Evidence suggests that large-scale brain network organization may differ between individuals with limited sensory input (e.g. deaf individuals) and those without sensory impairment. These neural differences may manifest as behavioural differences, such as enhanced visual attention in deaf individuals. Using independent component analyses (ICA) on resting-state fMRI data, we showed that the attention network was more widespread in the hearing compared to the deaf group. This suggests that the organization of the attention network is different in deaf and hearing adults. 

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. Twentytwo 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.

Few studies have examined which acoustic features of speech can be used to distinguish between different emotions, and how combinations of acoustic parameters contribute to identification of emotions. The aim of the present study was to investigate which acoustic parameters in Swedish speech are most important for differentiation between, and identification of, the emotions anger, fear, happiness, sadness, and surprise in Swedish sentences. One-way ANOVAs were used to compare acoustic parameters between the emotions and both simple and multiple logistic regression models were used to examine the contribution of different acoustic parameters to differentiation between emotions. Results showed differences between emotions for several acoustic parameters in Swedish speech: surprise was the most distinct emotion, with significant differences compared to the other emotions across a range of acoustic parameters, while anger and happiness did not differ from each other on any parameter. The logistic regression models showed that fear was the best-predicted emotion while happiness was most difficult to predict. Frequency- and spectral-balance-related parameters were best at predicting fear. Amplitude- and temporal-related parameters were most important for surprise, while a combination of frequency-, amplitude- and spectral balance-related parameters are important for sadness. Assuming that there are similarities between acoustic models and how listeners infer emotions in speech, results suggest that individuals with hearing loss, who lack abilities of frequency detection, may compared to normal hearing individuals have difficulties in identifying fear in Swedish speech. Since happiness and fear relied primarily on amplitude- and spectral-balance-related parameters, detection of them are probably facilitated more by hearing aid use.

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.

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.

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, previousresearch has only examined vocal emotion recognition using verbal material, in which emotions are expressed through emotional prosody. There is evidence that older individualswith hearing loss suffer from deficits in general prosody recognition, not specific to emotionalprosody. No study has examined the recognition of non-verbal vocalization, which constitutes another important source for the vocal communication of emotions. It might be thecase that individuals with hearing loss have specific difficulties in recognizing emotionsexpressed through prosody in speech, but not non-verbal vocalizations. We aim to examinewhether 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 sentencesand non-verbal expressions. Furthermore a, some of the studies which have concluded thatindividuals with mild-moderate hearing loss have deficits in vocal emotion recognition abilityhave also found that the use of hearing aids does not improve recognition accuracy in thisgroup. We aim to examine the effects of linear amplification and audibility on the recognitionof different emotions expressed both verbally and non-verbally. Besides examining accuracy for different emotions we will also look at patterns of confusion (which specific emotionsare mistaken for other specific emotion and at which rates) during both amplified and nonamplified 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 onthe perception of different emotions. For these purposes, a total of 70 middle-aged-olderindividuals, half with mild-moderate hearing loss and half with normal hearing will perform acomputerized forced-choice vocal emotion recognition task with and without amplification

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.

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.

Stimulus degradation adds to working memory load during speech processing.We investigated whether this applies to signprocessing and, if so, whether the mechanism implicates secondary auditory cortex.We conducted an fMRI experimentwhere 16 deaf early signers (DES) and 22 hearing non-signers performed a sign-based n-back task with three load levels andstimuli presented at high and low resolution.We found decreased behavioral performance with increasing load anddecreasing visual resolution, but the neurobiological mechanisms involved differed between the two manipulations and didso for both groups. Importantly, while the load manipulation was, as predicted, accompanied by activation in thefrontoparietal 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 strongeractivation and stronger connectivity between this and several other regions.We conclude that load and stimulus resolutionhave different neural underpinnings in the visual–verbal domain, which has consequences for current working memorymodels, and that for DES the secondary auditory cortex is involved in the binding of representations when task demandsare low.

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.

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.

Deafness is generally associated with poor mental arithmetic, possibly due to neuronal differences in arithmetic processing across language modalities. Here, we investigated for the first time the neuronal networks supporting arithmetic processing in adult deaf signers. Deaf signing adults and hearing non-signing peers performed arithmetic and phonological tasks during fMRI scanning. At whole brain level, activation patterns were similar across groups. Region of interest analyses showed that although both groups activated phonological processing regions in the left inferior frontal gyrus to a similar extent during both phonological and multiplication tasks, deaf signers showed significantly more activation in the right horizontal portion of the inferior parietal sulcus. This region is associated with magnitude manipulation along the mental number line. This pattern of results suggests that deaf signers rely more on magnitude manipulation than hearing non-signers during multiplication, but that phonological involvement does not differ significantly between groups.Abbreviations: AAL: Automated Anatomy Labelling; fMRI: functional magnetic resonance imaging; HIPS: horizontal portion of the intraparietal sulcus; lAG: left angular gyrus; lIFG: left inferior frontal gyrus; rHIPS: right horizontal portion of the intraparietal sulcus

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.

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.

Studies of sign languages have been used to test traditional cognitive models of working memory (WM) that distinguish between verbal and visuospatial WM (e.g. Baddeley, 2003), without considering that sign languages operate in the visuospatial domain. Previous studies have shown that WM mental representations and processes are largely similar for signed and spoken languages (e.g. Rönnberg et al., 2004). However, it is not clear to what extent visual WM processes aid and support sign language WM.

Here we characterise the neural substrates supporting sign language and visual WM, and the mechanisms that subserve differential processing for signers and for deaf individuals. We conducted a functional magnetic resonance imaging (fMRI) experiment with three groups of participants: deaf native signers, hearing native signers and hearing non-signers. Participants performed a 2-back WM task and a control task on two sets of stimuli: signs from British Sign Language or non-sense objects. Stimuli were composed of point-lights to control for differences in visual features.

Our results show activation in a fronto-parietal network for WM processing in all groups, independently of stimulus type, in agreement with previous literature. We also replicate previous findings in deaf signers showing a stronger right posterior superior temporal cortex (STC) activation for visuospatial processing, and stronger bilateral STC activation for sign language stimuli.

Group comparisons further reveal stronger activations in STC for WM in deaf signers, but not for the groups of hearing individuals. This activation is independent of the linguistic content of the stimuli, being observed in both WM conditions: signs and objects. These results suggest a cognitive role for STC in deaf signers, beyond sign language processing.

Semantic coherence and prior knowledge enhance perceptual clarity of degraded speech. Recent study by our team has shown that these two effects interact such that the perceptual clarity of noise-vocoded speech (NVS) is still enhanced by semantic coherence when prior knowledge is available from text cues and prior knowledge enhances perceptual clarity of NVS even when semantic coherence is low (Signoret et al., 2015). Here, we investigated the neural correlates of this interaction. We predicted 1) an effect of matching cues for both sentences with high and low semantic coherence in left-lateralized perisylvian areas (Zekveld et al., 2012) and right superior temporal gyrus (Wild et al., 2012), but stronger for low than for high coherent sentences since more resources are required to process sentences with low semantic coherence in the left inferior frontal gyrus (Obleser and Kotz, 2010) and 2) an effect of semantic coherence in temporal and inferior frontal cortex (Lau et al., 2008). The additive effect of semantic coherence when matching cues were provided should be observed in the angular gyrus (Obleser and Kotz, 2010). Twenty participants (age; M=25.14, SD=5.01) listened to sentences and performed an unrelated attentional task during sparse-imaging fMRI. The sentences had high or low semantic coherence, and were either clear, degraded (6-band NV) or unintelligible (1-band NV). Each spoken word was preceded (200 ms) by either a matching cue or a consonant string. Preliminary results revealed significant main effects of Cue (F(1,228) = 21.26; p < .05 FWE) in the left precentral gyrus, the left inferior frontal gyrus and the left middle temporal gyrus confirming the results of Zekveld et al (2012), but neither the main effect of Coherence nor the interaction between Cue and Coherence survived FWE correction. In accordance with our predictions, contrasts revealed a greater effect of matching cues for low than for high coherent sentences (t(19) = 6.25; p < .05 FWE) in the left superior temporal gyrus as well as left inferior frontal gyrus (BA 44 and 45), suggesting greater involvement of both top-down and bottom-up processing mechanisms during integration of prior knowledge with the auditory signal when sentence coherence is lower. There was a marginally greater effect of semantic coherence (t(19) = 3.58; p < .001unc) even when matching cues were provided in the left angular gyrus, the left middle frontal gyrus and the right superior frontal gyrus, suggesting greater involvement of top-down activation of semantic concepts, executive processes and the phonological store during integration of prior knowledge with the auditory signal when the semantic content of the speech is more readily available.

Evidence suggests that the lag reported in mathematics for deaf signers derives from difficulties related to verbal processing of numbers, whereas magnitude processing seems unaffected by deafness. Neuroimaging evidence from hearing individuals suggests that verbal processing of numbers engages primarily left angular gyrus (lAG), whereas magnitude processing engages primarily the horizontal portion of the right intraparietal sulcus (rHIP). In a ROI analysis of brain imaging data from 16 adult deaf signers and 16 adult hearing non-signers, who did not differ on sex, age or education, we examined if activity in lAG and rHIP changed as a result of task (multiplication vs subtraction) and group (deaf signers and hearing non-signers). We found a significant main effect of brain region (F(1,30) = 117.00, p < .001, η_p^2 = .80) and an interaction effect between region and group (F(1,30) = 20.70, p < .001, η_p^2 = .41). Further analyses showed that there were no significant differences in average activation between groups in lAG (F(1,30) = 0.16, p = .70). However, in rHIP deaf signers showed significantly greater average activation compared to non-signers (F(1,30) = 15.20, p < .001, η_p^2 = .34). There were no significant differences in activation between subtraction and multiplication (F(1,30) = 0.66, p = .42) and no behavioural differences between groups (F(1,30) = 1.70, p = .20). These results suggest that when engaging in arithmetic tasks deaf signers successfully make use of qualitatively difference processes, compared to hearing non-signers, with stronger emphasis on brain regions relating to magnitude manipulation.

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. (C) 2015 The Authors. Published by Elsevier Inc.

Degraded speech is rendered more intelligible both by semantic coherence and preceding text cues. Recently, we showed that the perceptual clarity of noise-vo-coded speech (NVS) is still enhanced by semantic coherence when cues are provided and that prior knowledge enhances perceptual clarity of NVS when semantic coherence is low (Signoret et al., 2015). Here, we investigated the neural correlates of this interaction. Twenty participants listened to sentences and performed an unrelated attentional task during sparse-imaging fMRI. The sentences had high or low semantic coherence, and were either clear, degraded (6-band NV) or unintelligible (1-band NV). Each spoken word was preceded (200 ms) by either a matching cue or a consonant string. Preliminary results revealed significant main effects of both Coherence and Cue in the superior temporal gyrus bilaterally and a significant interaction between Coherence and Cue when speech was degraded, in superior and middle temporal gyri bilaterally and left precentral gyrus. Investigation of this interaction revealed greater activation for high compared to low coherent sentences when cues were provided in the left-lateralized regions and greater activation without than with cues when semantic coherence was low in bilateral regions. The opposite contrasts elicited no significant activation. This pattern of results indicates that the increases in perceptual clarity of NVS attributable to semantic coherence and prior knowledge are supported by similar neural mechanisms organized in bilateral temporal regions, but that when perceptual clarity is optimized by both factors, it is supported by left-lateralized mechanisms.

Evidence suggests that the lag reported in mathematics for deaf signers derives from difficulties related to the verbal system of number processing as described in the triple code model. For hearing individuals the verbal system has been shown to be recruited for both arithmetic and language tasks. In the present study we investigate for the first time neuronal representations of arithmetic in deaf signers. We examine if the neural network supporting arithmetic and language, including the horizontal portion of the intraparietal sulcus (HIPS), the superior parietal lobule (SPL) bilaterally, the left angular gyrus (AG), pars opercularis (POPE) and pars triangularis (PTRI) of the left inferior frontal gyrus (IFG), is differently recruited for deaf and hearing individuals. Imaging data were collected from 16 deaf signers and 16 well-matched hearing nonsigners, using the same stimulus material for all tasks, but with different cues. During multiplication, deaf signers recruited rHIPS more than hearing non-signers, suggesting greater involvement of magnitude manipulation processes related to the quantity system, whereas there was no evidence that the verbal system was recruited. Further, there was no support for the notion of a common representation of phonology for sign and speech as previously suggested.

Deaf students generally lag several years behind hearing peers in arithmetic, but little is known about the mechanisms behind this. In the present study we investigated how phonological skills interact with arithmetic. Eighteen deaf signers and eighteen hearing non-signers took part in an experiment that manipulated arithmetic and phonological knowledge in the language modalities of sign and speech. Independent tests of alphabetical and native language phonological skills were also administered. There was no difference in performance between groups on subtraction, but hearing non-signers performed better than deaf signers on multiplication. For the deaf signers but not the hearing non-signers, multiplicative reasoning was associated with both alphabetical and phonological skills. This indicates that deaf signing adults rely on language processes to solve multiplication tasks, possibly because automatization of multiplication is less well established in deaf adults.

Dövhet har kopplats till bristande förmåga att hantera siffror inom områdena aritmetik och minne. Särskilt har språkmodalitetsspecifika skillnader i fonologisk likhet för siffror visat sig påverka korttidsminnet. Det övergripande syftet med den här avhandlingen var därför att undersöka om språkmodalitetsspecifika skillnader i fonologisk bearbetning mellan teckenoch talspråk kan förklara varför döva presterar sämre än hörande på sifferuppgifter. För att utforska det området undersöktes fonologisk bearbetning i sifferbaserade minnesuppgifter och aritmetik med hjälp av både beteendevetenskapliga metoder och hjärnavbildning hos grupper av teckenspråkiga döva och talspråkiga hörande som matchats noggrant på ålder, kön, utbildning och icke-verbal intelligens. För att testförhållandena skulle bli så likartade som möjligt för de båda grupperna, och för att förebygga materialeffekter, användes samma presentations- och svarssätt för båda grupperna. Resultaten visade att vid sifferbaserat korttidsminne påverkas de dövas prestation av de tecknade siffrornas fonologiska likhet. Däremot fanns det ingen skillnad mellan grupperna gällande sifferbaserat arbetsminne, vilket kan bero på att de båda grupperna fördelar sina kognitiva resurser på olika sätt. Dessutom fann vi att den grupp teckenspråkiga döva som deltog i studien presterade bättre på aritmetik än vad tidigare forskning visat och de skiljde sig bara från hörande på multiplikationsuppgifter, vilket kan bero på att grupperna var så noggrant matchade. Däremot fanns det skillnader mellan grupperna i vilka neurobiologiska nätverk som aktiverades vid aritmetik och fonologi. Vid multiplikationsuppgifter aktiverades cortex i höger parietallob och vänster frontallob för de teckenspråkiga döva, medan cortex i vänster frontal- och parietallob aktiverades för de talspråkiga hörande. Detta indikerar att de talspråkiga hörande förlitar sig på fonologiberoende minnesstrategier medan de teckenspråkiga döva förlitar sig på ickeverbal magnitudmanipulering och artikulatoriska processer. Under den fonologiska uppgiften aktiverade de talspråkiga hörande vänsterlateraliserade frontala och parietala områden inom det klassiska språknätverket. För de teckenspråkiga döva var fonologibearbetningen begränsad till cortex i vänster occipitallob, vilket tyder på att teckenspråksbaserad fonologi inte behöver aktivera det klassiska språknätverket. Sammanfattningsvis visar fynden i den här avhandlingen att språkmodalitetsspecifika skillnader mellan tecken- och talspråk på olika sätt kan förklara varför döva presterar sämre än hörande på vissa sifferbaserade uppgifter.

The literacy skills of deaf children generally lag behind those of their hearing peers. The mechanisms of reading in deaf individuals are only just beginning to be unraveled but it seems that native language skills play an important role. In this study 12 deaf pupils (six in grades 1?2 and six in grades 4?6) at a Swedish state primary school for deaf and hard of hearing children were trained on the connection between Swedish Sign Language and written Swedish using a pilot sign language version of the literacy training software program Omega-is. Literacy skills improved substantially across the 20 days of the study. These literacy gains may have rested upon the specific software-based intervention, upon regular classroom activities, or upon a combination of these factors. Omega-is-d, and similar software utilizing sign language as a component, targets an important mechanism supporting reading development in deaf children and could play an important role in bilingual education refinements.

Profoundly deaf individuals sometimes have difficulty with arithmetic and phonological tasks. In the present study we investigate if these differences can be attributed to differences in recruitment of neurobiological networks. Seventeen hearing non-signers (HN) and sixteen deaf signers (DS) matched on age, gender and non-verbal intelligence took part in an fMRI study. In the scanner three digit/letter pairs were visually presented and the participants performed six different blocked tasks tapping processing of digit and letter order, multiplication, subtraction and phonological ability. Data were analysed using two 2x2x2 ANOVAs; process (arithmetic, language) x level (high, low) x group (DS, HN). A main effect of process revealed language networks in the left inferior frontal gryus, supramarginal gyrus, fusiform gyrus and insula. Arithmetic networks included left middle orbital gyrus and superior medial gyrus. A main effect of level revealed low level processing (digit/letter order) in the right middle occipital gyrus and the right precuneus and high level processing (subtraction/multiplication/phonological ability) in left inferior frontal gyrus. There was no main effect of group but a significant task x group interaction in the right temporal pole which in DS (but not HN) was activated more for arithmetic than language processing (p_fwe = .022) when multiplication was included in the analysis. This region is implicated in conceptual representation. These results suggest that both arithmetic and language are processed similarly by DS and HN with possible between-group differences in the use of conceptual representation in arithmetic and language tasks.

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.

In hearing individuals, multiplication relies mainly on the phonological loop while subtraction relies on the visuo-spatial sketchpad (VSSP; Lee & Kang, 2002). Little is known about arithmetic neural networks in deaf signers (DS). Since DS often perform worse than hearing non-signers (NH) on arithmetic in general and multiplication in particular (Traxler, 2000), we hypothesized that there are strategic differences between how groups recruit the phonological loop in multiplication, but not in subtraction, leading to differential activation of phonological processing areas in left inferior frontal gyrus (Broca’s area). We investigated this using a blocked fMRI-design in which nine DS and 17 HN matched on age, gender, education and non-verbal intelligence (Raven & Raven, 1998) were tested on tasks of multiplication, subtraction and phonology (rhyme). The contrasts rhyme versus multiplication and rhyme versus subtraction were examined across groups within the region of interest defined by a probability map of Broca’s area (Amunts, 1999). We observed a significant interaction between task (multiplication and rhyme) and group (F = 12.64, p = .034, FWE-corrected), where the HN showed higher activation for rhyme than for multiplication (T = 4.55, p = .001, FWE-corrected) whereas there were no differences in activations between tasks for DS. For subtraction versus rhyme no interaction with group was found. These results suggest that there are differences between DS and HN in the phonology dependent neural networks in Broca’s area used during multiplication, which may be part of the explanation for poorer performance in DS.

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 Objectives Working memory (WM) capacity is similar for signed (SL) and spoken (SpL) language yet underlying temporal and spatial processing mechanisms may not be identical. To investigate this, two studies with deaf native signers (DS) and hearing non-signers (HN) were conducted. Methods DS and matched HN groups performed WM tasks with varying temporal and spatial demands in study 1 at encoding (temporal, spatial and mixed presentation styles) and in study 2 at retrieval (forward and backward span) and with abstract spatial demands (math span). Results DS performance was inferior with high temporal demands at encoding (temporal style) and retreival (forward span). There was no difference between groups with high spatial order demands at encoding (spatial style) or retrieval (backward span). DS performance was worse when abstract spatial processing was involved (math span). Conclusion WM processing mechanisms for SL and SpL differ for temporal information at encoding and retrieval and for abstract spatial information. 

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.

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. © 2007.

Glutamate is the principal excitatory neurotransmitter in the central nervous system and it is implicated in neural transmission, learning, memory processes and neuronal plasticity. In the glutamatergic synapse two main components are present; the glutamate receptors and the glutamate transporters. The receptors, the NMDA, AMPA, kainite and the metabotroptic receptors, are responsible for conveying neural transmission, including long term potentiation (LTP), synaptic strengthening and modification. The transporters, located to the neuronal membrane and to the membranes of surrounding astrocytes, regulates the extracellular concentration of glutamate and thereby the duration of the synaptic signal.

Alterations in both receptor and transporter systems have been suggested to be important in the pathogenesis of several acute and chronic nervous system diseases, such as psychosis, mood disorders, epilepsy, Parkinson's disease and Alzheimer's disease. The pathophysiology of these disorders is not yet completely understood and the involvement of glutamate is unclear. In this thesis we have sought to investigate the role of the glutamatergic system in the treatment of mood disorders and dementia. The antidepressant drug amitriptyline exerts its main effects on the serotonergic and noradrenergic systems and the antidementia drug rivastigmine acts mainly on the cholinergic system. However, given the close relationship between different neurotransmitter systems we have investigated the influence of amitriptyline and rivastigmine on the mRNA expression of the neuronal transporter, EAAC1, in rats. The results showed for the first time an involvement of EAAC1 in amitriptyline and rivastigmine treatment. Amitriptyline induced an acute increase in EAAC1 mRNA expression, which 24 hour after administration returned to baseline levels. Chronic treatment, on the other hand, induces a significant decrease in cortical areas, which we suggest results in enhanced neuronal transmission. Rivastigmine treatment, acute as well as chronic, induced increases in the mRNA expression in hippocampus. We hypothesize that this counteracts the excitotoxic glutamate levels seen in Alzheimer's disease.

Further, environmental enrichment has been shown to have beneficial effects on capillary supply, the number of glial cells and dendritic spines, the thickness and weight of cortex, the concentration of cholinesterase, LTP and synaptic strength in animals. It has also been reported that humans that lead an active life have a reduced risk of developing Alzheimer's disease. This suggests that an active and stimulated life may have a protective effect against dementia in man, by creating a cognitive reserve which provides a buffer against brain pathology or age-related changes. We investigated the influence of environmental enrichment on the mRNA expression of NMDA and AMPA receptors and on EAACl and showed for the first time that EAAC1 mRNA is decreased after environmental enrichment. This is probably followed by an increase of glutamate in the synapse, which in turn leads to enhanced neuronal transmission including enhanced memory formation and learning. Furthermore, we confirmed in greater detail previous findings on the upregulation of NMDA mRNA and show that the regulation is regionally and hemisphere specific. We also confirm that AMPA mRNA is not per se changed by environmental enrichment in adult animals.

This work provides further evidence about the involvement of the glutamatergic system in affective and cognitive disorders. Improved knowledge of the glutamatergic system will contribute to the development of strategies aimed at limiting pathological changes associated with glutamatergic dysfunctions.

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.