鸟类的发声系统和调控机制

鸟类是具有复杂声行为的动物,其拥有特殊的发声器官——鸣管。尽管鸣禽与非鸣禽的发声特性和发声器官解剖学差异较大,但是两者发声运动控制模式相似。文章综述了近年来鸟类呜声研究的新进展,重点比较了呜禽和非鸣禽发声器官的结构功能特点和发声特性调控的异同。作为一种动物模型,鸟类发声系统能为人类语言学习等研究提供借鉴。     

鸣禽与人类发声器官及发声行为神经控制通路的比较

与人类语言学习或形成一样,鸣禽鸣唱也是一种发声学习行为,二者具有一定的相似性,例如发声学习过程均需听觉反馈的参与,幼年期具有更强的发声学习能力,可对复杂的声学结构和音节序列进行控制等。尽管鸣禽和人类的发声器官在结构上有很大差异,但二者发声的物理机制仍表现出很强的相似性。虽然相比于其他哺乳动物,鸣禽和人类的亲缘关系很远,但通过对比发声行为产生的基础通路——脑干先天发声控制通路,以及与发声学习相关的更高神经水平的发声运动和学习通路脑区位置、相互联系、功能及基因表达谱,提示鸣禽鸣唱和人类语言的神经控制具有一定的进化相似性。这些共同特征使得鸣禽成为了研究发声学习的理想模型。本文对鸣禽与人类的发声器官及发声行为的神经控制通路进行了比较,并对鸣禽模型在人类失语症治疗研究中潜在的应用前景进行了展望,以期为研究人类语言学习的神经机制及语言障碍的治疗带来理论参考和借鉴。     

鸣禽发声控制神经通路的左侧优势

鸟类鸣叫机理是近年比较活跃的研究领域,已证明可为人类发声机理提供实验模型。本文作者通过自己的研究工作,对鸣禽控制发声的外围和中枢侧向优势作了简要介绍。     

鸣禽发声学习行为的神经生物学机制

鸣禽发声学习的控制系统主要由一条直接神经通路和一条辅助神经通路组成,由前脑控制发声学习的最高中枢ＨＶＣ、旁嗅叶的Ｘ区和巨细胞核外侧部（ｌＭＡＮ）组成的辅助通路,对鸟类发声学习行为的发育和调制具有重要作用。发声控制系统中神经元类型、数量及再生与更替、神经组构及其重组、神经介质和受体的分布等差异,决定了鸣禽在发声学习行为表现的差异以及性双态性。本文对近年鸟类控制发声学习行煌神经生物学机制的进展作了较为     

综述与专论: 鸣禽模型在语言学习中承担独特的生物学角色

鸣禽鸣唱与人类语言相似，是一种复杂的发声学习行为，并受脑中一组相互联系的神经核团调控。该组核团与人类发声控制相关脑区具有一定程度的结构同源性，并可能共享某些发声学习调控机制。因此，鸣禽成为研究发声学习神经机理的重要模式动物，不仅对鸟类语言学习，也可为揭示人类语言学习的神经过程和语言障碍的治疗提供重要参考借鉴。本文基于本课题组长期坚持的研究方向，较系统地概述了国内外鸣禽鸣唱行为研究的历史、重要发现和进展，及其为相关中枢神经系统疾病治疗带来的启示。     

鸟类鸣声初探

对鸟类鸣声的研究是生物声学研究中最为活跃的领域之一,对鸟鸣声的应用在很多方面具有十分广阔的前景。鸟类的鸣声可分为鸣唱和鸣叫,不同鸟类具有不同的鸣叫能力,可据此将鸟类分为鸣禽类和非鸣禽类两大类。文章初步综述了鸟类鸣声的相关知识,对鸣禽和非鸣禽发声器官的结构功能进行了比较。     

鸣禽前脑基底神经节X区与语言学习

成年鸣禽鸣唱语句的形成和维持依赖于听觉反馈。X区是鸣禽前脑回路的一个重要核团,对鸣禽的发声学习和语言结构的稳定有重要作用。X区在解剖结构、电生理学以及神经化学等方面的特性与哺乳类基底神经节极为相似。对鸣禽前脑X区的研究有助于揭示人类语言学习的中枢机制。本文对近年来鸣禽X区的相关研究进展,包括鸣禽X区的结构特征、电生理特性及神经化学特征予以阐述。     

鸣禽发声控制核团内的神经递质及其在发声学习中的作用

神经递质在鸣禽脑中不仅是神经元间信号传递中介物质,还有资料表明它们通过在习鸣敏感期影响发声控制团间的突触联系的形成和突触可塑性,从而对鸣转类型的确定和巩固起重要作用,本文着重介绍了鸣禽发声控制核团内神经递质的分布及变化情况,并就神经递质在发声学习中的作用进行了探讨。     

鸣禽发声学习记忆与即刻早期基因

鸣禽受到声音信号的刺激或自身表现出发声行为时,脑内即刻早期基因(immediate early gene,IEG)能迅速被激活而表达.其中zenk、c-fos和c-jun表达的脑区及水平与鸟在鸣唱时神经元的活动区域及活动程度相一致,暗示IEG在鸣禽发声学习记忆中起重要作用.     

成年鸣禽习鸣脑区的神经发生

成年鸣禽习鸣脑区可产生新的神经元。这些新生神经元沿神经胶质纤维迁移至鸣禽前脑高级发声控制与发声学习中枢,加入到原有的神经回路,参与发声学习记忆的感知与运动过程。这提示神经可塑性不仅适用于脑的发育,而且也是成年动物脑功能神经回路的一种正常过程。     

THE MEASUREMENT OF VOCAL AMPLITUDE AND VOCAL RADIATION PATTERN IN BLUE MONKEYS AND GREY-CHEEKED MANGABEYS

ABSTRACT

The substitution method was adopted from industrial acoustics (Francois and de Montussaint 1972) to “eliminate the influence of the environment” on measurements of the amplitude of vocalizations given by blue monkeysCercopithecus mitis and grey-cheeked mangabeysCercocebus albigena. Measurements were conducted of sound power and sound pressure level of representative utterances. Monkey vocal radiation patterns were also measured. The results showed that vocal amplitude ranged from 62 dB to 100 dB in sound pressure (re 1 pw). At a distance of 2 m, the loudest calls approached an amplitude of 110 dB SPL, a level about equal to the loudest human yell. The measurements of call amplitude conducted here exceeded those derived from the field by approximately 10 dB. It was shown that the discrepancy in amplitude between these laboratory based measurements and earlier measurements conducted under field conditions (Waser and Waser 1977) was probably due to destructive interference between the direct wave and the “ground wave”, a phase shifted wave reflected from the ground. Measurements of radiation patterns of primate vocalizations showed that, like human speech, directivity was a function of frequency, with high-frequency components being radiated more directionally than lower-frequency components. However, primate utterances were in general radiated more omnidirectionally than was human speech.     

鸣禽鸟发声行为的激素调节

讨论了性类固醇激素对鸣食发声行为及其神经回路的影响,从细胞回路水平,行为水平对性激素在幼年鸣食发声学习和成年鸣食鸣啭可塑性中的作用进行了全面论述,介绍了国内外在鸣禽发声这一研究领域的最新进展,对深入开展动物学习记忆神经机制的研究具有借鉴作用。     

PRIMARY SQUAMOUS CELL CARCINOMA OF THE FALSE VOCAL CORD

Primary squamous cell carcinoma of the false vocal cord may arise on the surface mucosa or may arise beneath it and continue to grow deeply, presenting only a smooth tumescence of the area. These lesions may not cause hoarseness until late in the course of development. Diagnosis of submucosal primary lesions may present difficulty.Widefield laryngectomy is recommended for small primary lesions of the surface mucosa of the false vocal cord. Such lesions do not show edema of the tissues or deep ulceration and do not cause limitation of motion of the false or true vocal cords.For advanced lesions of the false vocal cord which arise on the surface mucosa and cause edema, ulceration and limitation of motion without enlargement of cervical nodes, widefield laryngectomy and elective block dissection of the neck should be done at the primary operation. Patients with such a primary lesion and metastasis to cervical lymph nodes, which are resectable, should be treated in a like manner.Patients with submucosal primary squamous cell carcinoma of the false vocal cord should be treated with widefield laryngectomy and block dissection of the neck, whether or not palpable resectable lymph nodes are noted in the neck. Apparently these lesions metastasize early and widely.     

新纹状体巨细胞核外侧部（LMAN）与鸣禽鸣唱学习可塑性

鸣禽鸣唱控制系统的前端脑通路（anterior forebrain pathway, AFP）在鸣唱学习中发挥着重要作用。新纹状体巨细胞核外侧部（lateral magnocellular nucleus of the anterior neostriatum, LMAN）是AFP的最后一级输出核团,AFP中的信号通过LMAN传导到弓状皮质栎核（robust nucleus of the arcopallium, RA）,与高级发声中枢（high vocal centre,HVC）共同调节RA的活动,从而影响鸣禽的发声行为。LMAN可能通过其与RA的单突触连接来影响鸣唱可塑性。文章对近年来LMAN在鸣唱学习可塑性方面的研究进行综述。     

VOCAL RESPONSES OF GREEN BARBET POPULATIONS,AND THE TAXONOMIC IMPLICATIONS

Dowsett-Lemaire, F. & Dowsett, R. J. 1987. Vocal responses of Green Barbet populations, and the taxonomic implications. Ostrich 58:160-163.

The songs of the following populations of the Green Barbet Stactolaemu olivacea are described and found to be very similar: coastal Kenya (olivacea), northern (rung weensis) and southern Malawi (belcheri), and Zululand (woodwardi). Birds of these isolated populations react readily to the playback of songs from other areas. Vocal similarity between these isolates suggests that they are no more than subspecifically distinct, including the Zululand population.     

VOCAL AND OTHER BEHAVIOUR OF KENYAN BLACK-COLLARED BARBETS LYBIUS TORQUATUS

We describe 12 visual and six acoustical displays of Black-collared Barbets, all associated with interactions of these sexually monomorphic barbets in greeting ceremonies and in antiphonal duets. Certain of the displays, e. g., bill directing, smooth posture and chatter calls largely are associated with multiple-barbet interactions and greeting ceremonies involving more than two barbets, and seem to represent purely agonistic displays. Other displays, most notably the bobbing and floating flight displays, and the tyaw call, are associated with preduet greeting ceremonies involving the presumed pair. In every case duets were performed by a presumed pair, following a greeting ceremony and visual displays accompany the duetting, which usually becomes precisely synchronized after the initial few notes. Four to 17 notes of each of two types, a usually lower pitched, short and mainly double type A note that introduces most duets, and a higher pitched, usually single, clear type B note that often terminates the duet. Variation in the notes and in their tempos within duets suggests no clearly defined temporal indicators for achieving, maintaining, and concluding the duets. The A and B notes cannot yet be assigned to one or other sex and similarity in form and development of the notes, from notes of greeting ceremonies, suggests the possibility that under some circumstances a bird of one sex may be able to deliver notes of either type. The duets are delivered from singing posts within the group's (or pair's) territory and usually only one duet is uttered from a site before the birds shift to another singing post. The duetting is compared with that of other barbets and appears to be among the best synchronized and most precise in the family. Comparisons with non-barbet duetting species are difficult because of the special habits and ecology of the barbets and uncertainty as to the sexual rôle of the duetters.     

鸣禽前脑发声控制核团的雌雄差别

本文应用尼氏染色组织学方法,对黄喉鹀(Emberiza elegans)、黄雀(Carduclis spinus)和燕雀(Fringilla montifringilla)三种鸣禽的前脑发声控制核团(HVc,RA,Area X)进行了观察和比较。结果表明,这些核团的体积存在着显著的性双形性。雄鸟的核团体积均大于雌鸟(P<0.001)。说明鸟类鸣啭行为的性别差异是由其神经结构的形态不同所造成的。