菊头蝠捕食时对障碍物的回避能力

食虫蝙蝠依靠发射超声讯号并收听其回声来感知周围环境。Griffin(1944)将此过程命名为“回声定位”(Echolocation)。测定这种“回声定位”能力,通常都采用障碍物回避试验法(Griffin,1944,1974;Suga,1969;Jen,1980),即让蝙蝠在静止不动的细丝阵列之间飞行,记录碰撞和回避障碍物的次数,以判断蝙蝠“回声定位”能力的有无和高低。Jell与 McCarty(1978)发展了这一方法,让蝙蝠在障碍物间捕食飞行,并比较观察,发现蝙蝠能有效地回避障碍目标。了解自然界中以捕食昆虫为生的蝙蝠,在其捕食过程中,如何回避障碍物这个问题,对研究食虫蝙蝠的“回声定位”机制是有意义的。本文以鲁氏菊头蝠(Rhinolophus rouxi)为对象进行了实验研究。     

鲁氏菊头蝠回避障碍时抗干扰能力的研究

实验在30只鲁氏菊头蝠(Rhinolcphus rouxi)中进行。使用障碍回避测定法,观察了白噪声和不同频率超声干扰对回避障碍能力的影响。发现宽频带白噪声不能干扰动物回避障碍的成功率。分别使用该动物的特征频率85kHz正弦波,以及相近的65kHz、75kHz、95kHz和105kHz的超声干扰信号,都不能影响障碍回避成功率。实验结果提示,食虫蝙蝠的生物声纳系统具有很强的抵抗白噪声和超声信号干扰的能力,以保证它们从周围环境中获取有意义的生物信息。     

东方蝙蝠在交通噪声环境中的回声定位行为

环境噪声会降低动物声信号可探测性,是动物声信号进化的压力。为了避免人工噪声干扰,多数脊椎动物调整其叫声的频谱-时间结构,如采用延长叫声持续时间、提高频率和增加强度等策略。本研究选择回声定位声波频率范围与交通噪声频率范围相互重叠的东方蝙蝠(Vespertilio sinensis)为研究对象,在自然条件下开展实验,研究交通噪声对蝙蝠回声定位声波的影响。本实验选择交通噪声强度存在差异的两个样点,分别录制东方蝙蝠的回声定位声波,分析其结构参数。在交通噪声强度较高的样点,东方蝙蝠回声定位声波的持续时间、斜率均未显著改变,但起始频率、主频、终止频率及带宽均显著提高。结果表明,东方蝙蝠的回声定位行为在交通噪声干扰下具有明显的可塑性。     

我国学者发现蝙蝠具有暗视觉并非“瞎子”

<正>一直以来,人们认为蝙蝠是夜行性动物,食虫蝙蝠主要用回声定位来确定方位、捕捉昆虫,其眼睛退化,蝙蝠被认为是"瞎子"。然而,中国科学院昆明动物研究所张亚平院士及其博士生沈永义发现,旧大陆果蝠并没有回声定位能力,它们主要依赖视觉和嗅觉来寻找食物,眼睛很发达。     

马铁菊头蝠对同种咀嚼声的反应研究

咀嚼是哺乳动物食物吸收的重要组成部分,对动物的生存和繁殖极其重要。动物咀嚼时会发出低频低强度的咀嚼声,研究表明人类的咀嚼声可以增强自身或他人的食欲和愉悦度。蝙蝠作为哺乳动物中的第二大类群,其咀嚼声的特征和功能仍不清楚。本研究以吉林省集安市治安村的马铁菊头蝠Rhinolophus ferrumequinum为研究对象,通过回放蝙蝠咀嚼声和空白对照实验,同步录制蝙蝠的进食行为和回声定位声波,试图阐明蝙蝠咀嚼声对其进食行为和回声定位声波的影响。结果表明:进食次数在2种回放条件下的差异无统计学意义,但捕食尝试在2种回放条件下的差异有统计学意义。因此,咀嚼声能够显著地提高蝙蝠的进食欲望。此外,在蝙蝠咀嚼声的刺激下,其回声定位声波的频率增加,持续时间延长,脉冲速率减慢。这可能是因为蝙蝠在咀嚼声的影响下改变发声动机,从而改变了其回声定位声波的频谱时间结构。本研究第一次报道了蝙蝠咀嚼声对其进食行为的影响,为进一步阐明蝙蝠咀嚼声的功能和进化具有重要意义。     

六种共栖蝙蝠的回声定位信号及翼型特征的比较

本研究于2006 年5 ～ 8 月在桂林市七星公园七星岩洞进行,比较分析了共栖2 科(蹄蝠科和蝙蝠科)6 种共75 只蝙蝠的回声定位信号和翼型特征。普氏蹄蝠的回声定位叫声为短而多谐波的CF/ FM 型,主频率为61.2±0.8 kHz, 具有高翼载、低翼展比和中等翼尖指数; 大蹄蝠的回声定位叫声为单CF/ FM 型,主频率为68. 6 ±0.7 kHz,具有高翼载、低翼展比和中等翼尖指数;中蹄蝠的回声定位叫声为单CF / FM 型,主频率为85.2 ±0.5 kHz,具有中等翼载、低翼展比和中等翼尖指数;高颅鼠耳蝠的回声定位叫声为长带宽的FM 型,主频率为50.7 ±3.8 kHz,具有低翼载、中等翼展比和低翼尖指数;大足鼠耳蝠回声定位叫声为FM 型,主频率为39.9 ±3.2 kHz,具有中等翼载、低翼展比和高翼尖指数;绒山蝠回声定位叫声为短而多谐波的FM 型,主频率为49.0± 0. 4 kHz,具有高翼载、中等翼展比和低翼尖指数。经单因素方差分析表明,6 种蝙蝠之间绝大部分的形态和声音参数差异显著(One-way ANOVA,P < 0. 05)。以上结果说明,6 种同地共栖蝙蝠种属特异的回声定位叫声
和形态结构体现出了相互之间的生态位分离,从而降低了种间竞争压力,使得6 种蝙蝠能够同地共存。     

降雨噪声对菲菊头蝠出飞行为的影响

降雨噪声属于常见的自然噪声,由雨滴撞击物体表面产生。目前,有关降雨噪声对动物的潜在影响被普遍忽视。回声定位蝙蝠主要利用声信号在黑暗环境导航空间、探测猎物及社群交流,是开展降雨噪声影响研究的理想类群。本研究选择菲菊头蝠 (Rhinolophus pusillus)作为研究对象,检验降雨噪声是否影响蝙蝠出飞行为。我们在集群栖息地外,播放强降雨噪声、空白对照和种内回声定位声波,开展野外回放实验。利用单因素方差分析及其事后检验,评价菲菊头蝠对不同回放刺激的反应差异。研究发现,相比空白对照,强降雨噪声导致菲菊头蝠的通勤数量百分比平均降低2.82倍,回声定位脉冲数量平均减少4.86倍,集群出飞时间延长3.75 min。相比空白对照,同种回声定位声波对菲菊头蝠出飞行为的影响并不显著。研究结果证实强降雨噪声抑制菲菊头蝠的出飞行为。本研究表明,降雨引起 的噪声干扰可能是导致蝙蝠躲避降雨的重要因素,为野生蝙蝠物种保育与管理提供启示。     

环境噪声对蝙蝠回声定位叫声及飞行活动的影响

环境噪声影响动物的活动及其叫声特性,已成为动物面对的一种重要选择压力。为应对噪声的干扰,多数动物类群会远离噪声区域和改变其叫声的频谱时间结构,如延长叫声持续时间、提高叫声频率等,但有些动物的活动和叫声频谱时间结构并不受环境噪声的影响。本研究在自然条件下,研究不同环境噪声强度对蝙蝠活动和回声定位声波的影响。选取噪声强度有差异的12个样点,分别录制各样点大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的回声定位声波,分析其持续时间、起始频率、峰频、终止频率和带宽,统计蝙蝠通过次数。回归分析结果显示：环境噪声强度与大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的活动无显著相关性P > 0.05）,与回声定位声波的脉冲持续时间、起始频率、峰频、终止频率及带宽均不显著相关（P > 0.05）。暗示低频低强度（< 20 kHz, < 67.5 dB）的环境噪声可能对高频回声定位蝙蝠的叫声及活动没有显著影响。     

菲菊头蝠回声定位声波频率的性二态有利于性别识别

许多动物的叫声频率呈现性二态现象。蝙蝠夜间活动,主要利用声音信号导航空间、追踪猎物、传递交流信息。本研究选择成体菲菊头蝠作为研究对象,检验回声定位声波频率性二态是否有利于性别识别。研究发现,菲菊头蝠回声定位声波频率参数具有显著性别差异。播放白噪音、雄性回声定位声波及雌性回声定位声波期间,实验个体的反应叫声数量依次递减。播放白噪音、雌性回声定位声波及雄性回声定位声波后,实验个体的反应叫声数量依次递增。白噪音诱导反应叫声强度高于回声定位声波诱导反应叫声强度。研究结果表明,菲菊头蝠回声定位声波的频率参数编码发声者性别信息,有利于种群内部的性别识别。本研究暗示,回声定位声波可能在蝙蝠配偶选择中扮演一定作用。     

蝙蝠与超声波、回声定位(1)

回声定位是高度演化、极为复杂的过程,使蝙蝠可利用大多数动物不能利用的生态位——漆黑的洞穴和黑夜的天空。对蝙蝠的回声定位研究已有近80年的历史,科学家已经从生物声学层面基本了解和认识了蝙蝠回声定位的特征、机制、生物学意义等,关于分子和神经生物学方面的机制也得到深入研究。重点介绍蝙蝠回声定位的研究历史,以及蝙蝠的超声波和回声定位在生物学和声学层面的基础知识。     

Rapid jamming avoidance in biosonar

The sonar systems of bats and dolphins are in many ways superior to man-made sonar and radar systems, and considerable effort has been devoted to understanding the signal-processing strategies underlying these capabilities. A major feature determining the efficiency of sonar systems is the sensitivity to noise and jamming signals. Previous studies indicated that echolocating bats may adjust their signal structure to avoid jamming ('jamming avoidance response'; JAR). However, these studies relied on behavioural correlations and not controlled experiments. Here, we provide the first experimental evidence for JAR in bats. We presented bats (Tadarida brasiliensis) with 'playback stimuli' consisting of recorded echolocation calls at one of six frequencies. The bats exhibited a JAR by shifting their call frequency away from the presented playback frequency. When the approaching bats were challenged by an abrupt change in the playback stimulus, they responded by shifting their call frequencies upwards, away from the playback. Interestingly, even bats initially calling below the playback's frequency shifted their frequencies upwards, 'jumping' over the playback frequency. These spectral shifts in the bats' calls occurred often within less than 200 ms, in the first echolocation call emitted after the stimulus switch-suggesting that rapid jamming avoidance is important for the bat.     

普氏蹄蝠下丘谐波主频神经元的时程选择性

为探讨下丘(Inferior colliculus,IC)回声定位信号主频范围内的神经元的时程选择性,在自由声场刺激条件下,我们在4 只普氏蹄蝠的IC 采用不同时程的声刺激,研究了神经元的时程选择性。通过在体细胞外记录,共获得56 个声敏感下丘神经元,其记录深度、最佳频率和最小阈值的范围分别为1547 - 3967 (2878. 9 ±629.1)μm,20 -68 (49.0 ± 11. 1)kHz 和36.5 -95. 5 (59. 8 ±13. 0)dB SPL。根据所记录到的下丘神经元对不同时程的声刺激的反应,即对不同时程的选择性(Duration selectivity),将其分为6 种类型:短通型(Short-pass,SP,n = 11/56)、带通型(Band-pass,BP,n = 1/56)、长通型(Long-pass,LP,n = 5 /56)、反带通型(Band-reject,BR,n = 3 /56)、多峰型(Multi-peak,MP,n =6 /56)和全通型(All-pass,AP,n =30 /56)或非时程选择型(Nonduration-selective,NDS)。通过比较普氏蹄蝠下丘谐波主频内和主频外神经元的时程选择性,我们发现处于回声定位信号主频范围内神经元(n =32)比主频外神经元(n = 24)具有更短的最佳时程和更高的时程选择性。结果提示,在普氏蹄蝠回声定位过程中谐波主频内神经元较谐波主频外神经元发挥了更为重要的作用。     

Support for the allotonic frequency hypothesis in an insectivorous bat community

The allotonic frequency hypothesis proposes that certain insectivorous bat species can prey upon moths that can hear bat echolocation calls by using echolocation frequencies outside the sensitivity range of moth ears. The hypothesis predicts that the peak frequencies of bat echolocation calls are correlated with the incidence of moths in the diets of these bats. The aim of this study was to test this prediction on a bat community dominated by bats using low duty cycle echolocation calls, i.e. aerial foraging, insectivorous species using frequency modulated calls. The community consisted of nine species, two molossids, Sauromys petrophillus and Tadarida aegyptiaca, five vespertilionids, Eptesicus capensis, Eptesicus hottentotus, Miniopteris schreibersii, Myotis tricolor, and Myotis lesueuri, one rhinolophid, Rhinolophus clivosus, and one nycterid, Nycteris thebaica. The insect fauna in the habitat used by the bat community was suited to the testing of the allotonic frequency hypothesis because more than 90% of the moths comprising the insect fauna were tympanate. These included Pyralidae (3.8%), Geometridae (44.9%), Notodontidae (3.8%), Arctiidae (4.6%), Lymantriidae (0.8%) and Noctuidae (32.4%). As predicted, peak echolocation frequency was correlated with the incidence of moths in the diets of these nine species (r=0.98, df=7, P<0.01). Furthermore, multivariate analysis revealed that echolocation frequency (t=9.91, n=129, P<0.001) was a better predictor of diet than forearm length (t=5.51, n=129, P<0.001) or wing area (t=-3.41, n=129, P<0.001). This suggests that the selection pressure exerted by moth hearing might have acted directly on call frequency and secondarily on body size and wing morphology, as part of the same adaptive complex. It is unlikely that dietary differences were due to temporal and spatial differences in the availability of prey because the pattern of differences in skull morphology of the nine species supported our dietary analyses. The skull morphology of a bat represents a historical record of the kind of diet it has become adapted to over its evolutionary history. These results suggest that prey defences may mediate other factors structuring bat communities, e.g. competition. Competition may be reduced for those species of bats that can circumvent prey defences.     

Prey-Correlated Spectral Changes in Echolocation Sounds of the Indian False Vampire Megaderma lyra

False Vampires ( Megaderma lyra ) are gleaning bats which emit brief (1 ms) and faint echolocation signals consisting of four harmonics of a shallow frequency downward modulated fundamental (27–19 kHz). The complete signal spans a frequency range from 100 to 19 kHz. In sound recordings from three experimental animals we show that Megaderma lyra shifts the dominant frequency in the echolocation signals in relation to the type of prey offered and to flight style. During roaming flights the mean peak frequency was 63.2 ± 9 kHz (third harmonic). In prey catching flights, peak frequencies were shifted into the fourth harmonic. In flights towards a dish of crawling mealworms, mean peak frequency was raised to 91.2 ± 3.3 kHz. When the bats flew towards living mice the dominant frequency was further increased to 99.8 ± 5.2 kHz, and the second and third harmonic were at least 10 dB fainter or no longer recordable. The additional frequency shift when flying towards mice was not only a consequence of the dominance of the fourth harmonic but also of an additional rise of the fundamental harmonic by nearly 2 kHz. These prey-correlated frequency shifts in echolocation calls showed little variation between the three experimental animals and were reproducible over time. They occurred at or even before take-off of the bats. This is the first report of target-correlated transient adaptations in echolocation calls of any bat species.     

在不同状态和生境复杂度中大蹄蝠回声定位叫声的可塑性

蝙蝠回声定位声波的可塑性对其适应不同状态、生境以及捕食和社会交流具有重要的作用。为进一步研究大蹄蝠的回声定位声波在不同状态和生境下的可塑性,通过室内行为实验,对大蹄蝠在4 种不同状态(室内飞行、静息、布袋内和手持)和4 种不同生境复杂度(室外、室内0 棵树、室内1 棵树、室内5 棵树)条件下飞行的回声定位声波特征进行研究。结果表明:大蹄蝠的回声定位声波为CF - FM 型,通常连续发出2 - 4 个脉冲组成一个脉冲组。对比大蹄蝠在4 种不同状态下的回声定位叫声发现,主频按静息、布袋内、手持、飞行的顺序依次降低,后端FM 频宽则按手持、布袋内、飞行和静息的顺序依次变短;而脉冲间隔和脉冲时程则均按静 息、飞行、布袋内、手持的顺序依次增加。对比大蹄蝠在4 种不同生境复杂度中飞行的回声定位叫声发现,主频按室外、室内0 棵树、室内1 棵树、室内5 棵树依次提高,而脉冲时程及脉冲间隔则依次缩短;室外放飞条件下的后端FM 频宽比室内飞行的短。研究结果说明,大蹄蝠在不同状态、不同生境复杂度条件下的回声定位叫声具有明显的可塑性和生境适应性。     

‘Compromise’ in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus)

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the ‘compromise’ in echolocation calls might be used to ensure effective mutual communication among bats.     

皮氏菊头蝠回声定位声波与年龄的关系

皮氏菊头蝠 (Rhinolophuspearsoni)雌性成体 5只和幼体 2只采自贵州省贞丰县珉谷镇。采用超声波探测仪 (D980 ,ULTRASOUNDDETECTOR)接收皮氏菊头蝠的回声定位声波 ,转换到原频率的 1 / 1 0后导入计算机 ,然后用专业声谱分析软件 (Batsound 3 1 0 )进行分析。成蝠在飞行和悬挂状态下的声波结构相似 ,只是声波各项参数值略有不同 :它们发射FM CF FM型声波 ,具有 2～ 3个谐波 ,主频率在飞行时为 5 6 80± 0 6 2kHz ,悬挂时为 5 8 0 5± 0 2 4kHz ;声脉冲时间和间隔在飞行时分别为 3 4 6 2± 5 2 9ms和 86 5 0± 1 9 72ms ,悬挂时分别为 4 1 0 8± 5 87ms和 1 1 7 2 9± 6 6 4 4ms ;能率环飞行时为 ( 4 4 0 6± 1 2 5 8) % ,悬挂时为 ( 4 6 0 0±2 4 2 5 ) %。幼蝠声波为CF FM型 ,谐波数为 5～ 8个 ,主频率明显低于成体 ,FM带宽窄于成体 ,声脉冲时间和间隔短于成体 ,能率环低于成体。皮氏菊头蝠回声定位声波与年龄有关 ,这可能因成体的声波主要是探测食物和周围环境的详细信息 ,而幼体主要是与母蝠进行交流。     

Effect of auditory cortical ablation on functioning of the doppler echolocation system in horseshoe bats

The effect of unilateral and bilateral partial and total ablation of the auditory cortex on compensation of Doppler frequency shifts in echosignals was studied in greater horseshoe bats moving in space. The ability of the bats to compensate Doppler surges in the echolocation signal was found to be worsened even after partial ablation of the auditory cortex. Total bilateral ablation led to more marked and irreversible changes in the functioning of the Doppler echolocation system. In this case the degree of compensation in the decorticated animals was only 33% of normal; the return of the frequency of the constant part of the signal to the resting level after movement of the animal ceased was delayed. After total ablation of the auditory cortical projection area definite retrograde degeneration of cells of the medial geniculate body was observed. It is concluded from the results that the auditory cortex in bats plays a direct part in echolocation spatial analysis.A. A. Ukhtomskii Physiological Research Institute, A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 43–50, January–February, 1982.