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Morphological and behavioral mechanisms underlying sound localization in barn owls

机译:谷仓猫头鹰声音定位基础的形态和行为机制

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摘要

In this thesis, properties of the acoustic system and sound-localization behavior of barn owls were investigated. The influence of adaptation on sound localization behavior was examined in the first experiment. Payne (1971) was able to observe that barn owls wait for at least a second sound before they approach their prey. This situation was mimicked in a behavioral experiment to investigate how a preceding stimulus accuracy of the response to the second stimulus. It is known that the response of neurons to a second stimulus is decreased compared to the response to the first stimulus. This phenomenon is called response adaptation. This means that the detection threshold of the second stimulus may be elevated stimulus and, therefore, response-adaptation influences localization accuracy of the owl. Response adaptation was examined with a double stimulus paradigm. The owl had to locate a broadband noise token, which was preceded by another broadband noise token. I found out that the accuracy and precision with which the barn owls localized the sound source, decreased with double stimulation compared to the condition with only a single stimulus. By varying the interval between the end of the first and onset of the second stimulus I was able to show that the adaptive or masking effect of the first stimulus expires after a few hundred milliseconds. The results suggest that waiting for the second stimulus actually caused costs in terms of decreasing accuracy. In the second study, the head-turning behavior was used to compare responses to frequency-modulated and stimuli with stationary stimulus content. Barn owls detect time differences in the arrival of sound at both ears and can thus determine the azimuth of a sound source. When stimulated with narrow-band stationary stimuli, however, barn owls locate so-called phantom sources, i.e. they turn their head to a position that does not correspond to the actual sound source. The position of the phantom source can be predicted by the period of the center frequency and a known factor that converts the time differences in an angle. The percentage of phantom localization was determined as a function of stimulus bandwidth. Phantom sound sources are not localized at high stimulus bandwidths. Integration of frequency information in the auditory pathway of the barn owl leads to a reduction of phantom-source locations. Frequency-modulated tones offered the opportunity to present the same frequency content as with stationary noise, but within a certain time interval. This allows determination of the duration of the time window in which the frequency information is integrated. The behavioral data could be well explained with a model that simulates two important processes in the auditory pathway of the barn owl: 1) binaural interaction 2) integration of frequency information. The time constants of the time windows had a duration between 2 and 17 ms for both model steps and did not depend on the stimulus duration. In the third series of experiments I investigated whether the tympanic membrane of the barn owl functions as pressure receiver or as pressure-gradient receiver. In a pressure receiver, as it occurs in mammals, both middle ears are not acoustically coupled. This means that no sound is transmitted through an intracranial, interaural canal. Especially in small lizards, but also birds, however, there are cavities that couple both middle ears. Sound does not only reach the eardrum from outside, but also through the interaural canal. The incoming signals are phase shifted. The phase shift depends on stimulus location. In the case of lossless sound transmission through the interaural canal certain sound directions lead to complete extinction of the eardrum vibration. Consequently, the reduction of the eardrum vibration also depends on the degree of sound attenuation through the canal. To measure ear coupling the eardrum vibration was measured with a laser Doppler vibrometer. Eardrum vibration was measured as a function of stimulus frequency and azimuth. In addition, the actual attenuation of acoustic signals by the interaural canal was measured. The tympanic membrane was directional up to 3 kHz. That is, the eardrum vibration amplitude varied by more than 3 dB in 360° of stimulation angles. These data can be explained by attenuation of sound through the interaural canal. For frequencies higher than 3 kHz attenuation was too high to produce significant directionality.
机译:本文研究了acoustic的声学系统特性和声定位行为。在第一个实验中研究了适应对声音定位行为的影响。佩恩(Payne)(1971)能够观察到,bar接近它们的猎物之前至少要等待第二声。在行为实验中模仿了这种情况,以研究先前刺激对第二刺激的反应的准确性。众所周知,与对第一刺激的响应相比,神经元对第二刺激的响应降低了。这种现象称为响应适应。这意味着第二刺激的检测阈值可能会升高,因此响应适应会影响猫头鹰的定位精度。用双刺激范例检查了反应适应性。猫头鹰必须找到宽带噪声标记,然后再放置另一个宽带噪声标记。我发现,与只有一次刺激的情况相比,在两次刺激下,谷仓猫头鹰定位声源的准确性和精确度都会降低。通过改变第一个刺激的结束与第二个刺激的发作之间的间隔,我能够证明第一个刺激的适应性或掩盖效应在几百毫秒后就消失了。结果表明,等待第二轮刺激实际上会降低准确性。在第二项研究中,将转头行为用于比较对具有固定刺激内容的频率调制和刺激的响应。仓检测到声音到达两只耳朵时的时间差异,因此可以确定声源的方位角。但是,当受到窄带静止刺激刺激时,仓会定位所谓的幻像声源,即它们将头转向与实际声源不符的位置。幻象源的位置可以通过中心频率的周期和将角度的时间差转换为已知因子来预测。幻影定位的百分比被确定为刺激带宽的函数。幻像声源未定位在高刺激带宽下。频率信息在仓的听觉途径中的整合导致幻像源位置的减少。调频音调提供了在固定的时间间隔内呈现与固定噪声相同的频率含量的机会。这允许确定其中集成了频率信息的时间窗口的持续时间。可以用模拟仓鼠听觉途径中两个重要过程的模型来很好地解释行为数据:1)双耳相互作用2)频率信息的整合。对于两个模型步骤,时间窗口的时间常数的持续时间在2到17毫秒之间,并且与刺激的持续时间无关。在第三系列实验中,我研究了仓的鼓膜是作为压力接收器还是作为压力梯度接收器。在压力接收器中(就像在哺乳动物中一样),两只中耳在听觉上都不是耦合的。这意味着没有声音通过颅内,耳间管传输。尤其是在小蜥蜴中,在鸟类中也是如此,但是,有空腔将两只中耳耦合在一起。声音不仅从外部到达耳膜,而且还通过耳道。输入信号被相移。相移取决于激励位置。在通过耳间通道无损传输声音的情况下,某些声音方向会导致鼓膜振动完全消失。因此,鼓膜振动的减小还取决于通过耳道的声音衰减程度。为了测量耳耦合,用激光多普勒振动计测量鼓膜振动。测得的振膜振动是刺激频率和方位角的函数。另外,还测量了耳道对声信号的实际衰减。鼓膜的方向可达3 kHz。即,鼓膜振动振幅在360°的刺激角度变化超过3dB。这些数据可以通过穿过耳道的声音衰减来解释。对于高于3 kHz的频率,衰减太高而无法产生明显的方向性。

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    Kettler Lutz;

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  • 年度 2015
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  • 正文语种 eng
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