RESUMEN
Elephants have a unique auditory system that is larger than any other terrestrial mammal. To quantify the impact of larger middle ear (ME) structures, we measured 3D ossicular motion and ME sound transmission in cadaveric temporal bones from both African and Asian elephants in response to air-conducted (AC) tonal pressure stimuli presented in the ear canal (PEC). Results were compared to similar measurements in humans. Velocities of the umbo (VU) and stapes (VST) were measured using a 3D laser Doppler vibrometer in the 7-13,000 Hz frequency range, stapes velocity serving as a measure of energy entering the cochlea-a proxy for hearing sensitivity. Below the elephant ME resonance frequency of about 300 Hz, the magnitude of VU/PEC was an order of magnitude greater than in human, and the magnitude of VST/PEC was 5x greater. Phase of VST/PEC above ME resonance indicated that the group delay in elephant was approximately double that of human, which may be related to the unexpectedly high magnitudes at high frequencies. A boost in sound transmission across the incus long process and stapes near 9 kHz was also observed. We discuss factors that contribute to differences in sound transmission between these two large mammals.
Asunto(s)
Elefantes , Animales , Humanos , Oído Medio/fisiología , Sonido , Estribo/fisiología , Audición/fisiología , VibraciónRESUMEN
Elephants have a unique auditory system that is larger than any other terrestrial mammal. To quantify the impact of larger middle ear (ME) structures, we measured 3D ossicular motion and ME sound transmission in cadaveric temporal bones from both African and Asian elephants in response to air-conducted (AC) tonal pressure stimuli presented in the ear canal (P EC ). Results were compared to similar measurements in humans. Velocities of the umbo (V U ) and stapes (V ST ) were measured using a 3D laser Doppler vibrometer in the 7-13,000 Hz frequency range, stapes velocity serving as a measure of energy entering the cochlea-a proxy for hearing sensitivity. Below the elephant ME resonance frequency of about 300 Hz, the magnitude of V U /P EC was an order of magnitude greater than in human, and the magnitude of V ST /P EC was 5x greater. Phase of V ST /P EC above ME resonance indicated that the group delay in elephant was approximately double that of human, which may be related to the unexpectedly high magnitudes at high frequencies. A boost in sound transmission across the incus long process and stapes near 9 kHz was also observed. We discuss factors that contribute to differences in sound transmission between these two large mammals.
RESUMEN
Textbook descriptions of primary sensory cortex (PSC) revolve around single neurons' representation of low-dimensional sensory features, such as visual object orientation in primary visual cortex (V1), location of somatic touch in primary somatosensory cortex (S1), and sound frequency in primary auditory cortex (A1). Typically, studies of PSC measure neurons' responses along few (one or two) stimulus and/or behavioral dimensions. However, real-world stimuli usually vary along many feature dimensions and behavioral demands change constantly. In order to illuminate how A1 supports flexible perception in rich acoustic environments, we recorded from A1 neurons while rhesus macaques (one male, one female) performed a feature-selective attention task. We presented sounds that varied along spectral and temporal feature dimensions (carrier bandwidth and temporal envelope, respectively). Within a block, subjects attended to one feature of the sound in a selective change detection task. We found that single neurons tend to be high-dimensional, in that they exhibit substantial mixed selectivity for both sound features, as well as task context. We found no overall enhancement of single-neuron coding of the attended feature, as attention could either diminish or enhance this coding. However, a population-level analysis reveals that ensembles of neurons exhibit enhanced encoding of attended sound features, and this population code tracks subjects' performance. Importantly, surrogate neural populations with intact single-neuron tuning but shuffled higher-order correlations among neurons fail to yield attention- related effects observed in the intact data. These results suggest that an emergent population code not measurable at the single-neuron level might constitute the functional unit of sensory representation in PSC.SIGNIFICANCE STATEMENT The ability to adapt to a dynamic sensory environment promotes a range of important natural behaviors. We recorded from single neurons in monkey primary auditory cortex (A1), while subjects attended to either the spectral or temporal features of complex sounds. Surprisingly, we found no average increase in responsiveness to, or encoding of, the attended feature across single neurons. However, when we pooled the activity of the sampled neurons via targeted dimensionality reduction (TDR), we found enhanced population-level representation of the attended feature and suppression of the distractor feature. This dissociation of the effects of attention at the level of single neurons versus the population highlights the synergistic nature of cortical sound encoding and enriches our understanding of sensory cortical function.
Asunto(s)
Potenciales de Acción/fisiología , Atención/fisiología , Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Neuronas/fisiología , Estimulación Acústica , Animales , Femenino , Macaca mulatta , MasculinoRESUMEN
Selective attention is necessary to sift through, form a coherent percept of, and make behavioral decisions on the vast amount of information present in most sensory environments. How and where selective attention is employed in cortex and how this perceptual information then informs the relevant behavioral decisions is still not well understood. Studies probing selective attention and decision-making in visual cortex have been enlightening as to how sensory attention might work in that modality; whether or not similar mechanisms are employed in auditory attention is not yet clear. Therefore, we trained rhesus macaques on a feature-selective attention task, where they switched between reporting changes in temporal (amplitude modulation, AM) and spectral (carrier bandwidth) features of a broadband noise stimulus. We investigated how the encoding of these features by single neurons in primary (A1) and secondary (middle lateral belt, ML) auditory cortex was affected by the different attention conditions. We found that neurons in A1 and ML showed mixed selectivity to the sound and task features. We found no difference in AM encoding between the attention conditions. We found that choice-related activity in both A1 and ML neurons shifts between attentional conditions. This finding suggests that choice-related activity in auditory cortex does not simply reflect motor preparation or action and supports the relationship between reported choice-related activity and the decision and perceptual process.NEW & NOTEWORTHY We recorded from primary and secondary auditory cortex while monkeys performed a nonspatial feature attention task. Both areas exhibited rate-based choice-related activity. The manifestation of choice-related activity was attention dependent, suggesting that choice-related activity in auditory cortex does not simply reflect arousal or motor influences but relates to the specific perceptual choice.
Asunto(s)
Atención/fisiología , Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Conducta de Elección/fisiología , Desempeño Psicomotor/fisiología , Animales , Conducta Animal/fisiología , Electrocorticografía , Femenino , Macaca mulatta , MasculinoRESUMEN
In macaques, the middle lateral auditory cortex (ML) is a belt region adjacent to the primary auditory cortex (A1) and believed to be at a hierarchically higher level. Although ML single-unit responses have been studied for several auditory stimuli, the ability of ML cells to encode amplitude modulation (AM)-an ability that has been widely studied in A1-has not yet been characterized. Here, we compared the responses of A1 and ML neurons to amplitude-modulated (AM) noise in awake macaques. Although several of the basic properties of A1 and ML responses to AM noise were similar, we found several key differences. ML neurons were less likely to phase lock, did not phase lock as strongly, and were more likely to respond in a nonsynchronized fashion than A1 cells, consistent with a temporal-to-rate transformation as information ascends the auditory hierarchy. ML neurons tended to have lower temporally (phase-locking) based best modulation frequencies than A1 neurons. Neurons that decreased their firing rate in response to AM noise relative to their firing rate in response to unmodulated noise became more common at the level of ML than they were in A1. In both A1 and ML, we found a prevalent class of neurons that usually have enhanced rate responses relative to responses to the unmodulated noise at lower modulation frequencies and suppressed rate responses relative to responses to the unmodulated noise at middle modulation frequencies.NEW & NOTEWORTHY ML neurons synchronized less than A1 neurons, consistent with a hierarchical temporal-to-rate transformation. Both A1 and ML had a class of modulation transfer functions previously unreported in the cortex with a low-modulation-frequency (MF) peak, a middle-MF trough, and responses similar to unmodulated noise responses at high MFs. The results support a hierarchical shift toward a two-pool opponent code, where subtraction of neural activity between two populations of oppositely tuned neurons encodes AM.
Asunto(s)
Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Neuronas/fisiología , Estimulación Acústica , Potenciales de Acción , Animales , Femenino , Macaca mulatta , MasculinoRESUMEN
An anatomically based three-dimensional finite-element human middle-ear (ME) model is used to test the sensitivity of ME sound transmission to tympanic-membrane (TM) material properties. The baseline properties produce responses comparable to published measurements of ear-canal input impedance and power reflectance, stapes velocity normalized by ear-canal pressure (PEC), and middle-ear pressure gain (MEG), i.e., cochlear-vestibule pressure (PV) normalized by PEC. The mass, Young's modulus (ETM), and shear modulus (GTM) of the TM are varied, independently and in combination, over a wide range of values, with soft and bony TM-annulus boundary conditions. MEG is recomputed and plotted for each case, along with summaries of the magnitude and group-delay deviations from the baseline over low (below 0.75 kHz), mid (0.75-5 kHz), and high (above 5 kHz) frequencies. The MEG magnitude varies inversely with increasing TM mass at high frequencies. Increasing ETM boosts high frequencies and attenuates low and mid frequencies, especially with a bony TM annulus and when GTM varies in proportion to ETM, as for an isotropic material. Increasing GTM on its own attenuates low and mid frequencies and boosts high frequencies. The sensitivity of MEG to TM material properties has implications for model development and the interpretation of experimental observations.
Asunto(s)
Oído Medio/anatomía & histología , Oído Medio/fisiología , Audición , Modelos Anatómicos , Modelos Teóricos , Membrana Timpánica/anatomía & histología , Membrana Timpánica/fisiología , Oído Medio/diagnóstico por imagen , Módulo de Elasticidad , Análisis de Elementos Finitos , Humanos , Movimiento (Física) , Presión , Reproducibilidad de los Resultados , Sonido , Membrana Timpánica/diagnóstico por imagen , Vibración , Microtomografía por Rayos XRESUMEN
Sensory environments often contain an overwhelming amount of information, with both relevant and irrelevant information competing for neural resources. Feature attention mediates this competition by selecting the sensory features needed to form a coherent percept. How attention affects the activity of populations of neurons to support this process is poorly understood because population coding is typically studied through simulations in which one sensory feature is encoded without competition. Therefore, to study the effects of feature attention on population-based neural coding, investigations must be extended to include stimuli with both relevant and irrelevant features. We measured noise correlations (rnoise) within small neural populations in primary auditory cortex while rhesus macaques performed a novel feature-selective attention task. We found that the effect of feature-selective attention on rnoise depended not only on the population tuning to the attended feature, but also on the tuning to the distractor feature. To attempt to explain how these observed effects might support enhanced perceptual performance, we propose an extension of a simple and influential model in which shifts in rnoise can simultaneously enhance the representation of the attended feature while suppressing the distractor. These findings present a novel mechanism by which attention modulates neural populations to support sensory processing in cluttered environments.SIGNIFICANCE STATEMENT Although feature-selective attention constitutes one of the building blocks of listening in natural environments, its neural bases remain obscure. To address this, we developed a novel auditory feature-selective attention task and measured noise correlations (rnoise) in rhesus macaque A1 during task performance. Unlike previous studies showing that the effect of attention on rnoise depends on population tuning to the attended feature, we show that the effect of attention depends on the tuning to the distractor feature as well. We suggest that these effects represent an efficient process by which sensory cortex simultaneously enhances relevant information and suppresses irrelevant information.
Asunto(s)
Adaptación Fisiológica , Atención , Corteza Auditiva/fisiología , Percepción Auditiva , Ruido , Animales , Corteza Auditiva/citología , Potenciales Evocados Auditivos , Femenino , Macaca mulatta , Masculino , Neuronas/fisiologíaRESUMEN
The ability to segregate simultaneous sound sources based on their spatial locations is an important aspect of auditory scene analysis. While the role of sound azimuth in segregation is well studied, the contribution of sound elevation remains unknown. Although previous studies in humans suggest that elevation cues alone are not sufficient to segregate simultaneous broadband sources, the current study demonstrates they can suffice. Listeners segregating a temporally modulated noise target from a simultaneous unmodulated noise distracter differing in elevation fall into two statistically distinct groups: one that identifies target direction accurately across a wide range of modulation frequencies (MF) and one that cannot identify target direction accurately and, on average, reports the opposite direction of the target for low MF. A non-spiking model of inferior colliculus neurons that process single-source elevation cues suggests that the performance of both listener groups at the population level can be accounted for by the balance of excitatory and inhibitory inputs in the model. These results establish the potential for broadband elevation cues to contribute to the computations underlying sound source segregation and suggest a potential mechanism underlying this contribution.
Asunto(s)
Percepción Auditiva/fisiología , Localización de Sonidos/fisiología , Adulto , Señales (Psicología) , Femenino , Humanos , Colículos Inferiores/fisiología , Masculino , Persona de Mediana Edad , Modelos Biológicos , Ruido , Enmascaramiento Perceptual/fisiología , Psicofísica , Percepción del Habla/fisiología , Adulto JovenRESUMEN
We recorded from middle lateral belt (ML) and primary (A1) auditory cortical neurons while animals discriminated amplitude-modulated (AM) sounds and also while they sat passively. Engagement in AM discrimination improved ML and A1 neurons' ability to discriminate AM with both firing rate and phase-locking; however, task engagement affected neural AM discrimination differently in the two fields. The results suggest that these two areas utilize different AM coding schemes: a "single mode" in A1 that relies on increased activity for AM relative to unmodulated sounds and a "dual-polar mode" in ML that uses both increases and decreases in neural activity to encode modulation. In the dual-polar ML code, nonsynchronized responses might play a special role. The results are consistent with findings in the primary and secondary somatosensory cortices during discrimination of vibrotactile modulation frequency, implicating a common scheme in the hierarchical processing of temporal information among different modalities. The time course of activity differences between behaving and passive conditions was also distinct in A1 and ML and may have implications for auditory attention. At modulation depths ≥ 16% (approximately behavioral threshold), A1 neurons' improvement in distinguishing AM from unmodulated noise is relatively constant or improves slightly with increasing modulation depth. In ML, improvement during engagement is most pronounced near threshold and disappears at highly suprathreshold depths. This ML effect is evident later in the stimulus, and mainly in nonsynchronized responses. This suggests that attention-related increases in activity are stronger or longer-lasting for more difficult stimuli in ML.
Asunto(s)
Corteza Auditiva/fisiología , Discriminación en Psicología/fisiología , Percepción Sonora/fisiología , Neuronas/fisiología , Estimulación Acústica , Potenciales de Acción , Animales , Femenino , Macaca mulatta , Masculino , Microelectrodos , Pruebas Neuropsicológicas , Curva ROC , Procesamiento de Señales Asistido por Computador , Factores de TiempoRESUMEN
We recorded from middle-lateral (ML) and primary (A1) auditory cortex while macaques discriminated amplitude-modulated (AM) noise from unmodulated noise. Compared with A1, ML had a higher proportion of neurons that encoded increasing AM depth by decreasing their firing rates ("decreasing" neurons), particularly with responses that were not synchronized to the modulation. Choice probability (CP) analysis revealed that A1 and ML activity were different during the first half of the test stimulus. In A1, significant CP began before the test stimulus, remained relatively constant (or increased slightly) during the stimulus, and increased greatly within 200 ms of lever release. Neurons in ML behaved similarly, except that significant CP disappeared during the first half of the stimulus and reappeared during the second half and prerelease periods. CP differences between A1 and ML depend on neural response type. In ML (but not A1), when activity was lower during the first half of the stimulus in nonsynchronized, decreasing neurons, the monkey was more likely to report AM. Neurons that both increased firing rate with increasing modulation depth ("increasing" neurons) and synchronized their responses to AM had similar choice-related activity dynamics in ML and A1. These results suggest that, when ascending the auditory system, there is a transformation in coding AM from primarily synchronized increasing responses in A1 to nonsynchronized and dual (increasing/decreasing) coding in ML. This sensory transformation is accompanied by changes in the timing of activity related to choice, suggesting functional differences between A1 and ML related to attention and/or behavior.
Asunto(s)
Corteza Auditiva/citología , Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Conducta de Elección/fisiología , Neuronas/fisiología , Sonido , Estimulación Acústica , Potenciales de Acción/fisiología , Animales , Discriminación en Psicología , Femenino , Lateralidad Funcional , Macaca mulatta , Masculino , Curva ROC , Tiempo de Reacción , Estadísticas no ParamétricasRESUMEN
The effect of attention on single neuron responses in the auditory system is unresolved. We found that when monkeys discriminated temporally amplitude modulated (AM) from unmodulated sounds, primary auditory cortical (A1) neurons better discriminated those sounds than when the monkeys were not discriminating them. This was observed for both average firing rate and vector strength (VS), a measure of how well neurons temporally follow the stimulus' temporal modulation. When data were separated by nonsynchronized and synchronized responses, the firing rate of nonsynchronized responses best distinguished AM- noise from unmodulated noise, followed by VS for synchronized responses, with firing rate for synchronized neurons providing the poorest AM discrimination. Firing rate-based AM discrimination for synchronized neurons, however, improved most with task engagement, showing that the least sensitive code in the passive condition improves the most with task engagement. Rate coding improved due to larger increases in absolute firing rate at higher modulation depths than for lower depths and unmodulated sounds. Relative to spontaneous activity (which increased with engagement), the response to unmodulated sounds decreased substantially. The temporal coding improvement--responses more precisely temporally following a stimulus when animals were required to attend to it--expands the framework of possible mechanisms of attention to include increasing temporal precision of stimulus following. These findings provide a crucial step to understanding the coding of temporal modulation and support a model in which rate and temporal coding work in parallel, permitting a multiplexed code for temporal modulation, and for a complementary representation of rate and temporal coding.
Asunto(s)
Potenciales de Acción/fisiología , Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Discriminación en Psicología/fisiología , Neuronas/fisiología , Percepción del Tiempo/fisiología , Animales , Femenino , Macaca mulatta , MasculinoRESUMEN
Recent evidence is reshaping the view of primary auditory cortex (A1) from a unisensory area to one more involved in dynamically integrating multisensory- and task-related information. We found A1 single- (SU) and multiple-unit (MU) activity correlated with macaques' choices in an amplitude modulation (AM) discrimination task. Animals were trained to discriminate AM noise from unmodulated noise by releasing a lever for AM noise and holding down the lever for unmodulated noise. Activity for identical stimuli was compared between trials where the animals reported AM and trials where they did not. We found 47.4% of MUs and 22.8% of SUs significantly increased firing shortly before the animal's behavioral response to report AM when compared to the equivalent time period on trials where AM was not reported. Activity was also linked to lever release in a different task context, suggesting A1 modulation by somatosensory, or efference copy, input. When spikes were counted only during the stimulus, 19.6% of MUs and 13.8% of SUs increased firing rate when animals reported AM compared to when they did not, suggesting an attentional effect, or that A1 activity can be used by higher decision areas, or that such areas provide feedback to A1. Activity associated with AM reporting was correlated with a unit's AM sensitivity, suggesting AM sensitive neurons' involvement in task performance. A1 neurons' phase locking to AM correlated more weakly (compared to firing rate) with the animals' report of AM, suggesting a preferential role for rate-codes in A1 for this AM discrimination task.
Asunto(s)
Estimulación Acústica , Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Juicio/fisiología , Estimulación Acústica/métodos , Potenciales de Acción , Animales , Mapeo Encefálico/métodos , Femenino , Macaca mulatta , Masculino , Distribución AleatoriaRESUMEN
Amplitude modulation (AM) is a common feature of natural sounds, and its detection is biologically important. Even though most sounds are not fully modulated, the majority of physiological studies have focused on fully modulated (100% modulation depth) sounds. We presented AM noise at a range of modulation depths to awake macaque monkeys while recording from neurons in primary auditory cortex (A1). The ability of neurons to detect partial AM with rate and temporal codes was assessed with signal detection methods. On average, single-cell synchrony was as or more sensitive than spike count in modulation detection. Cells are less sensitive to modulation depth if tested away from their best modulation frequency, particularly for temporal measures. Mean neural modulation detection thresholds in A1 are not as sensitive as behavioral thresholds, but with phase locking the most sensitive neurons are more sensitive, suggesting that for temporal measures the lower-envelope principle cannot account for thresholds. Three methods of preanalysis pooling of spike trains (multiunit, similar to convergence from a cortical column; within cell, similar to convergence of cells with matched response properties; across cell, similar to indiscriminate convergence of cells) all result in an increase in neural sensitivity to modulation depth for both temporal and rate codes. For the across-cell method, pooling of a few dozen cells can result in detection thresholds that approximate those of the behaving animal. With synchrony measures, indiscriminate pooling results in sensitive detection of modulation frequencies between 20 and 60 Hz, suggesting that differences in AM response phase are minor in A1.
Asunto(s)
Corteza Auditiva/fisiología , Percepción Auditiva/fisiología , Modelos Neurológicos , Estimulación Acústica , Animales , Femenino , Macaca mulatta , Masculino , Neuronas/fisiologíaRESUMEN
Previous observations show that humans outperform non-human primates on some temporally-based auditory discrimination tasks, suggesting there are species differences in the proficiency of auditory temporal processing among primates. To further resolve these differences we compared the abilities of rhesus macaques and humans to detect sine-amplitude modulation (AM) of a broad-band noise carrier as a function of both AM frequency (2.5 Hz-2 kHz) and signal duration (50-800 ms), under similar testing conditions. Using a go/no-go AM detection task, we found that macaques were less sensitive than humans at the lower frequencies and shorter durations tested but were as, or slightly more, sensitive at higher frequencies and longer durations. Humans had broader AM tuning functions, with lower frequency regions of peak sensitivity (10-60 Hz) than macaques (30-120 Hz). These results support the notion that there are species differences in temporal processing among primates, and underscore the importance of stimulus duration when making cross-species comparisons for temporally-based tasks.
Asunto(s)
Vías Auditivas/fisiología , Conducta Animal , Discriminación en Psicología , Discriminación de la Altura Tonal , Estimulación Acústica , Adulto , Análisis de Varianza , Animales , Audiometría , Umbral Auditivo , Femenino , Humanos , Macaca mulatta , Masculino , Persona de Mediana Edad , Patrones de Reconocimiento Fisiológico , Psicoacústica , Especificidad de la Especie , Factores de TiempoRESUMEN
Conflicting results have led to different views about how temporal modulation is encoded in primary auditory cortex (A1). Some studies find a substantial population of neurons that change firing rate without synchronizing to temporal modulation, whereas other studies fail to see these nonsynchronized neurons. As a result, the role and scope of synchronized temporal and nonsynchronized rate codes in AM processing in A1 remains unresolved. We recorded A1 neurons' responses in awake macaques to sinusoidal AM noise. We find most (37-78%) neurons synchronize to at least one modulation frequency (MF) without exhibiting nonsynchronized responses. However, we find both exclusively nonsynchronized neurons (7-29%) and "mixed-mode" neurons (13-40%) that synchronize to at least one MF and fire nonsynchronously to at least one other. We introduce new measures for modulation encoding and temporal synchrony that can improve the analysis of how neurons encode temporal modulation. These include comparing AM responses to the responses to unmodulated sounds, and a vector strength measure that is suitable for single-trial analysis. Our data support a transformation from a temporally based population code of AM to a rate-based code as information ascends the auditory pathway. The number of mixed-mode neurons found in A1 indicates this transformation is not yet complete, and A1 neurons may carry multiplexed temporal and rate codes.
Asunto(s)
Relojes Biológicos/fisiología , Red Nerviosa/fisiología , Percepción de la Altura Tonal/fisiología , Animales , Corteza Auditiva , Femenino , Macaca mulatta , MasculinoRESUMEN
Electromagnetic floating-mass transducers for implantable middle-ear hearing devices (IMEHDs) afford the advantages of a simple surgical implantation procedure and easy attachment to the ossicles. However, their shortcomings include susceptibility to interference from environmental electromagnetic fields, relatively high current consumption, and a limited ability to output high-frequency vibrations. To address these limitations, a piezoelectric floating-mass transducer (PFMT) has recently been developed. This paper presents the results of a comparative study of these two types of vibration transducer developed for IMEHDs. The differential electromagnetic floating-mass transducer (DFMT) and the PFMT were implanted in two different sets of three cadaveric human temporal bones. The resulting stapes displacements were measured and compared on the basis of the ASTM standard for describing the output characteristics of IMEHDs. The experimental results show that the PFMT can produce significantly higher equivalent sound pressure levels above 3 kHz, due to the flat response of the PFMT, than can the DFMT. Thus, it is expected that the PFMT can be utilized to compensate for high-frequency sensorineural hearing loss.
Asunto(s)
Fenómenos Electromagnéticos , Audífonos , Hueso Temporal/fisiología , Transductores de Presión , Estimulación Acústica , Cadáver , Diseño de Equipo , Humanos , Presión , Procesamiento de Señales Asistido por Computador , Estribo/fisiología , Hueso Temporal/cirugía , VibraciónRESUMEN
The focus of most research on auditory cortical neurons has concerned the effects of rather simple stimuli, such as pure tones or broad-band noise, or the modulation of a single acoustic parameter. Extending these findings to feature coding in more complex stimuli such as natural sounds may be difficult, however. Generalizing results from the simple to more complex case may be complicated by non-linear interactions occurring between multiple, simultaneously varying acoustic parameters in complex sounds. To examine this issue in the frequency domain, we performed a parametric study of the effects of two global features, spectral pattern (here ripple frequency) and bandwidth, on primary auditory (A1) neurons in awake macaques. Most neurons were tuned for one or both variables and most also displayed an interaction between bandwidth and pattern implying that their effects were conditional or interdependent. A spectral linear filter model was able to qualitatively reproduce the basic effects and interactions, indicating that a simple neural mechanism may be able to account for these interdependencies. Our results suggest that the behavior of most A1 neurons is likely to depend on multiple parameters, and so most are unlikely to respond independently or invariantly to specific acoustic features.
RESUMEN
Middle-ear circuit model parameters are selected to produce overall magnitude and phase agreement with pressure to stapes velocity transfer function measurements made on 16 human temporal bones, up to approximately 12 kHz. The circuit model, which was previously used for the cat, represents the tympanic membrane (TM) as a distributed parameter acoustic transmission line, and ossicular chain and cochlea as a network of lumped circuit elements. For some ears the TM transmission line primarily affects the magnitude of the response, while for others it primarily affects the phase. Model responses also compare favorably with velocity ratio data between the umbo and stapes footplate as well as between the umbo and incus, and exhibit similar characteristics to three previous input impedance measurements, including two from living ears. Similarities are also shown between the model magnitude and adjusted pressure to stapes velocity measurements from living ears, suggesting that the model may suitably approximate the behavior of living ears. In addition to fitting individual measurements, a set of parameters is selected to produce agreement with the mean of the 16 measurements up to 10 kHz, to allow the main features of the ensemble to be reproduced from a single parameter set.
Asunto(s)
Oído Medio/fisiología , Percepción Auditiva/fisiología , Humanos , Estribo/fisiología , Hueso Temporal/anatomía & histología , Membrana Timpánica/fisiologíaRESUMEN
OBJECTIVE: To investigate the significance of tympanic membrane collagen fiber layers in high frequency sound transmission. STUDY DESIGN: Human cadaver temporal bone study. METHODS: Laser Doppler vibrometry was used to measure stapes footplate movement in response to acoustic stimulation. The tympanic membrane was altered by creating a series of slits and applying paper patches to isolate the effects of specifically oriented collagen fibers. Three groups of membrane alterations were evaluated: 1) circumferentially oriented slits involving each quadrant to primarily disrupt radial fibers, made sequentially within superior-anterior, inferior-anterior, inferior-posterior, and superior-posterior quadrants; 2) the same slits made in the reverse order; and 3) radially oriented slits from the umbo to the annulus to primarily disrupt circumferential fibers. For each group, measurements of the middle-ear cavity pressure, ear canal pressure, and stapes velocity were made each time the tympanic membrane was altered. RESULTS: Regardless of the order in which the circumferentially oriented slits were made, there was a consistent decrease in stapes velocity above 4 kHz for the third and fourth cuts compared to the control. The mean decrease in the range of 4 to 12.5 kHz was 11 dB for the third patched slit and 14 dB for the fourth patched slit (P < .01). Radially oriented slits appear to produce smaller effects. CONCLUSIONS: Radial collagen fibers in the tympanic membrane play an important role in the conduction of sound above 4 kHz.
Asunto(s)
Colágeno , Sonido , Membrana Timpánica/fisiología , Membrana Timpánica/ultraestructura , Fenómenos Biofísicos , Biofisica , Cadáver , HumanosRESUMEN
Experiments were conducted to evaluate a silicon accelerometer as an implantable sound sensor for implantable hearing aids. The main motivation of this study is to find an alternative sound sensor that is implantable inside the body, yet does not suffer from the signal attenuation from the body. The merit of the accelerometer sensor as a sound sensor will be that it will utilize the natural mechanical conduction in the middle ear as a source of the vibration. With this kind of implantable sound sensor, a totally implantable hearing aid is feasible. A piezoresistive silicon accelerometer that is completely encapsulated with a thin silicon film and long flexible flex-circuit electrical cables were used for this study. The sensor is attached on the middle ear ossicles and measures the vibration transmitted from the tympanic membrane due to the sound in the ear canal. In this study, the sensor is fully characterized on a human cadaveric temporal bone preparation.