Key points
A popular conception of mammalian cochlear physiology is that tuned mechanical vibration of the basilar membrane defines the frequency response of the innervating auditory nerve fibres
...However, the data supporting these concepts come from vibratory measurements at cochlear locations tuned to high frequencies (>7 kHz).
Here, we measured the travelling wave in regions of the guinea pig cochlea that respond to low frequencies (<2 kHz) and found that mechanical tuning was broad and did not match auditory nerve tuning characteristics.
Non‐linear amplification of the travelling wave functioned over a broad frequency range and did not substantially sharpen frequency tuning.
Thus, the neural encoding of low‐frequency sounds, which includes most of the information conveyed by human speech, is not principally determined by basilar membrane mechanics.
The popular notion of mammalian cochlear function is that auditory nerves are tuned to respond best to different sound frequencies because basilar membrane vibration is mechanically tuned to different frequencies along its length. However, this concept has only been demonstrated in regions of the cochlea tuned to frequencies >7 kHz, not in regions sensitive to lower frequencies where human speech is encoded. Here, we overcame historical technical limitations and non‐invasively measured sound‐induced vibrations at four locations distributed over the apical two turns of the guinea pig cochlea. In turn 3, the responses demonstrated low‐pass filter characteristics. In turn 2, the responses were low‐pass‐like, in that they occasionally did have a slight peak near the corner frequency. The corner frequencies of the responses were tonotopically tuned and ranged from 384 to 668 Hz. Non‐linear gain, or amplification of the vibrations in response to low‐intensity stimuli, was found both below and above the corner frequencies. Post mortem, cochlear gain disappeared. The non‐linear gain was typically 10–30 dB and was broad‐band rather than sharply tuned. However, the gain did reach nearly 50 dB in turn 2 for higher stimulus frequencies, nearly the amount of gain found in basal cochlear regions. Thus, our data prove that mechanical responses do not match neural responses and that cochlear amplification does not appreciably sharpen frequency tuning for cochlear regions that respond to frequencies <2 kHz. These data indicate that the non‐linear processing of sound performed by the guinea pig cochlea varies substantially between the cochlear apex and base.
Key points
A popular conception of mammalian cochlear physiology is that tuned mechanical vibration of the basilar membrane defines the frequency response of the innervating auditory nerve fibres
However, the data supporting these concepts come from vibratory measurements at cochlear locations tuned to high frequencies (>7 kHz).
Here, we measured the travelling wave in regions of the guinea pig cochlea that respond to low frequencies (<2 kHz) and found that mechanical tuning was broad and did not match auditory nerve tuning characteristics.
Non‐linear amplification of the travelling wave functioned over a broad frequency range and did not substantially sharpen frequency tuning.
Thus, the neural encoding of low‐frequency sounds, which includes most of the information conveyed by human speech, is not principally determined by basilar membrane mechanics.
A subset of neurons in the cochlear nucleus (CN) of the auditory brainstem has the ability to enhance the auditory nerve's temporal representation of stimulating sounds. These neurons reside in the ...ventral region of the CN (VCN) and are usually known as highly synchronized, or high-sync, neurons. Most published reports about the existence and properties of high-sync neurons are based on recordings performed on a VCN output tract--not the VCN itself--of cats. In other species, comprehensive studies detailing the properties of high-sync neurons, or even acknowledging their existence, are missing.Examination of the responses of a population of VCN neurons in chinchillas revealed that a subset of those neurons have temporal properties similar to high-sync neurons in the cat. Phase locking and entrainment--the ability of a neuron to fire action potentials at a certain stimulus phase and at almost every stimulus period, respectively--have similar maximum values in cats and chinchillas. Ranges of characteristic frequencies for high-sync neurons in chinchillas and cats extend up to 600 and 1000 Hz, respectively. Enhancement of temporal processing relative to auditory nerve fibers (ANFs), which has been shown previously in cats using tonal and white-noise stimuli, is also demonstrated here in the responses of VCN neurons to synthetic and spoken vowel sounds.Along with the large amount of phase locking displayed by some VCN neurons there occurs a deterioration in the spectral representation of the stimuli (tones or vowels). High-sync neurons exhibit a greater distortion in their responses to tones or vowels than do other types of VCN neurons and auditory nerve fibers.Standard deviations of first-spike latency measured in responses of high-sync neurons are lower than similar values measured in ANFs' responses. This might indicate a role of high-sync neurons in other tasks beyond sound localization.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The currently recognized mechanisms of the biology of cancer are not yet enough to explain the high incidence of the disease in industrialized countries. Survival and proliferation of cancer cells ...demand a well‐orchestrated combination of functional capabilities, or hallmarks, which requires complex signalling networks that often exceed the tumour boundaries. Based on latest research on environmental health and aiming to provide cancer with a coherent set of organizing principles, we propose an integrative model of carcinogenesis founded on tumour growth activation by the central nervous system as an adaptive, allostatic response to both environmental and emotional challenges. In this way, chronicity of physical as well as psychological stressors may be directly involved in cancer genesis and progression, after an early inflammatory stage. The model also contemplates accidental activation of the tumour growth programme following direct DNA damage, but as a rare event that does not account for most cancers in humans. Bodily and cellular mechanisms designed to facilitate tumorigenesis may include exacerbation of the sympathetic activity, overexpression of membrane ion channels, promotion of selected mutations and methylations, degradation of the mitochondria and reprogramming of adult stem cells.
Georg von Békésy observed that the onset times of responses to brief-duration stimuli vary as a function of distance from the stapes, with basal regions starting to move earlier than apical ones. He ...noticed that the speed of signal propagation along the cochlea is slow when compared with the speed of sound in water. Fast traveling waves have been recorded in the cochlea, but their existence is interpreted as the result of an experiment artifact. Accounts of the timing of vibration onsets at the base of the cochlea generally agree with Békésy's results. Some authors, however, have argued that the measured delays are too short for consistency with Békésy's theory. To investigate the speed of the traveling wave at the base of the cochlea, we analyzed basilar membrane (BM) responses to clicks recorded at several locations in the base of the chinchilla cochlea. The initial component of the BM response matches remarkably well the initial component of the stapes response, after a 4-μs delay of the latter. A similar conclusion is reached by analyzing onset times of time-domain gain functions, which correspond to BM click responses normalized by middle-ear input. Our results suggest that BM responses to clicks arise from a combination of fast and slow traveling waves.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Basilar-membrane responses to single tones were measured, using laser velocimetry, at a site of the chinchilla cochlea located 3.5 mm from its basal end. Responses to low-level (< 10-20 dB SPL) ...characteristic-frequency (CF) tones (9-10 kHz) grow linearly with stimulus intensity and exhibit gains of 66-76 dB relative to stapes motion. At higher levels, CF responses grow monotonically at compressive rates, with input-output slopes as low as 0.2 dB/dB in the intensity range 40-80 dB. Compressive growth, which is significantly correlated with response sensitivity, is evident even at stimulus levels higher than 100 dB. Responses become rapidly linear as stimulus frequency departs from CF. As a result, at stimulus levels > 80 dB the largest responses are elicited by tones with frequency about 0.4-0.5 octave below CF. For stimulus frequencies well above CF, responses stop decreasing with increasing frequency: A plateau is reached. The compressive growth of responses to tones with frequency near CF is accompanied by intensity-dependent phase shifts. Death abolishes all nonlinearities, reduces sensitivity at CF by as much as 60-81 dB, and causes a relative phase lead at CF.
Road traffic noise is a major public health issue, given the documented association with several diseases and the growing number of exposed persons all over the world. The effects widely investigated ...pertain to cardiovascular health, and to a lesser extent to respiratory and metabolic health. The epidemiological design of most studies has made it possible to ascertain long-term associations of urban noise with a number of cardiovascular, respiratory, and metabolic disorders and diseases; additionally, time series studies have reported short-term associations.
To review the various biological mechanisms that may account for all long-term as well as short-term associations between road traffic noise and cardiovascular, respiratory, and metabolic health. We also aimed to review the neuroendocrine processes triggered by noise as a stressor and the role of the central nervous system in noise-induced autonomic responses.
Review of the literature on road traffic noise, environmental noise in general, psychosomatics, and diseases of the cardiovascular, respiratory, and metabolic systems. The search was done using PubMed databases.
We present a comprehensive, integrative stress model with all known connections between the body systems, states, and processes at both the physiological and psychological levels, which allows to establish a variety of biological pathways linking environmental noise exposure with health outcomes.
The long- and short-term associations between road traffic noise and health outcomes found in latest noise research may be understood in the light of the integrative model proposed here.
Octopus cells in the ventral cochlear nucleus (VCN) have been difficult to study because of the very features that distinguish them from other VCN neurons. We performed in vivo recordings in cats on ...well-isolated units, some of which were intracellularly labeled and histologically reconstructed. We found that responses to low-frequency tones with frequencies < 1 kHz reveal higher levels of neural synchrony and entrainment to the stimulus than the auditory nerve. In responses to higher frequency tones, the neural discharges occur mostly near the stimulus onset. These neurons also respond in a unique way to 100 % amplitude-modulated (AM) tones with discharges exhibiting a bandpass tuning. Responses to frequency-modulated sounds (FM) are unusual: Octopus cells react more vigorously during the ascending than the descending parts of the FM stimulus. We examined responses of neurons in the ventral nucleus of the lateral lemniscus (VNLL) whose discharges to tones and AM sounds are similar to octopus cells. Repeated stimulation with short tone pips of VCN and VNLL onset neurons evokes trains of action potentials with gradual shifts toward later times in their first spike latency. This behavior parallels short-term post-synaptic depression observed by other authors in in vitro VCN recordings of octopus cells. VCN and VNLL onset units in cats respond to frozen noise stimuli with gaps as narrow as 1 ms with a robust discharge near the stimulus onset following the gap. This finding suggests that VCN and VNLL onset cells play a role in gap detection, which is of great importance to speech perception.
When stimulated by tones, the ear appears to emit tones of its own, stimulus-frequency otoacoustic emissions (SFOAEs). SFOAEs were measured in 17 chinchillas and their group delays were compared with ...a place map of basilar-membrane vibration group delays measured at the characteristic frequency. The map is based on Wiener-kernel analysis of responses to noise of auditory-nerve fibers corroborated by measurements of vibrations at several basilar-membrane sites. SFOAE group delays were similar to, or shorter than, basilar-membrane group delays for frequencies >4 kHz and <4 kHz, respectively. Such short delays contradict the generally accepted "theory of coherent reflection filtering" Zweig and Shera, J. Acoust. Soc. Am. 98, 2018-2047 (1995), which predicts that the group delays of SFOAEs evoked by low-level tones approximately equal twice the basilar-membrane group delays. The results for frequencies higher than 4 kHz are compatible with hypotheses of SFOAE propagation to the stapes via acoustic waves or fluid coupling, or via reverse basilar membrane traveling waves with speeds corresponding to the signal-front delays, rather than the group delays, of the forward waves. The results for frequencies lower than 4 kHz cannot be explained by hypotheses based on waves propagating to and from their characteristic places in the cochlea.
Key points
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Basilar membrane responses to tone and noise stimuli presented simultaneously were measured in chinchillas and a gerbil. Overall responses increase monotonically with stimulus level in ...a compressive manner.
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Response components elicited by the weaker stimulus, whether tone or noise, were reduced by the stronger one. That is, the suppressor stimulus could be either the tone or the noise. Suppression by noise stimuli occurred only when the tone frequency was in the non‐linear region of the basilar membrane recording site.
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Basilar membrane responses to click and noise stimuli were also recorded in some experiments. The click response component was suppressed by the noise.
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Mutual suppression, meaning the simultaneous suppression of the tone and noise response components, was also observed under certain stimulus conditions.
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Detection thresholds of the tone stimuli increased in a near‐linear fashion with noise level increments.
In the search for cochlear correlates of auditory masking by noise stimuli, we recorded basilar membrane (BM) vibrations evoked by either tone or click signals in the presence of varying levels of background noise. The BM vibrations were recorded from basal regions in healthy cochleae of anaesthetized chinchilla and gerbil. Non‐linear interactions that could underpin various aspects of psychophysical masking data, including both compression and suppression at the BM level, were observed. The suppression effects, whereby the amplitude of the responses to each stimulus component could be reduced, depended on the relative intensities of the noise and the tones or clicks. Only stimulus components whose frequencies fell inside the non‐linear region of the recording site, i.e. around its characteristic frequency (CF), were affected by presentation of the ‘suppressing’ stimulus (which could be either the tone or the noise). Mutual suppression, the simultaneous reduction of the responses to both tones and noise components, was observed under some conditions, but overall reductions of BM vibration were rarely observed. Moderate‐ to high‐intensity tones suppressed BM responses to low‐intensity Gaussian stimuli, including both broadband and narrowband noise. Suppression effects were larger for spectral components of the noise response that were closer to the CF. In this regime, the tone and noise stimuli became the suppressor and probe signals, respectively. This study provides the first detailed observations of cochlear mechanical correlates of the masking effects of noise. Mechanical detection thresholds for tone signals, which were arbitrarily defined using three criteria, are shown to increase in almost direct proportion to the noise level for low and moderately high noise levels, in a manner that resembles the findings of numerous psychophysical observations.
We review the mechanical origin of auditory-nerve excitation, focusing on comparisons of the magnitudes and phases of basilar-membrane (BM) vibrations and auditory-nerve fiber responses to tones at a ...basal site of the chinchilla cochlea with characteristic frequency ≈ 9 kHz located 3.5 mm from the oval window. At this location, characteristic frequency thresholds of fibers with high spontaneous activity correspond to magnitudes of BM displacement or velocity in the order of 1 nm or 50 μ m/s. Over a wide range of stimulus frequencies, neural thresholds are not determined solely by BM displacement but rather by a function of both displacement and velocity. Near-threshold, auditory-nerve responses to low-frequency tones are synchronous with peak BM velocity toward scala tympani but at 80-90 dB sound pressure level (in decibels relative to 20 microPascals) and at 100-110 dB sound pressure level responses undergo two large phase shifts approaching 180 degrees. These drastic phase changes have no counterparts in BM vibrations. Thus, although at threshold levels the encoding of BM vibrations into spike trains appears to involve only relatively minor signal transformations, the polarity of auditory-nerve responses does not conform with traditional views of how BM vibrations are transmitted to the inner hair cells. The response polarity at threshold levels, as well as the intensity-dependent phase changes, apparently reflect micromechanical interactions between the organ of Corti, the tectorial membrane and the subtectorial fluid, and/or electrical and synaptic processes at the inner hair cells.
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