Nolen T G, Hoy R R
J Comp Physiol A. 1986 Oct;159(4):441-56. doi: 10.1007/BF00604164.
The effects of two-tone stimuli on the high frequency bat-avoidance steering behavior of flying crickets (Teleogryllus oceanicus) were studied during tethered flight. Similarly, the effects of two-tone stimuli on the ultrasound sensitive auditory interneuron, Int-1, which elicits this behavior, were studied using intracellular staining and recording techniques. When a low frequency tone (3-8 kHz) was presented simultaneously with an aversive high frequency tone (in a two-tone stimulus paradigm), the high frequency avoidance steering behavior was suppressed. Suppression was optimal when the low frequency tone was between 4 and 5 kHz and about 10-15 dB louder than the high frequency tone (Figs. 2, 3). Best suppression occurred when the low frequency tone-pulse just preceded or overlapped the high frequency tone-pulse, indicating that the suppressive effects of 5 kHz could last for up to 70 ms (Fig. 4). The threshold for avoidance of the bat-like stimulus was elevated when model bat biosonar (30 kHz) was presented while the animal was performing positive phonotaxis toward 5 kHz model calling song, but only if the calling song intensity was relatively high (greater than 70-80 dB SPL) (Fig. 1). However, avoidance steering could always be elicited as long as the calling song was not more than 10 dB louder than the ultrasound (Fig. 1). This suppressive effect did not require performance of positive phonotaxis to the calling song (Fig. 2) and was probably due to the persistence of the suppressive effects of the 5 kHz model calling song (Fig. 4). The requirement for relatively high intensities of calling song suggest that the suppression of bat-avoidance by the calling song is not likely to be of great significance in nature. The high frequency harmonics of the male cricket's natural calling song overlap the lower frequency range used by insectivorous bats (10-20 kHz) and are loud enough to elicit avoidance behavior in a flying female as she closely approaches a singing male (Fig. 5). The high frequency 'harmonics' of a model calling song were aversive even if presented with a normally attractive temporal pattern (pulse repetition rate of 16 pps) (Fig. 6A). When the 5 kHz 'fundamental' was added to one of the high frequency 'harmonics', in a two-tone stimulus paradigm, this complex model calling song was attractive; the high frequency 'harmonic' no longer elicited the avoidance behavior (Fig. 6) and the animals steered toward the model CS. Thus, addition of 5 kHz to a high frequency harmonic of the calling song 'masked' the aversive nature of this stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)
在系留飞行过程中，研究了双音刺激对飞行蟋蟀（Teleogryllus oceanicus）高频避蝠转向行为的影响。同样，使用细胞内染色和记录技术，研究了双音刺激对引发这种行为的超声敏感听觉中间神经元Int-1的影响。当低频音（3-8千赫）与厌恶高频音同时呈现时（在双音刺激范式中），高频回避转向行为受到抑制。当低频音在4至5千赫之间且比高频音大约高10-15分贝时，抑制效果最佳（图2、3）。当低频音脉冲刚好先于或与高频音脉冲重叠时，出现最佳抑制效果，这表明5千赫的抑制作用可持续长达70毫秒（图4）。当动物朝着5千赫的模型求偶鸣声进行正趋声行为时，呈现模型蝙蝠生物声纳（30千赫）会提高对蝙蝠样刺激的回避阈值，但前提是求偶鸣声强度相对较高（大于70-80分贝声压级）（图1）。然而，只要求偶鸣声不比超声响超过10分贝，就总能引发回避转向（图1）。这种抑制作用并不需要对求偶鸣声进行正趋声行为（图2），可能是由于5千赫模型求偶鸣声的抑制作用持续存在（图4）。对相对高强度求偶鸣声的要求表明，求偶鸣声对避蝠行为的抑制在自然界中可能意义不大。雄性蟋蟀自然求偶鸣声的高频谐波与食虫蝙蝠使用的较低频率范围（10-20千赫）重叠，并且强度足以在飞行中的雌性接近歌唱的雄性时引发回避行为（图5）。即使以通常有吸引力的时间模式（脉冲重复率为16次/秒）呈现，模型求偶鸣声的高频“谐波”也具有厌恶作用（图6A）。在双音刺激范式中，当将5千赫的“基频”添加到高频“谐波”之一时，这种复杂的模型求偶鸣声具有吸引力；高频“谐波”不再引发回避行为（图6），动物转向模型CS。因此，在求偶鸣声的高频谐波中添加5千赫“掩盖”了这种刺激的厌恶性质。（摘要截断于400字）
