience on Visuospatial Attention in Deaf but Not in Hearing Individuals.

Auditory cortex in congenitally deaf early sign language users reorganizes to support cognitive processing in the visual domain. However, evidence suggests that the potential benefits of this reorganization are largely unrealized. At the same time, there is growing evidence that experience of playing computer and console games improves visual cognition, in particular visuospatial attentional processes. In the present study, we investigated in a group of deaf early signers whether those who reported recently playing computer or console games (deaf gamers) had better visuospatial attentional control than those who reported not playing such games (deaf non-gamers), and whether any such effect was related to cognitive processing in the visual domain. Using a classic test of attentional control, the Eriksen Flanker task, we found that deaf gamers performed on a par with hearing controls, while the performance of deaf non-gamers was poorer. Among hearing controls there was no effect of gaming. This suggests that deaf gamers may have better visuospatial attentional control than deaf non-gamers, probably because they are less susceptible to parafoveal distractions. Future work should examine the robustness of this potential gaming benefit and whether it is associated with neural plasticity in early deaf signers, as well as whether gaming intervention can improve visuospatial cognition in deaf people.

Working memory, deafness and sign language.

Working memory (WM) for sign language has an architecture similar to that for speech-based languages at both functional and neural levels. However, there are some processing differences between language modalities that are not yet fully explained, although a number of hypotheses have been mooted. This article reviews some of the literature on differences in sensory, perceptual and cognitive processing systems induced by auditory deprivation and sign language use and discusses how these differences may contribute to differences in WM architecture for signed and speech-based languages. In conclusion, it is suggested that left-hemisphere reorganization of the motion-processing system as a result of native sign-language use may interfere with the development of the order processing system in WM.

Influence of environmental enrichment on steady-state mRNA levels for EAAC1, AMPA1 and NMDA2A receptor subunits in rat hippocampus.

Interaction with the environment has a key role in refining the neuronal circuitry required for normal brain function throughout life. Profound effects of enriched environment have been shown on neuronal structure and chemistry in experimental animals. Epidemiological studies imply that this is true also in man, thus cognitive stimulation has a protective effect on neurodegeneration, e.g., in Alzheimer's disease. Glutamatergic pathways are imperative for cognitive functions, such as memory, learning and long-term potentiation, and relies on the AMPA and NMDA glutamate receptors and the hippocampus, with its specific subregions, is an important anatomical substrate in this. The glutamate signalling is also dependent on a fine-tuned transport system, in the hippocampus primarily achieved by the glutamate transporter EAAC1. In this study we show how environmental enrichment modulates these parts of the glutamatergic system using quantitative in situ hybridisation. This work demonstrates for the first time that environmental enrichment modulates the mRNA expression of EAAC1 which is significantly and region specifically decreased in the hippocampus. We also provide evidence for regional and hemisphere-specific upregulation of NMDA mRNA in the hippocampus after environmental enrichment. The current work also shows that AMPA mRNA of the hippocampus is not per se changed by environmental enrichment in adult animals. Taken together, our results extend the knowledge of the glutamatergic system of specific regions of the hippocampus and its modulation by environmental enrichment and could contribute to the development of strategies aimed at limiting pathological changes associated with glutamatergic dysfunctions.

Phonology and arithmetic in the language-calculation network.

Arithmetic and language processing involve similar neural networks, but the relative engagement remains unclear. In the present study we used fMRI to compare activation for phonological, multiplication and subtraction tasks, keeping the stimulus material constant, within a predefined language-calculation network including left inferior frontal gyrus and angular gyrus (AG) as well as superior parietal lobule and the intraparietal sulcus bilaterally. Results revealed a generally left lateralized activation pattern within the language-calculation network for phonology and a bilateral activation pattern for arithmetic, and suggested regional differences between tasks. In particular, we found a more prominent role for phonology than arithmetic in pars opercularis of the left inferior frontal gyrus but domain generality in pars triangularis. Parietal activation patterns demonstrated greater engagement of the visual and quantity systems for calculation than language. This set of findings supports the notion of a common, but regionally differentiated, language-calculation network.

Modulation of neuronal glutamate transporter rEAAC1 mRNA expression in rat brain by amitriptyline.

Glutamate transporters regulate the glutamate concentration in the synaptic cleft within the CNS, a regulation required for normal brain function. In several neurological conditions, the amount of glutamate is altered. One reason for the changes in glutamate concentration might be impaired glutamate transporter function. In this study, an in situ hybridisation technique has been used to elucidate changes in mRNA expression of the glutamate transporter, excitatory amino acid carrier 1 (EAAC1), after treatment with the tricyclic antidepressant (TCA) amitriptyline. The results lead to the suggestion that treatment with tricyclic antidepressants leads to changes in the EAAC1 mRNA expression in rat brain suggesting involvement of the glutamate system in the tricyclic treatment of depression.

Similar digit-based working memory in deaf signers and hearing non-signers despite digit span differences.

Similar working memory (WM) for lexical items has been demonstrated for signers and non-signers while short-term memory (STM) is regularly poorer in deaf than hearing individuals. In the present study, we investigated digit-based WM and STM in Swedish and British deaf signers and hearing non-signers. To maintain good experimental control we used printed stimuli throughout and held response mode constant across groups. We showed that deaf signers have similar digit-based WM performance, despite shorter digit spans, compared to well-matched hearing non-signers. We found no difference between signers and non-signers on STM span for letters chosen to minimize phonological similarity or in the effects of recall direction. This set of findings indicates that similar WM for signers and non-signers can be generalized from lexical items to digits and suggests that poorer STM in deaf signers compared to hearing non-signers may be due to differences in phonological similarity across the language modalities of sign and speech.

Rivastigmine as a modulator of the neuronal glutamate transporter rEAAC1 mRNA expression.

Alzheimer's disease is a neurodegenerative disorder that affects the cholinergic, glutamatergic and monoaminergic systems in the neocortex and hippocampus. Today, the major pharmacological treatment involves the use of acetylcholinesterase inhibitors (AChEIs). In this study, an in situ hybridisation technique (using digoxigenin-labelled cRNA probes) was used to elucidate changes in mRNA expression of the neuronal glutamate transporter, rat excitatory amino carrier 1 (rEAAC1), after treatment with the AChEI rivastigmine. Compared with saline-treated rats, the rats subchronically (3 days) and chronically (21 days), but not acutely, treated with rivastigmine showed a significant increase in rEAAC1 mRNA expression in the hippocampal areas cornu anterior 1 (CA1), CA2, CA3 and dentate gyrus (p < 0.01), but not in the cortical areas. These results provide the first evidence that the glutamatergic system is modulated following acetylcholinesterase inhibition by rivastigmine, a finding, which is likely to be of importance for the clinical effects.