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Author: admin | 22.05.2014 | Category: Tinnitus Causes And Treatments

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Tinnitus is the perception of a sound in the absence of a corresponding external sound source. Non-pulsatile tinnitus is the perception of a sound in the absence of a corresponding external sound source. Based on phenomenological similarities it has been proposed that tinnitus is the auditory analog of phantom pain (1–4) with auditory deafferentation representing its causative factor. A second complementary concept proposes that deafferentiation-induced alterations in the central auditory pathways are not sufficient for the conscious perception of tinnitus. Recently, the concept has been forwarded that tinnitus might be the expression of a global workspace hyperactivity rather than being limited to the auditory system (22), which has been extended in a heuristic model integrating the deafferentiation model, the noise-cancelling deficit, and the global workspace model. What all models, the deafferentation neurophysiological model, the noise-cancelling deficit, and the multiple parallel overlapping dynamical network model have in common, is that they rely on a final common pathway called TCD (39) (see Figure 1, yellow box). Tinnitus is perceived as a unified percept, incorporating a sound, which can be different in quality, location, loudness, constancy, duration, as well as affective components such as mood and distress. Gamma-band activity (>30 Hz) is important for motor control (55) and processing of sensory stimuli, whether in the olfactory bulb (56), visual (57), auditory (58), or somatosensory cortex (59). Based on the principle that gamma-band activity is associated with the conscious percept of auditory stimuli (58, 66, 82), Llinas proposed that tinnitus, as a conscious percept, should be associated with persistent gamma-band activity in the auditory cortex (39) confirmed in latter Ca V2.1-null experiments (81). An explanation for the occurrence of this theta–gamma coupled activity in sensory deafferentation is provided by the concept of TCD (39, 85).
How is this TCD generated at a cellular level and what mechanism can link TCD to a more widespread phenomenon in the brain? It has been proposed that the tectal system, with its rapid multisensory input, organizes the instantaneous orienting responses crucial to the maintenance of life (95).
In summary, in the non-bursting regions subthreshold oscillations are capable of performing a filtering task without increasing neuronal output. Gamma-band activity in the auditory cortex is a prerequisite for auditory conscious perception and, therefore, likely also contributes to the perception of a phantom sound (Table 1). When checked, Shutterstock's safe search screens restricted content and excludes it from your search results.
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In this study, we describe differences between neural plasticity in auditory cortex (AC) of animals that developed subjective tinnitus (group T) after noise-induced hearing loss (NIHL) compared to those that did not [group non-tinnitus (NT)].
Despite these severe impairments and the large number of patients affected, there is still no established treatment available to safely and effectively alleviate tinnitus.
Although short or acute tinnitus percepts are experienced occasionally by virtually everyone, chronic tinnitus develops only in a sub-population of humans as well as in the different animal model systems. We have recently published the first report comparing neuronal plasticity following NIHL in animals that develop a tinnitus percept (group T) with plastic changes in animals that do not [group non-tinnitus (NT)], even though both animal groups did not differ in NIHL (17). Based on these results we have proposed a model for tinnitus prevention that points to a global inhibitory mechanism in auditory cortex (AC) that may prevent tinnitus genesis in animals with high overall activity in the auditory system, whereas this mechanism does not seem to be available for tinnitus prevention in animals with low overall activity. In the present follow-up study we present a further, more detailed analysis compared to the Ahlf et al.
A total of 35 eight to ten weeks old male gerbils purchased from Charles River Laboratories, Inc. All animals were handled before the beginning of the experiments and accustomed to the setup environment to minimize stress.
The setups for behavioral and electrophysiological recordings have been described in detail earlier (17, 18). Three different types of PPI modulated ASR paradigms (17) were performed to assess, first, hearing capacities [behavioral and reflex based audiogram (19)] and, second, the potential existence of a tinnitus percept (20) after the noise trauma. For tinnitus testing we used two modified ASR paradigms in all animals before and after trauma.
Of the 35 animals investigated behaviorally, 20 were used for electrophysiological recording in brainstem and AC.
We statistically analyzed behavioral and neuronal response amplitude data as well as FFT frequency spectra by parametric tests; response latencies and durations were assessed by non-parametric tests as one cannot assume normal distribution for these parameters. Our patented on-ear design lies so flat, Bedphones practically disappear between your ears and the pillow, helping you fall asleep fasterOur earhooks are made from gentle, rubber-coated memory wire, INFINITELY ADJUSTABLE for a custom fit. Add our optional in-line microphone with single-buttom remote to answer phone calls, play, pause, and skip music tracks without touching your phone. This means that you will not need to remember your user name and password in the future and you will be able to login with the account you choose to sync, with the click of a button. This page doesn't support Internet Explorer 6, 7 and 8.Please upgrade your browser or activate Google Chrome Frame to improve your experience.
As an auditory phantom phenomenon, it has been conceptualized to be either linked to bottom-up deafferentation (see Figure 1, green box) with or without behaviorally measurable hearing loss, or a deficient top-down noise-cancelling mechanism (see Figure 1, blue box) or a combination of both. Many subsequent models are refinements of this concept and limit themselves to changes in the auditory system, zooming in on the specific aspects linked to auditory deprivation, such as hyperactivity (5), an increase in gain (6), plasticity (7), including map plasticity (8) and homeostatic plasticity (9–11), synchrony (5, 12), filling the missing information (3, 13, 14), or sensitization (15). Tinnitus only arises, if auditory deafferentiation is accompanied by a deficient inhibitory top-down mechanism (19, 20), analogous to the anti-nociceptive system deficit linked to some forms of spontaneous pain such as fibromyalgia (21).
The global workspace model was first proposed by Baars (23) as a model for conscious cognitive processing.
However, the question arises, how to theoretically integrate TCD in the setting of consciousness as an emergent property of network activity (31, 40) and more specifically in tinnitus (41). It is assumed that each of these characteristics is related to a separate subnetwork, but linked by hubs that are part of multiple subnetworks (41).
In physiological sound processing, sound triggers gamma-band activity in the auditory cortex. Indeed, it has been shown both by EEG (83), MEG (39, 84, 85), and intracranial recordings (46) that tinnitus is related to gamma-band activity in the auditory cortex and nested on theta activity (39, 46, 84). By contrast, the thalamocortical system, with its extensive corticothalamic recurrence and temporal binding properties, creates the complex combinatorial sensorimotor activities required for intentional behavior, characteristic of vertebrate behavior (95). Thus, T-type calcium channels can be considered first-responders to small depolarization (100), generating a low-threshold Ca2+ potential upon the crest of which sodium and potassium channel-mediated action potentials fire, creating low-threshold bursts (101). Contrastingly, bursting regions may spread network oscillations and recruit interconnected cortical areas, while simultaneously acting as a theta frequency filter (100). Fitted into soft, durable, silicone earpieces which can be worn under protective head gear. To this end, our analysis focuses on the input activity of cortical neurons based on the temporal and spectral analysis of local field potential (LFP) recordings and an in-depth analysis of auditory brainstem responses (ABR) in the same animals. The development of effective therapies for tinnitus is mainly hampered by the fact that the exact neuronal mechanisms leading to the phantom percept and its chronic manifestation are still largely unknown.
We currently do not know why some patients with hearing impairments develop chronic tinnitus while others with the same shift in hearing threshold do not, and vice versa why some but not all subjects without detectable noise-induced hearing loss (NIHL) still develop tinnitus, although first models that may solve this enigma have been published (14–17). As we could demonstrate, NT animals showed higher overall auditory cortical and brainstem activity before noise trauma compared to T-animals. We suggest that by activating this global inhibitory mechanism the NT animals may actively counteract the development of maladaptive increased response gains in AC seen in T-animals that possibly reflect the tinnitus development. During the first week, animals were tested first, with three different paradigms of pre-pulse inhibition (PPI) of the acoustic startle reflex (ASR) to evaluate the audiograms for each animal and acquire baseline data for the tinnitus percept evaluation.
The measurements were performed before and usually 7 days after the acoustic trauma in a single 4 h block. Mean PPI effect (±95% confidence interval) in six exemplary animals classified as non-tinnitus (NT) and tinnitus (T) perceiving. Root mean square (RMS) values of the LFP recordings in AC with 70 dB SPL stimulation intensity. Signals were recorded using a Plexon Multichannel Acquisition Processor (Plexon Inc., Dallas, TX, USA) after amplification by a JHM NeuroAmp 401 (J. Among these, the complete recording protocol in AC could be accomplished in four T and two NT animals. Wear the cable down your back and tighten with the cable slider to keep Bedphones secure all nightBedphones are the PERFECT TRAVEL HEADPHONES.
Both mechanisms are proposed to alter auditory thalamocortical signal transmission, resulting in thalamocortical dysrhythmia (TCD).
Many of these models are still a matter of debate, such as the involvement of synchrony (16), or whether changes in gain explain really tinnitus and not only hyperacusis.


It is a model for the interface between multiple unconscious, parallel processing modules on one side, and conscious experience on the other side, proposing how different sources of information are integrated into one percept with internally consistent content. Furthermore, there are common associated symptoms in tinnitus, such as hearing loss, which can also generate distress (42, 43), and as distress is generated by a non-specific brain network (3, 44–49) this can be perceived or confounded as tinnitus distress. It is likely that different forms or subtypes of tinnitus exist, each with a different pathophysiology, which could explain the interindividual differences seen in recent electrophysiological studies (53, 54).
The externally presented sound only reaches consciousness if consciousness enabling network (=global workspace) is co-activated with sound perception. Whereas, initially, it was thought that the gamma-band activity was contralateral (83, 84), it has been shown that gamma-band activity is increased in both auditory cortices, even in unilateral tinnitus (86).
Conceptually, this can be explained that the firing rate decreases when less information needs to be processed (91) due to the deafferentation, and that firing and oscillation rate are coupled at the thalamocortical level (85). It has been proposed that this theta could then act as a long-range carrier wave (96–98) on which the tinnitus information can be nested by means of high-frequency oscillatory activity (41).
Thalamic neurons, on being depolarized from resting potential levels positive to −55 mV, fire tonically. As such theta burst-firing increases network synchrony and recruitment (100), a mechanism, which might enable long-range synchrony, which in turn could represent a means for finding the missing thalamocortical information (due to deafferentation) (105).
It can be conceived that the theta activity is the carrier wave connecting widespread areas (72), and that focal gamma-band activity in geographically separated brain areas is synchronized by nesting on the theta phase, as exemplified in auditory attention (106). In response to NIHL in NT animals we find a significant general reduction in overall cortical activity and spectral power as well as changes in all ABR wave amplitudes as a function of loudness.
Hence a significant amount of data from both human (6, 7) and animal studies (8–12) on the plastic changes occurring in the involved neuronal networks during tinnitus development is available, a number of different and in part controversial neurobiological models trying to explain the phenomenon do exist. We concluded that animals with low overall neuronal activity in the auditory system might have a predisposition for the development of tinnitus early after noise trauma while animals with higher neuronal activity did not develop such a percept within the time of observation. According to this model, the high overall AC activity of NT animals before the trauma is a prerequisite for the activation of this hypothetical global inhibition. The use and care of animals was approved by the state of Bavaria (Regierungspräsidium Mittelfranken, Ansbach, Germany). All methods used in this paper (except the LFP recordings) have been described previously (17) but still will be explained in detail here for better intelligibility.
Second, ABR measurements of the healthy animals were performed to obtain individual audiograms under anesthesia and the far field responses of the auditory pathway in the brainstem up to the inferior colliculus (IC). For obtaining the behavioral hearing thresholds we used a PPI of ASR paradigm in all animals. Keep them safe on-the-go with the included sturdy-sided travel case and ONE-YEAR MANUFACTURER'S WARRANTY. In deafferentation, TCD is characterized by a slowing down of resting state alpha to theta activity associated with an increase in surrounding gamma activity, resulting in persisting cross-frequency coupling between theta and gamma activity. Also, the memory-based filling in mechanism is still a rather speculative model with limited experimental evidence.
The global workspace model suggests that multiple, highly specialized; quasi-independent input processors compete for access to a broadcasting capability by which the winning processor can disseminate its information globally throughout the brain.
The lateralization of the tinnitus percept depends on gamma-band activity in the parahippocampal area, associated with functional connectivity between the parahippocampal area and auditory cortex. This could suggest that gamma reflects a difference, a prediction error (68), a detection of change to the reference (69), or status quo, the latter, which would be reflected by beta activity (70). The last sound presentation is stored in the (para)hippocampal area as a reference for future input to calculate the prediction error. The amount of gamma-band activity has been shown to be related to the subjectively perceived tinnitus loudness (83). As a result, GABAA-mediated lateral inhibition is reduced (85), inducing gamma (>30 Hz) band activity (39) surrounding the deafferented theta area, also known as the edge effect (85). This notion has been confirmed by a MEG study, which demonstrated increased coupled gamma–theta wave activity in tinnitus patients (39, 84, 85) and recently demonstrated in EEG as well (99).
When a burst is generated in a thalamocortical neuron anywhere on the dendrite or cell body, the low-threshold Ca2+ potential is conducted not only to the soma but also back-propagates throughout the entire dendritic tree (100), generating bursts in multiple postsynaptic neurons in the projected nucleus through axonal arborization to multiple synapses.
In contrast, T-animals show no significant change in overall cortical activity as assessed by root mean square analysis of LFP amplitudes, but a specific increase in LFP spectral power and in the amplitude of ABR wave V reflecting activity in the inferior colliculus (IC). Furthermore, within a week after the acoustic trauma T-animals showed increased activity in auditory cortical neurons representing the tinnitus frequencies, whereas NT animals exhibited decreased activity at moderate sound intensities at that point in time. We suggest that a certain dynamic range for unspecific gain reduction across the whole AC is needed to successfully counteract the response gain increase that is associated with tinnitus development. The authors declare that there is no conflict of interests regarding the publication of this paper. After these measurements surgery for implantation of the head holder and the recording chamber over the left AC was performed and electrophysiological single electrode recordings in primary AC field AI of the healthy anesthetized animal were carried out on two to three sessions within 1 week after the surgery.
We startled the animals with 90 dB SPL pure tones (6 ms length including 2 ms rise and fall ramps) ranging from 0.5 to 16 kHz in a pseudo-randomized order in octave steps.
0) and no changes in PPI effect were found comparing pre (blue) and post trauma (red) data by t-tests. The recordings were identical to those described in the earlier report (17) allowing us to additionally analyze single unit responses within AC. And for long rides, they also come with a comfy satin eye maskBedphones now come with an OPTIONAL INLINE MICROPHONE with remote - a necessity for the many who use Bedphones as their all-day everyday headphones. Theta burst-firing increases network synchrony and recruitment, a mechanism, which might enable long-range synchrony, which in turn could represent a means for finding the missing thalamocortical information and for gaining access to consciousness. Moreover, some models focus on somatosensory compensatory mechanisms in the dorsal cochlear nucleus (17), which in itself has been proposed to be a core region in the development of tinnitus (18). In general, gamma-band activity is present only in locally restricted areas of the cortex for short periods of time (34, 35, 71, 72). A decrease in tinnitus loudness has been found to be associated with a decrease in gamma-band activity in the auditory cortex (87–89) and a worsening in tinnitus loudness with an increase in auditory cortex gamma-band activity (90), even if such a relationship has been called into question by a recent study (54). Indeed, in tinnitus, a decrease in alpha power is associated with an increase in gamma power (92), and gamma goes together with theta activity (84), confirming the initial studies by Llinas et al.
These bursting neurons will in turn recruit additional neurons in the reciprocally connected initiating nucleus and this burst amplification will progress on each cycle of the oscillation, generating synchronization in a network of reciprocally connected CNS nuclei (103, 104). The synchrony would permit the (para)hippocampal information to be boosted to conscious perception by recruiting consciousness enabling network areas (41). If the overall neuronal activity in AC is already low before the trauma, as in the case of T-animals, this activity may not be reduced further and tinnitus develops because of un-resisted response gain increase in those regions of AC representing the tinnitus percept.
Acoustic trauma (2 kHz, 115 dB SPL, 75 min) under deep anesthesia was used to induce a mild NIHL (evaluated by ABR analysis directly after the trauma and PPI audiogram after 1 week) and a persistent tinnitus percept in the majority of the animals (26 of 35 animals, 74%; evaluated by PPI measurement one week after trauma). The same pure tones were used as pre-stimulus probes ranging from 0 to 50 dB SPL in 10 dB steps 100 ms before the startle stimulus.
In both cases, a silent 15 ms gap within the noise (white or band pass) 100 ms before the startle stimulus served as pre-pulse. The right panel gives the comparison of time of recording averaged over all frequencies as part of the 2-factorial ANOVA. Our single-buttom remote is compatible with most smartphones and allows you to answer phone calls, play, pause, and skip music tracks without touching your phoneBecause Bedphones lie against your ear without entering the ear canal, they are the PERFECT ALTERNATIVE TO UNCOMFORTABLE EARBUDS Description:Bedphones are the most comfortable headphones for sleep. Theta oscillations could function as a carrier wave to integrate the tinnitus-related focal auditory gamma activity in a consciousness enabling network, as envisioned by the global workspace model.
Any such global pattern can inhibit alternative activity patterns among workspace neurons, thus preventing the conscious processing of alternative stimuli and enabling a uniform conscious percept. Whereas normally gamma activity waxes and wanes, related to the presence of a novel external stimulus, persisting gamma activity localized in one brain area has been considered pathological (39) (see Figure 2). In this paper (54), gamma-band activity increases and decreases were noted associated with an increase in tinnitus loudness after residual inhibition or a decrease in loudness after residual excitation, respectively. This is in agreement with clinical data demonstrating that the dysrhythmia can be recorded with MEG over widespread areas (39). What remains an open question in our model is the source of the hypothetical global inhibition, whether it is generated intra-cortically (within AC), sub-cortically or even in higher cortical or limbic structures. Within the first week after the trauma all electrophysiological recordings in AC were performed in two to three recording sessions. The rationale of these paradigms is that an animal that perceives a tinnitus would be impaired in gap detection because it would hear its own tinnitus within the silent gap [cf. Stimulation was pseudo-randomized using a fixed list of all combinations of stimulus frequencies and sound pressure levels (0–90 dB SPL in 5 dB steps). This model suggests that focal activity in the brain does not reach consciousness, except if the focal activity becomes functionally coupled to a consciousness enabling network, aka the global workspace. Moreover, top-down attentional amplification is the mechanism by which modular processes can be temporarily mobilized and made available to the global workspace, and therefore to consciousness. However, in contrast to the waxing and waning of gamma-band activity, recent studies related to language processing do demonstrate there are also spontaneous endogenous longer lasting gamma rhythms present at rest in monkeys (73) and humans (74). This could reflect that gamma represents a homeostatic change in loudness per se, on the condition that the tinnitus state has become the reference, or it could potentially be attributed to other factors, such as associated hearing loss in these patients.
Other studies have since demonstrated the presence of both low- (delta or theta) and high-frequency activity in the auditory cortex of tinnitus patients (88, 89, 93, 94), and the coupled activity has also been confirmed on electrode recordings of implanted patients (46).


Thus, in short, thalamic neurons have the unique property to fire both at depolarization and at hyperpolarization.
But the burst-firing process may also act as a neuronal frequency filter, preventing the conduction of recent incoming postsynaptic activity to the soma, and thereby preventing other activities contributing to the neuronal output (99). All experiments were carried out in an IAC (Industrial Acoustics Company GmbH, Niederkrüchten, Germany) acoustic chamber on a TMC (Technical Manufacturing Corporation, Peabody, MA, USA) low-vibration table. For sound generation, the frequency response function of the speaker was calibrated to produce an output spectrum that was flat within ±1 dB measured within the acrylic tube. Note that we did not find any significant change over time and no interaction of the two predictors in these animals. To obtain ABR-based audiograms the mean ABR wave amplitudes were compared to the mean amplitude 200–100 ms before the stimulus (baseline).
In limited deafferentation, the missing information can be retrieved from the auditory cortical neighborhood, decreasing surround inhibition, resulting in TCD. These are likely generated by inhibitory interneurons (75) or interactions between pyramidal cells and interneurons (76, 77) and might reflect attentional processes (76, 77). The data obtained were checked by eye via a custom-made Matlab program, trials in which the animals moved within 100 ms before the startle stimulus were discarded; in the valid trials only peak-to-peak amplitudes of responses within the first 50 ms after startle stimulus onset were used for further analysis.
Animals were again ketamine-xylazine anesthetized, placed on a 37°C warm heating pad and fixated via the aluminum head-post. When the deafferentation is too wide in bandwidth, it is hypothesized that the missing information is retrieved from theta-mediated parahippocampal auditory memory.
The emergence of a unified percept relies on the coordination of scattered mosaics of functionally specialized brain regions (31).
These complement the exogenous stimulus driven gamma-band activity, which usually increases gamma-band activity in sensory areas, presumably clustering spiking activity that is propagated to higher hierarchic processing stages (74).
Finally, the exact relations between cochlear damage, hearing impairment, and tinnitus development are still elusive. The evaluation was performed independently by the principal investigator and a technical assistant, who was blind to the state of the animal. Consequently, when using a gap as pre-pulse, animals with tinnitus should produce smaller PPI of ASR than animals without tinnitus (cf. For additional analysis the root mean square (RMS) value of the ABR signal was calculated from 1 to 5 ms after stimulus onset and single waves of the ABR responses representing the different levels within the brainstem (25) were detected by a custom-made Matlab program at 70 dB SPL for each stimulation frequency.
This suggests that based on the amount of deafferentation TCD might change to parahippocampocortical persisting and thus pathological theta–gamma rhythm. This is proposed to be related to oscillatory activity and connectivity within the brain (32). Thus, it has been envisaged that these spontaneous gamma oscillations mechanistically might optimize the extraction of relevant sensory input in time (74).
From a Bayesian point of view, in which the brain is conceived as a prediction machine that updates its memory-based predictions through sensory updating, tinnitus is the result of a prediction error between the predicted and sensed auditory input. Large-scale integration has been proposed to bind the distributed anatomical and functional organization of brain activity enabling the emergence of coherent behavior and cognition.
We made sure that the animals were always (pre and post trauma = trauma status) responding to the 90 dB SPL startle stimulus (cf. In humans, it is known that the majority of tinnitus percepts (around 70%) exhibit a pure tone-like character while the remaining part consists of ringing tones, noises or even more complex sounds (22). The recorded LFPs were analyzed with custom-made Matlab programs and statistically evaluated with Statistica. Although the mechanisms involved in large-scale integration are still largely unknown, it has been convincingly argued that the most plausible candidate is the formation of dynamic links mediated by synchrony over multiple frequency bands (31). It has been proposed that oscillations correspond to the alternation of phases of high- and low-neuronal excitability, which temporally constrain sensory processing (78). We used the two different paradigms to determine possible tinnitus percepts in the animals because the exact nature of these percepts is unknown. Three response measures were evaluated, first, the amplitude and timing of the first negative peak (N1) and the second positive peak (P2) waves of the recorded data (reflecting mainly the slower extralemniscal and intracortical input [for review see Ref.
Thus, TCD can be considered as an adaptive mechanism to retrieve missing auditory input in tinnitus. Even though within-frequency phase synchronization may support the binding of anatomically distributed processing, it cannot coordinate neuronal processing distributed into distinct time windows or frequency bands (33). This means that gamma oscillations, which have a period of approximately 25 ms, provide a 10- to 15-ms window for integrating spectrotemporal information followed by a 10- to 15-ms window for propagating the output (74).
For validation of the PPI effect of the pre-stimuli we performed 1-factorial ANOVAs for the valid response amplitudes dependent on the pre-stimulus intensity for each frequency and trauma status separately for each individual animal. With the pure tone stimulus over a white noise and the click stimulus over band pass filtered noise we aimed to cover as many different percepts as possible. Thus, cross-frequency coupling might be important for large-scale integration via low-frequency coherence of distributed geographically focal high-frequency activity (34).
However, because the average length of a phoneme is about 50 ms, a 10- to 15-ms window might be too short for integrating this information, but a phoneme can correctly be represented by three gamma cycles, which act as a three-bit code (74).
The mean responses before and after trauma of the six animals used for electrophysiological recordings in AC are given in Figure S1 in Supplementary Material.
If a tinnitus is detected, the different tone frequencies and noise spectra used should give a rough estimate of the spectral content of the tinnitus percept.
Further studies should evaluate whether tinnitus could be represented by such a 3-bit gamma code.
Responses in this threshold paradigm were fitted with a sigmoidal Boltzmann function for each frequency, trauma status, and animal separately. Each frequency and gap-condition was repeated 15 times and each test was performed before and after the acoustic trauma. This notion is supported by very recent results demonstrating tinnitus-related cross-frequency-coupling (37). In the visual system, stimuli that reach consciousness and those which do not reach consciousness are characterized by a similar increase of local gamma oscillations in the visual cortex (79, 80). Hearing thresholds were defined as the sound level at the inflection point of the Boltzmann function for each frequency before and after trauma (18). Again the data obtained were checked by eye by the same two experimenters as above via a custom-made Matlab program.
Thus, gamma-band activity, per se, is not related to conscious perception, but could be a condition sine qua non, an essential prerequisite, for conscious perception (30).
Trials in which the animals moved within 100 ms before the startle stimulus were discarded; in the valid trials only peak-to-peak amplitudes of responses within the first 50 ms after startle stimulus onset were used for further analysis (17).
This approach allowed the determination of a possible frequency-specific tinnitus-related behavior at one octave precision. In tinnitus, intra-area cross-frequency coupling is known as thalamocortical dysrhythmia (TCD), and inter-area phase–amplitude coupling between anterior cingulate and auditory cortex and dorsolateral prefrontal cortex has been shown as well (37). Only when focal gamma activity is embedded in a larger network of long-range functional connectivity, this will lead to the emergence of a conscious percept (41) (see Figure 2).
After the trauma only some animals showed an undisturbed gap-effect at all frequencies tested while other animals showed no gap-effect at some but not all frequencies (cf.
Figure 1), which gave a first hint of a possible tinnitus percept but was not yet used as the final classification of the animals in the tinnitus or NT group (cf. To avoid possible effects of the acoustic trauma on different stimulation frequencies all startle response data were normalized to minimize variance of the response amplitudes. Normalization was performed as described earlier (17, 23); briefly, we divided each amplitude by the corresponding median amplitude of the 90 dB SPL only condition (which reflects the full startle response of the animal for the loudest condition at each specific frequency, 0 dB SPL pre-stimulus intensity Figure S1 in Supplementary Material). Thus we attempted to control for differences in the startle amplitudes resulting from the hearing loss at the trauma frequency.
This normalization also guarantees that the reduced ASR response after acoustic trauma is not due to hearing loss but likely due to a tinnitus percept (23). Significant positive values for PPI change reflect impaired PPI and therefore indicate the development of a tinnitus percept. Only such animals were therefore classified as probably perceiving a tinnitus (T group) that showed at least one impaired frequency after the trauma independent of the affected frequency itself. As it turned out, in all cases where a tinnitus was detected according to one of the two gap-ASR paradigms used, the second gap-ASR paradigm was also positive for tinnitus. Animals without such a significant increase in PPI change were classified as NT perceiving animals (NT group). It became apparent that animals classified as T or NT based on these behavioral measures also differed in neurophysiological response measures (ABR and AC responses; cf. While the tests used here may only detect acute tinnitus, exemplary animals were re-tested in regular intervals for up to 16 weeks and did not show any changes in tinnitus frequencies.
Based on these results we think that the classification of NT and T-animals is not only valid in acute but also in the chronic state of the disease.



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