普氏蹄蝠、鲁氏菊头蝠、长翼蝠的生态、形态、及耳蜗结构和听觉功能的研究
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摘要
蝙蝠在中国的种类丰富，共有7科31属117种，其特殊的回声定位功能引起了许多学者的兴趣。目前我国对蝙蝠的研究主要集中在以下方面：蝙蝠的分布和分类；蝙蝠的生态学，包括了其食性、栖息地、亲缘关系、昼夜活动节律、冬眠等；蝙蝠的化学通讯；蝙蝠的生殖和发育；蝙蝠的染色体和核型研究；蝙蝠携带病毒的研究；蝙蝠的生化研究；蝙蝠的听觉和回声定位研究，包括了回声定位蝙蝠的捕食策略、声通讯、听觉系统特化的研究；回声定位和蝙蝠行为状态、生态位、体形、性别的关系；以及听觉的神经生理学和神经组织化学方面的研究。
本文对陕西境内柞水地区一个洞穴中的蝙蝠进行为期一年的研究，内容包括了普氏蹄蝠、鲁氏菊头蝠、长翼蝠的生态、形态、及耳蜗结构和听觉功能。提出了研究中存在的问题，并对研究方向进行了展望。研究结果如下：
1．在陕西境内柞水地区进行蝙蝠生态的区系调查，从2005年到2006年的蝙蝠非冬眠期，在不同月份白天进入洞穴中调查。调查的内容包括洞穴中蝙蝠的种类，以及在不同月份内蝙蝠的种群数量和栖息分布的位置变化情况。观察发现蝙蝠对洞穴生境具有选择性，蝙蝠喜欢在农田—草灌植被状态良好生境的洞穴中栖息，偏爱人为干扰较少的洞穴，喜欢较深、黑暗、潮湿、温暖的洞穴。普氏蹄蝠、鲁氏菊头蝠、长翼蝠在春末，夏季，初秋都存在着集群分布现象，是与集群捕食和集群繁殖行为相关。而蝙蝠分布的层次性使得不同种类的蝙蝠在生存空间上存在了显著的异质性，引起了生态位上的分化，生态位分化是多种蝙蝠同地共栖的关键。蝙蝠在春末到夏季雄性多于雌性，出现性比失调推测可能是因为：一是蝙蝠迁徙路过此地，二是雌性寻找合适地点越冬，而雄性闻讯前来交配。菊头蝠是这个蝙蝠种群中的优势种群，证明了栖息地有良好的阔叶植被，是健康的生态环境。而对于蝙蝠数量减少这一现象，很大程度是由于人为的干扰和对栖息地的影响所致。以上结果为中国蝙蝠分布区系研究提供了直接证据和资料，对蝙蝠种群的保护，探讨蝙蝠对环境的适应和进化提供了依据。
2．对柞水同一个洞穴中的蝙蝠进行种类鉴定，外形特征描述和相关数据测量。经过中国科学院动物研究所鉴定为以下种类：普氏蹄蝠指名亚种(Hipposideros pratti pratti Thomas)、鲁氏菊头蝠中国亚种(Rhinolophus rouxi sinicus Andersen)、长翼蝠(折翼蝠)中国亚种(Miniopterus schreibersi chinensisThomas)。数据测量结果表明蹄蝠、菊头蝠、长翼蝠在形态学数据和耳相关数据上的差异，决定了这几种共栖同一山洞的蝙蝠的回声定位信号、和听觉相关的行为模式、捕食策略、生态位的差别。也就是说，蝙蝠的回声定位行为以及相关的形态特征与其捕食策略及捕食生境有着密切的联系。从蹄蝠的外部特征来看，体重最重、前臂长最长、体形最大、有较长的翼、耳长和耳宽最大，决定了它们适于远距离飞行，是快速飞行者，但飞行的灵活性不高。蹄蝠捕食环境较为复杂，捕食体形较大，飞行较快的昆虫，来满足其生活的能量需求。菊头蝠具有较小的体重、头体长、前臂长、可判断其飞行速度较慢，但是飞行的灵敏性很高，在相对简单的环境中，比如树木枝叶或其周围的狭窄空间内捕食较小的翼拍动昆虫。长翼蝠具有较小的体重、头体长、前臂长，可判断其飞行速度较慢，但是飞行的灵敏性很高，适合在复杂的环境中捕食，比如较为狭窄，较为繁乱的空间中进行拾遗式捕食。以上结果为中国蝙蝠形态学数据库提供了补充和直接证据，再一次证实了蝙蝠的回声定位行为以及相关的形态特征与其捕食策略及捕食生境有着密切联系这一理论基础。
3．对柞水同一个洞穴中的蝙蝠进行听觉功能测定和耳蜗基底膜的毛细胞超微结构观察。通过使用Tucker Davis Technology(TDT)BioSig SystemⅢ测定，发现蝙蝠听觉诱发电位测定在12-48kH是其敏感区，蝙蝠的听觉敏感区与小鼠的听觉敏感区12—32kHz基本一致，而棕色田鼠的听觉敏感区是在6—8kHz。蝙蝠耳蜗毛细胞超微结构观察结果是：蝙蝠耳蜗结构与人类耳蜗极为相似。蝙蝠耳蜗内毛细胞基本与其他哺乳类动物一样呈单排，静纤毛较短，基底部的静纤毛束根数较少，表皮板相互分离；顶部的纤毛根数较多，表皮板相互连接。而外毛细胞静纤毛与其他哺乳类动物具有明显的差异，表现为外毛细胞静纤毛特别的短，均可见到静纤毛的断缺、融合现象，这些现象与小鼠老年性聋的表现相似。蝙蝠耳蜗外毛细胞体，主要特征有两点：其一是细胞体明显短，三排Deiters细胞的指突明显与顶表面相连，而且毛细胞体基本被Deiters细胞的杯状膜包裹；其二是外毛细胞体在基底部呈烧瓶状，并不是一般哺乳类动物的试管样形态。以上研究结果表明蝙蝠和大多数其他小型哺乳动物在一定范围内虽有着相似的听力敏度图，却还是有不同之处的，回声定位蝙蝠经常拥有对它们来说更为有用频率的敏锐听力。蝙蝠的耳蜗结构在部分相似于其它哺乳类动物的同时，也有部分特化现象的存在，这些特征都被认为是处理超高频声波的形态学适应。以上研究结果不仅能对回声定位的动物行为学和神经生物学提供帮助，而且能为人类的听觉机制和耳聋防治提供借鉴。
There are 117 species of chiropters belonging to 31 genera, 7 families inChina. Their particular echolocation attracts many researchers' focus and interesting. Atpresent, the studies of bats in China mostly focus on these aspects: bats' distribution andclassification; the ecology of bats, including their prey. selection, habitat, theirrelationship, their activity rhythm in day and night, their hibernation and so on; thechemical communication of bats; their reproduction and growth; chromosome andkaryotype; bats as reservior host of virus; physilology and biochemistry; hearing andecholocation, which contain echolocating bats' foraging strategy, audio communication,and the specific of their auditory system; The relationship of echolocation and differentactivity states, ecology niche, body size, sexual differences are also investigated.Meanwhile, the review shows aspects of neurophysiology and neurobiochemistry ofauditory system.
The bats in one cave in QinLing Mountain in ZhaShui area of ShannXi wereinvestigated during one year in present research, which contains the echology,morphology, structure of cochlea and hearing function of Hipposideros pratti prattiTomas, Rhinolophus rouxi sinicus Andersen and Miniopterus schreibersi chinensisThomas. Suggestions for future research are also made. The results are summarized asfollows:
1.From 2005 to 2006,bats in one cave in ZhaShui area of ShannXi wereinvestigated.We entered into the cave in the daytime in different months of one year,which exclude period of bats' hibernation. The content consist of the species of bats, thequantity of population and the diversification of distribution in different months.According to the records and investigating, bats have the ability to select more suitableliving space of caves for them. They prefer to inhabit caves, which are surrounded withgood vegetation of farmland-grassland. They also like to select the habitats of caves which are more deep, dark, moisture, warmer, low elevation and lesser jamming. At theend of the spring, in summer and at the beginging of the autumn, there is a phenomenonof assembly distribution of Hipposideros pratti pratti Tomas, Rhinolophus rouxi sinicusAndersen and Miniopterus schreibersi chinensis Thomas, which relate with groupforaging and group reproduction. The hiberarchy of distribution of bats bring onremarkable differences of different species in their living spaces. This differentiation ofecology niche is a key for different species living in the same cave. During the end ofspring and whole summer, the number of male is much more than female's. Thismaladjustment of sex ratio may result from migration of bats and their mating system.Rhinolophus rouxi sinicus Andersen is the dominant species, which demonstrate that thehabitat holding good broad-leaved vegetation is a healthy environment. Moreover, togreat extent, jamming and influence of habitat cause the decrease of bat population. Theoutcome above provide the straight evidence and data of distribution of bats in China,which offer some theory basis for bat' protection and discussion of how bats adaptenvironment and evolve.
2.Bats living in the same cave in ZhaShui region was identified, described andsome related data also are measured. There are three species of bats, Hipposideros prattipratti Thomas, Rhinolophus rouxi sinicus Andersen and Miniopterus schreibersichinensis Thomas, which are identified by Zoological Institute, CAS.The findings showthat there are some distinctions in morphologic datum and ear related datum amongthese bats inhabiting the same cave, which determine the differences of echolocaioncalls, activity patterns related to hearing, foraging strategy and ecology niche. That is tosay, bats' echolocation activities and their morphologic characters have close connectionwith their foraging strategy and foraging habitat. From the morphologic characters ofHipposideros pratti pratti Tomas, we can conclude that they own the heaviest weight,the longest forearm length, the biggest figure, the longest wing, ear length and ear width.These factors decide that they are fit for long-distance flying, flying quickly andunskillfully. Hipposideros pratti pratti Tomas often forage in complicated region, attacktargets with the bigger shape and the ability of flying quickly, which can meet theirenergy needs. Rhinolophus rouxi sinicus Andersen with the lightweight, short totallength and forarm length, which we can conclude that their flying speed is slow, buthave the skillful flying ability. They can detect the little wing-beating insects incomparative simple condition, such as in dense foliages and narrow habitat around. Miniopterus schreibersi chinensis Thomas has the similarities with Rhinolophus rouxisinicus Andersen, but they ofen prey on in complicated room by gleaning, such as innarrow space. These results provide supplement and proof of morphologic database ofbats in China, and approve the theory again, which bats' echolocation activities andtheir morphologic characters have close connection with their foraging strategy andforaging habitat.
3. Bats living in the same cave in ZhaShui region were studied by mensuratingtheir hearing function and observing their aultra-structure of hair cells of cochlea. Byusing Tucker-Davis Technology(TDT)Biosig SystemⅢ, we gain the result that auditorybrainstem responses(ABR) of bats,which sensitive region is 12-48kHz.This outcome isbasically consistent with the ICR mouse's sensitive region that is 12-32kHz,and thatMicrotus mandarinus's sensitive region is 6-8kHz. The bats' cochlear hair cells wereobserved by scanning electron microscope, and the structure of bats'cochlear is similarto human being's. The inner hair cells (IHCs) of bats resemble any other mammals,which own only one row, and their stereocilia are short. At the basal of basilarmembrane, the stereocilia bundles of IHCs are thin and cuticular plate (CP) separatewith each other. On the contrary, at the apex of basilar membrane, the stereociliabundles are bushy and cuticular plate join together. At the same time, there aresignificant differences between the outer hair cells (OHCs) stereocilia of bats and othermammals's. The stereocilia of OHCs are extremely short. We can observe thephenomena of rupture and inosculation of stereocilia, which is alike senile deafness ofICR mouse. The OHCs bodies of bats have two leading characteristics. Firstly, theOHCs bodies are obviously short. The Deiters cell phalanges are of exaggerated, andthe cup formation of Deiters cell body enwrap the bottom of the OHCs.Secondly, TheOHCs body possess the flask-shaped in the basal turn of the cochlea, which don't likeother mammals'tube-shaped OHCs body. All of the results above indicate that bats andmostly any other little mammals have the similar audiogram in definite range, but alsohave some difference. Echolocating bats always possess of acuity hearing, whichinclude more available frequency. The structure of bats'cochlea is partly resembled withother mammals; meanwhile, there are some specilized aspects, which are considered asmorphologic adaptation to dealing with ultra-high frequency sound wave. All thefindings not only supply helps for the researchesof ethology and neurobiology ofecholocating animals, but also provide suggestions of hearing mechanism and deafness prevention in human beings.
本文对陕西境内柞水地区一个洞穴中的蝙蝠进行为期一年的研究，内容包括了普氏蹄蝠、鲁氏菊头蝠、长翼蝠的生态、形态、及耳蜗结构和听觉功能。提出了研究中存在的问题，并对研究方向进行了展望。研究结果如下：
1．在陕西境内柞水地区进行蝙蝠生态的区系调查，从2005年到2006年的蝙蝠非冬眠期，在不同月份白天进入洞穴中调查。调查的内容包括洞穴中蝙蝠的种类，以及在不同月份内蝙蝠的种群数量和栖息分布的位置变化情况。观察发现蝙蝠对洞穴生境具有选择性，蝙蝠喜欢在农田—草灌植被状态良好生境的洞穴中栖息，偏爱人为干扰较少的洞穴，喜欢较深、黑暗、潮湿、温暖的洞穴。普氏蹄蝠、鲁氏菊头蝠、长翼蝠在春末，夏季，初秋都存在着集群分布现象，是与集群捕食和集群繁殖行为相关。而蝙蝠分布的层次性使得不同种类的蝙蝠在生存空间上存在了显著的异质性，引起了生态位上的分化，生态位分化是多种蝙蝠同地共栖的关键。蝙蝠在春末到夏季雄性多于雌性，出现性比失调推测可能是因为：一是蝙蝠迁徙路过此地，二是雌性寻找合适地点越冬，而雄性闻讯前来交配。菊头蝠是这个蝙蝠种群中的优势种群，证明了栖息地有良好的阔叶植被，是健康的生态环境。而对于蝙蝠数量减少这一现象，很大程度是由于人为的干扰和对栖息地的影响所致。以上结果为中国蝙蝠分布区系研究提供了直接证据和资料，对蝙蝠种群的保护，探讨蝙蝠对环境的适应和进化提供了依据。
2．对柞水同一个洞穴中的蝙蝠进行种类鉴定，外形特征描述和相关数据测量。经过中国科学院动物研究所鉴定为以下种类：普氏蹄蝠指名亚种(Hipposideros pratti pratti Thomas)、鲁氏菊头蝠中国亚种(Rhinolophus rouxi sinicus Andersen)、长翼蝠(折翼蝠)中国亚种(Miniopterus schreibersi chinensisThomas)。数据测量结果表明蹄蝠、菊头蝠、长翼蝠在形态学数据和耳相关数据上的差异，决定了这几种共栖同一山洞的蝙蝠的回声定位信号、和听觉相关的行为模式、捕食策略、生态位的差别。也就是说，蝙蝠的回声定位行为以及相关的形态特征与其捕食策略及捕食生境有着密切的联系。从蹄蝠的外部特征来看，体重最重、前臂长最长、体形最大、有较长的翼、耳长和耳宽最大，决定了它们适于远距离飞行，是快速飞行者，但飞行的灵活性不高。蹄蝠捕食环境较为复杂，捕食体形较大，飞行较快的昆虫，来满足其生活的能量需求。菊头蝠具有较小的体重、头体长、前臂长、可判断其飞行速度较慢，但是飞行的灵敏性很高，在相对简单的环境中，比如树木枝叶或其周围的狭窄空间内捕食较小的翼拍动昆虫。长翼蝠具有较小的体重、头体长、前臂长，可判断其飞行速度较慢，但是飞行的灵敏性很高，适合在复杂的环境中捕食，比如较为狭窄，较为繁乱的空间中进行拾遗式捕食。以上结果为中国蝙蝠形态学数据库提供了补充和直接证据，再一次证实了蝙蝠的回声定位行为以及相关的形态特征与其捕食策略及捕食生境有着密切联系这一理论基础。
3．对柞水同一个洞穴中的蝙蝠进行听觉功能测定和耳蜗基底膜的毛细胞超微结构观察。通过使用Tucker Davis Technology(TDT)BioSig SystemⅢ测定，发现蝙蝠听觉诱发电位测定在12-48kH是其敏感区，蝙蝠的听觉敏感区与小鼠的听觉敏感区12—32kHz基本一致，而棕色田鼠的听觉敏感区是在6—8kHz。蝙蝠耳蜗毛细胞超微结构观察结果是：蝙蝠耳蜗结构与人类耳蜗极为相似。蝙蝠耳蜗内毛细胞基本与其他哺乳类动物一样呈单排，静纤毛较短，基底部的静纤毛束根数较少，表皮板相互分离；顶部的纤毛根数较多，表皮板相互连接。而外毛细胞静纤毛与其他哺乳类动物具有明显的差异，表现为外毛细胞静纤毛特别的短，均可见到静纤毛的断缺、融合现象，这些现象与小鼠老年性聋的表现相似。蝙蝠耳蜗外毛细胞体，主要特征有两点：其一是细胞体明显短，三排Deiters细胞的指突明显与顶表面相连，而且毛细胞体基本被Deiters细胞的杯状膜包裹；其二是外毛细胞体在基底部呈烧瓶状，并不是一般哺乳类动物的试管样形态。以上研究结果表明蝙蝠和大多数其他小型哺乳动物在一定范围内虽有着相似的听力敏度图，却还是有不同之处的，回声定位蝙蝠经常拥有对它们来说更为有用频率的敏锐听力。蝙蝠的耳蜗结构在部分相似于其它哺乳类动物的同时，也有部分特化现象的存在，这些特征都被认为是处理超高频声波的形态学适应。以上研究结果不仅能对回声定位的动物行为学和神经生物学提供帮助，而且能为人类的听觉机制和耳聋防治提供借鉴。
There are 117 species of chiropters belonging to 31 genera, 7 families inChina. Their particular echolocation attracts many researchers' focus and interesting. Atpresent, the studies of bats in China mostly focus on these aspects: bats' distribution andclassification; the ecology of bats, including their prey. selection, habitat, theirrelationship, their activity rhythm in day and night, their hibernation and so on; thechemical communication of bats; their reproduction and growth; chromosome andkaryotype; bats as reservior host of virus; physilology and biochemistry; hearing andecholocation, which contain echolocating bats' foraging strategy, audio communication,and the specific of their auditory system; The relationship of echolocation and differentactivity states, ecology niche, body size, sexual differences are also investigated.Meanwhile, the review shows aspects of neurophysiology and neurobiochemistry ofauditory system.
The bats in one cave in QinLing Mountain in ZhaShui area of ShannXi wereinvestigated during one year in present research, which contains the echology,morphology, structure of cochlea and hearing function of Hipposideros pratti prattiTomas, Rhinolophus rouxi sinicus Andersen and Miniopterus schreibersi chinensisThomas. Suggestions for future research are also made. The results are summarized asfollows:
1.From 2005 to 2006,bats in one cave in ZhaShui area of ShannXi wereinvestigated.We entered into the cave in the daytime in different months of one year,which exclude period of bats' hibernation. The content consist of the species of bats, thequantity of population and the diversification of distribution in different months.According to the records and investigating, bats have the ability to select more suitableliving space of caves for them. They prefer to inhabit caves, which are surrounded withgood vegetation of farmland-grassland. They also like to select the habitats of caves which are more deep, dark, moisture, warmer, low elevation and lesser jamming. At theend of the spring, in summer and at the beginging of the autumn, there is a phenomenonof assembly distribution of Hipposideros pratti pratti Tomas, Rhinolophus rouxi sinicusAndersen and Miniopterus schreibersi chinensis Thomas, which relate with groupforaging and group reproduction. The hiberarchy of distribution of bats bring onremarkable differences of different species in their living spaces. This differentiation ofecology niche is a key for different species living in the same cave. During the end ofspring and whole summer, the number of male is much more than female's. Thismaladjustment of sex ratio may result from migration of bats and their mating system.Rhinolophus rouxi sinicus Andersen is the dominant species, which demonstrate that thehabitat holding good broad-leaved vegetation is a healthy environment. Moreover, togreat extent, jamming and influence of habitat cause the decrease of bat population. Theoutcome above provide the straight evidence and data of distribution of bats in China,which offer some theory basis for bat' protection and discussion of how bats adaptenvironment and evolve.
2.Bats living in the same cave in ZhaShui region was identified, described andsome related data also are measured. There are three species of bats, Hipposideros prattipratti Thomas, Rhinolophus rouxi sinicus Andersen and Miniopterus schreibersichinensis Thomas, which are identified by Zoological Institute, CAS.The findings showthat there are some distinctions in morphologic datum and ear related datum amongthese bats inhabiting the same cave, which determine the differences of echolocaioncalls, activity patterns related to hearing, foraging strategy and ecology niche. That is tosay, bats' echolocation activities and their morphologic characters have close connectionwith their foraging strategy and foraging habitat. From the morphologic characters ofHipposideros pratti pratti Tomas, we can conclude that they own the heaviest weight,the longest forearm length, the biggest figure, the longest wing, ear length and ear width.These factors decide that they are fit for long-distance flying, flying quickly andunskillfully. Hipposideros pratti pratti Tomas often forage in complicated region, attacktargets with the bigger shape and the ability of flying quickly, which can meet theirenergy needs. Rhinolophus rouxi sinicus Andersen with the lightweight, short totallength and forarm length, which we can conclude that their flying speed is slow, buthave the skillful flying ability. They can detect the little wing-beating insects incomparative simple condition, such as in dense foliages and narrow habitat around. Miniopterus schreibersi chinensis Thomas has the similarities with Rhinolophus rouxisinicus Andersen, but they ofen prey on in complicated room by gleaning, such as innarrow space. These results provide supplement and proof of morphologic database ofbats in China, and approve the theory again, which bats' echolocation activities andtheir morphologic characters have close connection with their foraging strategy andforaging habitat.
3. Bats living in the same cave in ZhaShui region were studied by mensuratingtheir hearing function and observing their aultra-structure of hair cells of cochlea. Byusing Tucker-Davis Technology(TDT)Biosig SystemⅢ, we gain the result that auditorybrainstem responses(ABR) of bats,which sensitive region is 12-48kHz.This outcome isbasically consistent with the ICR mouse's sensitive region that is 12-32kHz,and thatMicrotus mandarinus's sensitive region is 6-8kHz. The bats' cochlear hair cells wereobserved by scanning electron microscope, and the structure of bats'cochlear is similarto human being's. The inner hair cells (IHCs) of bats resemble any other mammals,which own only one row, and their stereocilia are short. At the basal of basilarmembrane, the stereocilia bundles of IHCs are thin and cuticular plate (CP) separatewith each other. On the contrary, at the apex of basilar membrane, the stereociliabundles are bushy and cuticular plate join together. At the same time, there aresignificant differences between the outer hair cells (OHCs) stereocilia of bats and othermammals's. The stereocilia of OHCs are extremely short. We can observe thephenomena of rupture and inosculation of stereocilia, which is alike senile deafness ofICR mouse. The OHCs bodies of bats have two leading characteristics. Firstly, theOHCs bodies are obviously short. The Deiters cell phalanges are of exaggerated, andthe cup formation of Deiters cell body enwrap the bottom of the OHCs.Secondly, TheOHCs body possess the flask-shaped in the basal turn of the cochlea, which don't likeother mammals'tube-shaped OHCs body. All of the results above indicate that bats andmostly any other little mammals have the similar audiogram in definite range, but alsohave some difference. Echolocating bats always possess of acuity hearing, whichinclude more available frequency. The structure of bats'cochlea is partly resembled withother mammals; meanwhile, there are some specilized aspects, which are considered asmorphologic adaptation to dealing with ultra-high frequency sound wave. All thefindings not only supply helps for the researchesof ethology and neurobiology ofecholocating animals, but also provide suggestions of hearing mechanism and deafness prevention in human beings.
引文
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