Wenstrup J J
Department of Neurobiology, Northeastern Ohio Universities College of Medicine, Rootstown, Ohio 44272-0095, USA.
J Neurophysiol. 1999 Nov;82(5):2528-44. doi: 10.1152/jn.1999.82.5.2528.
The auditory cortex of the mustached bat (Pteronotus parnellii) displays some of the most highly developed physiological and organizational features described in mammalian auditory cortex. This study examines response properties and organization in the medial geniculate body (MGB) that may contribute to these features of auditory cortex. About 25% of 427 auditory responses had simple frequency tuning with single excitatory tuning curves. The remainder displayed more complex frequency tuning using two-tone or noise stimuli. Most of these were combination-sensitive, responsive to combinations of different frequency bands within sonar or social vocalizations. They included FM-FM neurons, responsive to different harmonic elements of the frequency modulated (FM) sweep in the sonar signal, and H1-CF neurons, responsive to combinations of the bat's first sonar harmonic (H1) and a higher harmonic of the constant frequency (CF) sonar signal. Most combination-sensitive neurons (86%) showed facilitatory interactions. Neurons tuned to frequencies outside the biosonar range also displayed combination-sensitive responses, perhaps related to analyses of social vocalizations. Complex spectral responses were distributed throughout dorsal and ventral divisions of the MGB, forming a major feature of this bat's analysis of complex sounds. The auditory sector of the thalamic reticular nucleus also was dominated by complex spectral responses to sounds. The ventral division was organized tonotopically, based on best frequencies of singly tuned neurons and higher best frequencies of combination-sensitive neurons. Best frequencies were lowest ventrolaterally, increasing dorsally and then ventromedially. However, representations of frequencies associated with higher harmonics of the FM sonar signal were reduced greatly. Frequency organization in the dorsal division was not tonotopic; within the middle one-third of MGB, combination-sensitive responses to second and third harmonic CF sonar signals (60-63 and 90-94 kHz) occurred in adjacent regions. In the rostral one-third, combination-sensitive responses to second, third, and fourth harmonic FM frequency bands predominated. These FM-FM neurons, thought to be selective for delay between an emitted pulse and echo, showed some organization of delay selectivity. The organization of frequency sensitivity in the MGB suggests a major rewiring of the output of the central nucleus of the inferior colliculus, by which collicular neurons tuned to the bat's FM sonar signals mostly project to the dorsal, not the ventral, division. Because physiological differences between collicular and MGB neurons are minor, a major role of the tecto-thalamic projection in the mustached bat may be the reorganization of responses to provide for cortical representations of sonar target features.
髯蝠(Pteronotus parnellii)的听觉皮层展现出了一些在哺乳动物听觉皮层中所描述的最为高度发达的生理和组织特征。本研究考察了内侧膝状体(MGB)中的反应特性和组织结构,这些特性和结构可能促成了听觉皮层的这些特征。在427个听觉反应中,约25%具有简单的频率调谐,呈现单一的兴奋性调谐曲线。其余反应则使用双音或噪声刺激展现出更复杂的频率调谐。其中大多数对组合敏感,对声纳或社交发声中不同频段的组合有反应。它们包括FM - FM神经元,对声纳信号中调频(FM)扫描的不同谐波成分有反应,以及H1 - CF神经元,对蝙蝠的第一声纳谐波(H1)和恒定频率(CF)声纳信号的更高谐波的组合有反应。大多数对组合敏感的神经元(86%)表现出易化性相互作用。调谐到生物声纳范围之外频率的神经元也表现出对组合敏感的反应,这可能与对社交发声的分析有关。复杂的频谱反应分布在MGB的背侧和腹侧区域,构成了这种蝙蝠对复杂声音分析的一个主要特征。丘脑网状核的听觉部分也以对声音的复杂频谱反应为主。腹侧区域基于单音调谐神经元的最佳频率和对组合敏感神经元的更高最佳频率进行音调定位组织。最佳频率在腹外侧最低,向背侧然后向腹内侧增加。然而,与FM声纳信号更高谐波相关的频率表征大幅减少。背侧区域的频率组织不是音调定位的;在MGB的中间三分之一区域内,对第二和第三谐波CF声纳信号(60 - 63千赫兹和90 - 94千赫兹)的组合敏感反应出现在相邻区域。在吻侧三分之一区域,对第二、第三和第四谐波FM频段的组合敏感反应占主导。这些FM - FM神经元被认为对发出的脉冲和回声之间的延迟具有选择性,表现出一定的延迟选择性组织。MGB中频率敏感性的组织表明下丘中央核的输出发生了重大的重新布线,通过这种方式,调谐到蝙蝠FM声纳信号的下丘神经元大多投射到背侧而非腹侧区域。由于下丘和MGB神经元之间的生理差异较小,在髯蝠中顶盖 - 丘脑投射的一个主要作用可能是对反应进行重新组织,以实现声纳目标特征的皮层表征。