华中师范大学生命科学学院,武汉 430079
国家自然科学基金(31200832),湖北省自然科学基金 (2019CFB758)和华中师范大学中央高校基本科研业务费 (CCNU20TS018)资助项目。
School of Life Sciences, Central China Normal University, Wuhan 430079, China
This work was supported by grants from The National Natural Science Foundation of China (31200832), Hubei Provincial Natural Science Foundation of China (2019CFB758) and the Fundamental Research Funds of Central China Normal University (CCNU20TS018).
科学家以蝙蝠为模式动物,从听觉、回声定位和生态适应与演化等方面开展了研究,取得了令人瞩目的成果。为适应回声定位,蝙蝠听觉系统的结构和功能产生了明显的特化。从外周到中枢形成了对声频率极为有序的表征,甚至在恒频-调频(constant frequency-frequency modulation,CF-FM)蝙蝠耳蜗形成了所谓的听觉凹,以及听皮质功能组构也模块化,成为了具有代表性的特化象征。神经元反应的潜伏期对蝙蝠不仅是基本特性,也是回声定位行为调控的一部分;研究发现,有较长潜伏期的神经元具有较尖锐的回声-延迟调谐特性,而较短潜伏期的神经元则有较宽的回声-延迟调谐特性。蝙蝠听神经元对频率调谐的精准度亦远胜于人类和其他非回声定位动物;而且,源于耳蜗听觉凹的传入在各级听中枢均显示出对回声定位信号第二谐波CF成分的过度表征,以满足对靶物回声多普勒频移探测的需要。时程是回声定位蝙蝠发声信号主动改变的参数之一,而时程调谐神经元则提供了一种编码声音时相特征的重要神经机制,匹配了对回声定位信号时相信息加工的需要。在多种回声定位蝙蝠的听中枢还发现,有回声-延迟调谐神经元,它们不仅能对靶物距离进行调谐,而且也能对回声的方位角和俯仰方位进行调谐,从而在靶物位置的三维(3D)表征方面发挥重要的作用。在CF-FM蝙蝠下丘,神经元对行为相关的CF-FM声信号表现出single-on和double-on两种反应模式,可能分别由不同的局部神经回路所定型,它们分别行使对回声定位信号的加工。基于胡须蝠的听皮质神经元对回声定位信号的反应特性,可将其划分为处理不同回声信息的功能模块,如CF/CF区的神经元,负责处理多普勒频移幅度即靶物速度,FM/FM区神经元,对回声延迟或靶物距离敏感。对离皮质调控的研究表明,这种调控系统对皮质下的声信号加工,以及在成年后为适应环境而产生可塑性改变等提供了一种结构和功能的保障。研究还发现,在蝙蝠的听皮质内存在目标选择性神经元,它们对听觉目标以一种尺度不变性或大小恒常性方式产生反应,这一发现亦为哺乳动物在听觉的尺度不变性方面遵循共同机制的假说提供了证据。蝙蝠在飞航期间对3D空间的动态表征不仅存在于大脑皮质,也存在于听觉中脑,目前认为,蝙蝠在3D空间和飞行状态下,脑内不同类型的导航细胞(即神经元)能各自行使相应的功能,引导蝙蝠到达目的地。
Bats have been used as model animals to study their hearing, echolocation, ecological adaptation and evolution, and lots of remarkable results have been achieved. In order to adapt to echolocation, the structure and function of the auditory system in bats have developed distinct specializations. In the constant frequency-frequency modulation (CF-FM) bat cochlea, the so-called auditory fovea is formed, and the functional organization of auditory cortex is modular, which has become a representative specialized symbol. The latency of neuronal response is not only a fundamental characteristic of bats, but also a part of the regulation of echolocation behavior. It is found that neurons with longer latencies have sharper echo-delay tuning characteristics, while neurons with shorter latencies have wider echo-delay tuning characteristics. Frequency tuning of bat auditory neurons is far greater precision than humans and other non-echolocation animals. Moreover, the afferents originating from the cochlear auditory fovea show overrepresentation of CF components of the second harmonic of echolocation signals at all levels of auditory centers, so as to meet the needs of target echo Doppler drift detection. Duration is one of the actively changeable parameters of echolocating bat vocalization signals. The duration-tuned neurons provide an important neural mechanism for encoding the temporal features of sound, matching the need for processing the temporal information of echolocation signals. Echo-delay tuned neurons have been found in the auditory center of many echolocation bats, which not only can tune the target range, but also the azimuth and elevation of the echo, thus playing an important role in the three-dimensional representation of the target location. In the inferior colliculus of CF-FM bats, neurons showed single-on and double-on response patterns to the CF-FM sound signal, which may be shaped by different local neural circuits. Based on the response properties of the auditory cortex neurons of mustached bats to echolocation signals, the auditory cortex neurons of Pteronotus parnellii can be divided into functional modules for processing different echo information. For example, neurons in CF/CF area are responsible for processing Doppler-shift magnitude, i.e. target velocity. The neurons in FM/FM area are sensitive to echo delay or target distance. Studies on corticofugal control indicate that this control system provides a structural and functional guarantee for subcortical acoustic signal processing and plasticity changes in adaptation to the environment for adult bats. It has also been found that object-selective neurons exist in the auditory cortex of bats, which respond to auditory objects in a scale-invariant or size-constancy manner. This finding also provides evidence for the hypothesis that mammals follow a common mechanism for scale invariance of hearing. The dynamic representation of 3D space during flight not only exists in the cerebral cortex, but also in the auditory midbrain. It is now thought that different types of navigation neurons in the bats’ brains perform their respective functions to guide them to their destinations in 3D space and flight.
付子英,唐佳,陈其才.综述与专论: 蝙蝠,听觉和回声定位研究的模型动物[J].生物化学与生物物理进展,2022,49(5):883-896
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