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shows plots derived from SPM data analysis (one-sample t-tests) (standard approach) and 388 plots representing active region sizes using avareged individual data (alternative approach). Visual 389 inspection provides the following observations, among others. The tone with CF=400 Hz produced larger 390 active regions in SNHL than NH in both hemispheres. Size of activation for the 800Hz CF cortical 391 representations were similar in both groups. For the 1600 Hz CF stimulation, the region sizes were smaller 392 in patients, especially with the standard SPM approach and at lower statistical thresholds. In addition, 393 for 1600 Hz CF , the curve representing the relationship between the t-value and the activation size sloped 394 very rapidly at low t-thresholds until there was no activation detected in SNHL for the t-value 395 corresponding to p=0.05 FWE. Furthermore, some activation was detected in the patient group for high396 frequency stimulation (3200 Hz CF and 6400 Hz CF ) only when averaged activation sizes were looked at. 397 This shows that there were few patients in the group who were able to hear the sounds. This effect was 398 not present when the standard one-sample t-test was used as implemented in SPM, as only group 399 overlapping areas are revealed here. Also in both approaches very similar areas were revealed for pairs 400 of tones, i.e. 400 Hz CF and 800 Hz CF , 1600 Hz CF and 3200 Hz CF in NH. In SNHL, however, the 401 representations of these pairs of sounds were not overlapping in neither approach. 402 403 

shows plots derived from SPM data analysis (one-sample t-tests) (standard approach) and 388 plots representing active region sizes using avareged individual data (alternative approach). Visual 389 inspection provides the following observations, among others. The tone with CF=400 Hz produced larger 390 active regions in SNHL than NH in both hemispheres. Size of activation for the 800Hz CF cortical 391 representations were similar in both groups. For the 1600 Hz CF stimulation, the region sizes were smaller 392 in patients, especially with the standard SPM approach and at lower statistical thresholds. In addition, 393 for 1600 Hz CF , the curve representing the relationship between the t-value and the activation size sloped 394 very rapidly at low t-thresholds until there was no activation detected in SNHL for the t-value 395 corresponding to p=0.05 FWE. Furthermore, some activation was detected in the patient group for high396 frequency stimulation (3200 Hz CF and 6400 Hz CF ) only when averaged activation sizes were looked at. 397 This shows that there were few patients in the group who were able to hear the sounds. This effect was 398 not present when the standard one-sample t-test was used as implemented in SPM, as only group 399 overlapping areas are revealed here. Also in both approaches very similar areas were revealed for pairs 400 of tones, i.e. 400 Hz CF and 800 Hz CF , 1600 Hz CF and 3200 Hz CF in NH. In SNHL, however, the 401 representations of these pairs of sounds were not overlapping in neither approach. 402 403 

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Although the tonotopic organisation of the human primary auditory cortex (PAC) has already been studied, the question how its responses are affected in sensorineural hearing loss remains open. Twenty six patients (aged 38.1 ± 9.1 years; 12 men) with symmetrical sloping sensorineural hearing loss (SNHL) and 32 age- and gender-matched controls (NH) p...

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Currently, clinical characterization of hearing deficits for hearing-aid fitting is based on the pure-tone audiogram. Implicitly, this assumes that the audiogram can predict performance in complex, supra-threshold tasks. Sanchez-Lopez et al. (2018) hypothesized that the hearing deficits of a given listener, both at threshold and supra-threshold lev...

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... Given that ARHL begins in the high-frequency region of the auditory spectrum and spreads toward the low-frequency regions with age (Wang and Puel, 2020), the steep slope of the hearing threshold at high frequency is a very typical characteristic of patients with ARHL (Wolak et al., 2017). Accordingly, we used frequency division and the "steepness" of the audiogram to further refine the high-frequency region in this study. ...
... Previous study has found that patients with sensorineural hearing loss showed significantly reduced functional connectivity in the cingulate gyrus (Xu et al., 2019). Meanwhile, it has also been found that the hearing threshold of patients with ARHL began to decline rapidly at 2-4 kHz (Wolak et al., 2017). In addition, we found a significant positive correlation between the GMV of MCC and anxiety scores. ...
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... Due to the increased SNR and spatial resolution at 7T, fMRI studies focusing on each of the auditory relays separately have consistently reported that the cochlear frequency decomposition is conveyed as a clear, specific frequency gradient ranging from low to high frequencies in all the different subcortical auditory structures (collicular nuclei and Superior Olivary Complex: [1], inferior colliculi: [3,10] and medial geniculate bodies: [3]), until the auditory cortices. In hearing impaired humans, the damages concomitant with hearing loss or Meniere's disease at the level of the cochlea has been reported to induce a broadening of the auditory filters at the cochlear level and then an enlargement of the cortical representations near the "lesionedge" frequency (i.e. the frequency lost at the cochlear level) [9,11]. However, none of the aforementioned studies above investigated simultaneously the tonotopic gradient reorganization within the other auditory relays. ...
... Note that the functional results presented here are only descriptive, as the scope of this study was to evaluate the functional activation within all relays simultaneously using the pTx system and not their individual frequency mappings. Nevertheless, the results indicate potential for clinical applications as functional activations of the auditory pathway in less than 12 min were measured, compared to classical fMRI sessions ranging from 24 min [11] to~80 min [3]. Further studies will have to (a) adapt the MR sequences to match SAR requirements, (b) improve the anatomical localization of the subcortical regions using finer alignments with post mortem data or with functional atlases from other studies, (c) acquire a larger group of healthy subjects to quantify individual variability with specific analysis, and (d) compare the results between different auditory stimuli (pure tones vs. natural sounds) and different clinical populations. ...
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... Decreased frequency-modulation detection, in turn, is associated with reduced speech-in-noise perception (Holmes and Griffiths 2019;Parthasarathy et al. 2020). Moreover, it has been reported that severe hearing loss leads to tonotopic map reorganization in human auditory cortex (Wolak et al. 2017;Koops et al. 2020); however, this finding is not consistent across studies (Saenz and Langers 2014;Ouda et al. 2015). Tonotopic map reorganization (Mühlnickel et al. 1998) and broadened frequency-tuning (Sekiya et al. 2017) have also been observed in individuals with tinnitus (albeit inconsistently; e.g., Koops et al. 2020). ...
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... One example of a spectacular use of the fMRI technique is to gauge the tonotopic organisation of the auditory cortex ( Figure 4). The frequency-specific organisation of the cortex has been shown to exactly mirror sound frequency encoding in the cochlea [69][70][71][72][73][74][75][87][88][89][90]. ...
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... Thus, it is possible that the differences observed by Koops et al. between hearing-impaired subjects with and without tinnitus (2020) reflect differences in the peripheral pattern of the hearing loss (Tan et al., 2013) rather than changes arising centrally as a consequence of hearing loss. Consistent with this idea, earlier studies, which used constant stimulus levels, found either no changes in cortical responses associated with hearing loss (Ghazaleh et al., 2017;Lanting et al., 2008;Wolak et al., 2017) or increased responses at lower frequencies (Ghazaleh et al., 2017), opposite to the findings of Koops et al. (2020). Importantly, research in humans is complicated by the heterogeneity of tinnitus presentation, and likely also of the underlying causes and mechanisms (reviewed in Cederroth et al., 2019). ...
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... Ferrucci, 2011) and a higher risk for developing dementia and neurodegenerative disorders (Gates et al., 2010;Golub, 2017;Lin et al., 2011). Previous blood oxygen level dependent (BOLD) functional MRI (fMRI) studies have shown a reduced cortical activation in response to tonal stimulation in the tonotopic areas of the auditory cortex (Langers, van Dijk, Schoenmaker, & Backes, 2007;Zhang, Geng, Zhang, Li, & Zhang, 2006), especially at middle and higher frequencies (3.2 and 6.4 kHz) (Wolak et al., 2017), substantially mirroring the peripheral impairment of HL patients onto the auditory central perception. More recently, BOLD-fMRI studies conducted without stimulations (resting state; see, e.g., Chen et al., 2018) have similarly highlighted a significant reduction of spontaneous neural activity in the primary and secondary auditory cortex of HL patients which correlated with the reduced cognitive performances (including, e.g., language processing). ...
... The steeply sloping high frequency loss is a highly typical audiological pattern of the age-related HL condition, which is sometimes also called "sloping" sensori-neural HL (see, e.g., Wolak et al., 2017). ...
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... SŁOWA KLUCZOWE: czynnościowy rezonans magnetyczny, kora słuchowa, intensywność dźwięku BACKGROUND Functional magnetic resonance imaging (fMRI) has proven a valuable tool for studying central mechanisms of auditory perception. A number of works have been published which use this method to examine the relationship between sound parameters, including frequency and intensity, and activation patterns in the auditory cortex [1][2][3]. Notably, it has been shown that tonotopic organisation is preserved throughout the whole auditory system. This means that the systematic tuning of the basilar membrane in the inner ear, with filters tuned sequentially in frequency, is also reflected in the auditory cortex. ...
... Similar outcomes have been presented in fMRI studies by Hart and collaborators (2002) [6] and Sigalovsky and Melcher (2006) [9] who showed the same location of activity for a given frequency range at increased intensity levels with simultaneous spatial expansion of active regions. The high-frequency gradient forming a Vshape around the HG, revealed here for intensities 60 dB and 80 dB, has been reported in numerous papers using the fMRI technique to study auditory cortical responses [1,2,14,18,19,24,25,[27][28][29][30][31]. ...
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Background Despite rapid developments in fMRI, there is still ongoing debate on the optimal paradigm for evaluating the level of auditory cortex activation. Material and Methods A number of modern neuroimaging methods can be used to assess brain responses to acoustic stimulation, but new paradigms are still needed. Here the sparse fMRI approach is used to examine frequency-specific activation in auditory cortex in 12 normal hearing individuals. Results The size of activation expanded with increasing sound intensity and decreasing sound frequency. At the same time, the main site of frequency-specific activation remained the same across intensities, indicating fixed tonotopic organization. The findings of the study are explained in terms of basilar membrane phenomena such as the travelling wave pattern and spread of activation. Conclusions Stimulation levels of at least 60 dB are necessary in order to obtain robust maps of group activation in auditory cortex.
Chapter
Hearing loss is common among normally aging adults and often arises from damage to auditory peripheral structures, including cochlear hair cell loss and a degradation of synapses connecting hair cells to auditory nerves. However, it has become increasingly clear that age-related hearing loss involves dysfunction of the entire auditory system, including cortex. Peripheral damage induces neuroplastic changes in auditory cortex, most drastically reflected in a loss of neural inhibition that occurs in addition to a broader age-related decrease in inhibition. Here we review the impact of this combined loss of inhibition in auditory cortex on auditory processing. We discuss how reduced inhibition leads to hyperexcitability, reduced neural adaptation, altered periodicity processing, and changes in spectral and spatial tuning. We suggest that these functional changes, resulting from a loss of inhibition, underlie the hearing difficulties commonly experienced by older people, such as tinnitus, hyperacusis, and impaired speech understanding in noisy environments.
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Background Sensory deprivation, such as hearing loss, has been demonstrated to change the intrinsic functional connectivity (FC) of the brain, as measured with resting-state functional magnetic resonance imaging (rs-fMRI). Patients with sloping sensorineural hearing loss (SNHL) are a unique population among the hearing impaired, as they have all been exposed to some auditory input throughout their lifespan and all use spoken language. Materials and Methods Twenty patients with SNHL and 21 control subjects participated in a rs-fMRI study. Whole-brain seed-driven FC maps were obtained, with audiological scores of patients, including hearing loss severity and speech performance, used as covariates. Results Most profound differences in FC were found between patients with prelingual (before language development, PRE) vs. postlingual onset (after language development, POST) of SNHL. An early onset was related to enhancement in long-range network connections, including the default-mode network, the dorsal-attention network and the fronto-parietal network, as well as in local sensory networks, the visual and the sensorimotor. A number of multisensory brain regions in frontal and parietal cortices, as well as the cerebellum, were also more internally connected. We interpret these effects as top-down mechanisms serving optimization of multisensory experience in SNHL with a prelingual onset. At the same time, POST patients showed enhanced FC between the salience network and multisensory parietal areas, as well as with the hippocampus, when they were compared to those with PRE hearing loss. Signal in several cortex regions subserving visual processing was also more intra-correlated in POST vs. PRE patients. This outcome might point to more attention resources directed to multisensory as well as memory experience. Finally, audiological scores correlated with FC in several sensory and high-order brain regions in all patients. Conclusion The results show that a sloping hearing loss is related to altered resting-state brain organization. Effects were shown in attention and cognitive control networks, as well as visual and sensorimotor regions. Specifically, we found that even in a partial hearing deficit (affecting only some of the hearing frequency ranges), the age at the onset affects the brain function differently, pointing to the role of sensitive periods in brain development.