An ultra-sparse code underlies the generation of neural sequences in a songbird

Sequences of motor activity are encoded in many vertebrate brains by complex spatio-temporal patterns of neural activity; however, the neural circuit mechanisms underlying the generation of these pre-motor patterns are poorly understood. In songbirds, one prominent site of pre-motor activity is the forebrain robust nucleus of the archistriatum (RA), which generates stereotyped sequences of spike bursts during song and recapitulates these sequences during sleep. We show that the stereotyped sequences in RA are driven from nucleus HVC (high vocal centre), the principal pre-motor input to RA. Recordings of identified HVC neurons in sleeping and singing birds show that individual HVC neurons projecting onto RA neurons produce bursts sparsely, at a single, precise time during the RA sequence. These HVC neurons burst sequentially with respect to one another. We suggest that at each time in the RA sequence, the ensemble of active RA neurons is driven by a subpopulation of RA-projecting HVC neurons that is active only at that time. As a population, these HVC neurons may form an explicit representation of time in the sequence. Such a sparse representation, a temporal analogue of the 'grandmother cell' concept for object recognition, eliminates the problem of temporal interference during sequence generation and learning attributed to more distributed representations.     

Support for a synaptic chain model of neuronal sequence generation

In songbirds, the remarkable temporal precision of song is generated by a sparse sequence of bursts in the premotor nucleus HVC. To distinguish between two possible classes of models of neural sequence generation, we carried out intracellular recordings of HVC neurons in singing zebra finches (Taeniopygia guttata). We found that the subthreshold membrane potential is characterized by a large, rapid depolarization 5-10 ms before burst onset, consistent with a synaptically connected chain of neurons in HVC. We found no evidence for the slow membrane potential modulation predicted by models in which burst timing is controlled by subthreshold dynamics. Furthermore, bursts ride on an underlying depolarization of ～10-ms duration, probably the result of a regenerative calcium spike within HVC neurons that could facilitate the propagation of activity through a chain network with high temporal precision. Our results provide insight into the fundamental mechanisms by which neural circuits can generate complex sequential behaviours.     

Contributions of an avian basal ganglia-forebrain circuit to real-time modulation of song

Cortical-basal ganglia circuits have a critical role in motor control and motor learning. In songbirds, the anterior forebrain pathway (AFP) is a basal ganglia-forebrain circuit required for song learning and adult vocal plasticity but not for production of learned song. Here, we investigate functional contributions of this circuit to the control of song, a complex, learned motor skill. We test the hypothesis that neural activity in the AFP of adult birds can direct moment-by-moment changes in the primary motor areas responsible for generating song. We show that song-triggered microstimulation in the output nucleus of the AFP induces acute and specific changes in learned parameters of song. Moreover, under both natural and experimental conditions, variability in the pattern of AFP activity is associated with variability in song structure. Finally, lesions of the output nucleus of the AFP prevent naturally occurring modulation of song variability. These findings demonstrate a previously unappreciated capacity of the AFP to direct real-time changes in song. More generally, they suggest that frontal cortical and basal ganglia areas may contribute to motor learning by biasing motor output towards desired targets or by introducing stochastic variability required for reinforcement learning.     

断鸣管神经支对白腰文鸟鸣声的影响

为探究鸣管神经支（NXⅡts）对发声控制的作用以及验证鸣禽发声的侧别优势和性别差异,以成年白腰文鸟为实验材料,记录其正常鸣声及切断鸣管神经支后的鸣声.用Cool Edit Pro和Wavesurfer 等声音处理和分析软件,比较和分析雄鸟和雌鸟、断单侧和双侧鸣管神经前后声学特征的差异.结果表明,成年白腰文鸟的NXⅡts可单侧支配鸣肌,切断左侧NXⅡts将导致声音严重受损,而断右侧则影响不大,表明具有左侧优势.两侧NXⅡts既相互协调又各有侧重,左侧产生较高频率的声音,而右侧则有较高的熵值,使得音节稳定性和调频一致性增强.雄鸟左侧NXⅡts控制较高的音量和较长的音节,而雌鸟的音量和音节时长两侧均可以独立调节.断双侧神经对音量、音调和音色都产生明显影响.     

Species-typical songs in white-crowned sparrows tutored with only phrase pairs

Modern theories of learned vocal behaviours, such as human speech and singing in songbirds, posit that acoustic communication signals are reproduced from memory, using auditory feedback. The nature of these memories, however, is unclear. Here we propose and test a model for how complex song structure can emerge from sparse sequence information acquired during tutoring. In this conceptual model, a population of combination-sensitive (phrase-pair) detectors is shaped by early exposure to song and serves as the minimal representation of the template necessary for generating complete song. As predicted by the model, birds that were tutored with only pairs of normally adjacent song phrases were able to assemble full songs in which phrases were placed in the correct order; birds that were tutored with reverse-ordered phrase pairs sang songs with reversed phrase order. Birds that were tutored with all song phrases, but presented singly, failed to produce normal, full songs. These findings provide the first evidence for a minimal requirement of sequence information in the acoustic model that can give rise to correct song structure.     

分布式多空间数据库复杂时态数据提取技术

传统方法实现过程复杂、历史复杂时态数据的片面性,导致其无法全面地描述时态数据;且相似性计算无法准确匹配具有动态性与复杂性的时态数据,造成提取精度低。为此,提出一种新的分布式多空间数据库复杂时态数据提取技术。设计动态RBF神经网络,对分布式多空间数据库中未知动态进行识别和建模;通过建模结果完成对复杂时态数据的描述。依据加权关联规则与时态关联规则对支持度和置信度的定义,获取T-FS-tree加权时态关联规则中支持度和置信度。将复杂时态数据描述序列、最小支持度、最小置信度作为输入,将加权时态关联规则作为输出,建立T-FS-tree加权时态关联规则挖掘算法。按照向量计算获取加权时态频繁1项集以及频繁2项集,依据获取的加权时态频繁项集建立初始频繁项集树;依据初始频繁项集树获取全部时态频繁项集;通过获取的频繁项集产生加权时态关联规则。从所有关联规则中选择优先度高的规则,构建的复杂时态数据提取器,实现复杂时态数据提取。实验结果表明,所提方法复杂性低,提取结果更加全面、可靠,有很高的准确性。     

Decrystallization of adult birdsong by perturbation of auditory feedback.

Young birds learn to sing by using auditory feedback to compare their own vocalizations to a memorized or innate song pattern; if they are deafened as juveniles, they will not develop normal songs. The completion of song development is called crystallization. After this stage, song shows little variation in its temporal or spectral properties. However, the mechanisms underlying this stability are largely unknown. Here we present evidence that auditory feedback is actively used in adulthood to maintain the stability of song structure. We found that perturbing auditory feedback during singing in adult zebra finches caused their song to deteriorate slowly. This 'decrystallization' consisted of a marked loss of the spectral and temporal stereotypy seen in crystallized song, including stuttering, creation, deletion and distortion of song syllables. After normal feedback was restored, these deviations gradually disappeared and the original song was recovered. Thus, adult birds that do not learn new songs nevertheless retain a significant amount of plasticity in the brain.     

Complex auditory behaviour emerges from simple reactive steering

The recognition and localization of sound signals is fundamental to acoustic communication. Complex neural mechanisms are thought to underlie the processing of species-specific sound patterns even in animals with simple auditory pathways. In female crickets, which orient towards the male's calling song, current models propose pattern recognition mechanisms based on the temporal structure of the song. Furthermore, it is thought that localization is achieved by comparing the output of the left and right recognition networks, which then directs the female to the pattern that most closely resembles the species-specific song. Here we show, using a highly sensitive method for measuring the movements of female crickets, that when walking and flying each sound pulse of the communication signal releases a rapid steering response. Thus auditory orientation emerges from reactive motor responses to individual sound pulses. Although the reactive motor responses are not based on the song structure, a pattern recognition process may modulate the gain of the responses on a longer timescale. These findings are relevant to concepts of insect auditory behaviour and to the development of biologically inspired robots performing cricket-like auditory orientation.     

斑胸草雀鸣肌的形态观察

运用解剖学技术对斑胸草雀的左右侧鸣肌进行解剖,并对其进行测量.结果表明,斑胸草雀有2对外鸣肌和4对内鸣肌.两侧鸣肌质量差异显著,右侧较重.两侧NX IIts的直径差异显著,右侧较粗.在形态学上验证了鸣肌和NX IIts的右侧优势.初步分析了鸣肌各部分的结构及功能,为进一步研究鸣肌在鸣禽鸣唱过程中的作用提供了基础资料.     

鸣禽鸟脑的季节可塑性

呜禽鸟的发声控制脑区具有明显的季节可塑性变化，它是鸣啭季节差异的神经基础．目前鸣禽成鸟的发声控制系统已成为研究中枢神经系统形态和功能可塑性的重要模型．本文着重就引起发声控制核团或脑区体积改变的诱导因素和触发体积改变的机制进行探讨．     

Role of experience and oscillations in transforming a rate code into a temporal code

In the vast majority of brain areas, the firing rates of neurons, averaged over several hundred milliseconds to several seconds, can be strongly modulated by, and provide accurate information about, properties of their inputs. This is referred to as the rate code. However, the biophysical laws of synaptic plasticity require precise timing of spikes over short timescales (<10 ms). Hence it is critical to understand the physiological mechanisms that can generate precise spike timing in vivo, and the relationship between such a temporal code and a rate code. Here we propose a mechanism by which a temporal code can be generated through an interaction between an asymmetric rate code and oscillatory inhibition. Consistent with the predictions of our model, the rate and temporal codes of hippocampal pyramidal neurons are highly correlated. Furthermore, the temporal code becomes more robust with experience. The resulting spike timing satisfies the temporal order constraints of hebbian learning. Thus, oscillations and receptive field asymmetry may have a critical role in temporal sequence learning.     

An oscillatory neural network and dynamical associative memory

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鸣禽鸣啭系统的可塑性及长时程增强研究

鸣禽的鸣唱控制系统已成为研究神经系统与学习、行为和发育相关的一个重要模型.鸣禽的鸣啭表现出一种复杂的学习过程.鸣禽学习鸣啭的过程可以分为两个阶段.在感觉学习期,幼鸟必须听到成鸟的鸣啭,并形成鸣啭模板记忆;在感觉运动学习期,鸣禽通过听觉反馈与模板匹配逐步建立稳定的鸣啭.该文对近年来鸣禽鸣啭学习过程中的新生神经元及长时程增强研究进展进行综述.     

Performance variability enables adaptive plasticity of 'crystallized' adult birdsong

Significant trial-by-trial variation persists even in the most practiced skills. One prevalent view is that such variation is simply 'noise' that the nervous system is unable to control or that remains below threshold for behavioural relevance. An alternative hypothesis is that such variation enables trial-and-error learning, in which the motor system generates variation and differentially retains behaviours that give rise to better outcomes. Here we test the latter possibility for adult bengalese finch song. Adult birdsong is a complex, learned motor skill that is produced in a highly stereotyped fashion from one rendition to the next. Nevertheless, there is subtle trial-by-trial variation even in stable, 'crystallized' adult song. We used a computerized system to monitor small natural variations in the pitch of targeted song elements and deliver real-time auditory disruption to a subset of those variations. Birds rapidly shifted the pitch of their vocalizations in an adaptive fashion to avoid disruption. These vocal changes were precisely restricted to the targeted features of song. Hence, birds were able to learn effectively by associating small variations in their vocal behaviour with differential outcomes. Such a process could help to maintain stable, learned song despite changes to the vocal control system arising from ageing or injury. More generally, our results suggest that residual variability in well learned skills is not entirely noise but rather reflects meaningful motor exploration that can support continuous learning and optimization of performance.     

离散化四足机器人自由步态控制方法

为了实现离散化四足机器人自由步态的控制,提出一种新的基于中枢神经模式发生器(CPG)的自由步态控制方法。介绍了离散化四足机器人模型,在已确定的地形中,设定四足机器人起始点与抵达点的状态。将连续步态按照离散化步态完成排序,形成排序集合。在此基础上,利用中枢神经模式发生器CPG,采用周期性振荡信号对离散化四足机器人腿部各关节进行控制,给出单独神经元模型。为了便于分析,使用互抑神经元构成的振荡器对神经元的输出信号进行改善,通过该振荡器产生规律的振荡信号,以控制离散化四足机器人完成自由步态移动。实验结果表明,所提方法能够有效控制离散化四足机器人实现自由步态移动。     

基于多尺度时序交互的第一人称行为识别方法

对于第一人称行为识别任务,现有方法大多使用了目标边界框和人眼视线数据等非行为类别标签对深度神经网络进行辅助监督,以使其关注视频中手部及其交互物体所在区域。这既需要更多的人工标注数据,又使得视频特征的提取过程变得更为复杂。针对该问题,提出了一种多尺度时序交互模块,通过不同尺度的3D时序卷积使2D神经网络提取的视频帧特征进行时序交互,从而使得单一视频帧的特征融合其近邻帧的特征。在只需行为类别标签作监督的情况下,多尺度时序交互能够促使网络更加关注第一人称视频中手部及其交互物体所在区域。实验结果表明,提出的方法在识别准确率优于现有第一人称行为识别方法。     

Interruption of a basal ganglia-forebrain circuit prevents plasticity of learned vocalizations

Birdsong, like speech, is a learned vocal behaviour that relies greatly on hearing; in both songbirds and humans the removal of auditory feedback by deafening leads to a gradual deterioration of adult vocal production. Here we investigate the neural mechanisms that contribute to the processing of auditory feedback during the maintenance of song in adult zebra finches. We show that the deleterious effects on song production that normally follow deafening can be prevented by a second insult to the nervous system--the lesion of a basal ganglia-forebrain circuit. The results suggest that the removal of auditory feedback leads to the generation of an instructive signal that actively drives non-adaptive changes in song; they also suggest that this instructive signal is generated within (or conveyed through) the basal ganglia-forebrain pathway. Our findings provide evidence that cortical-basal ganglia circuits may participate in the evaluation of sensory feedback during calibration of motor performance, and demonstrate that damage to such circuits can have little effect on previously learned behaviour while conspicuously disrupting the capacity to adaptively modify that behaviour.     

How sleep affects the developmental learning of bird song

Sleep affects learning and development in humans and other animals, but the role of sleep in developmental learning has never been examined. Here we show the effects of night-sleep on song development in the zebra finch by recording and analysing the entire song ontogeny. During periods of rapid learning we observed a pronounced deterioration in song structure after night-sleep. The song regained structure after intense morning singing. Daily improvement in similarity to the tutored song occurred during the late phase of this morning recovery; little further improvement occurred thereafter. Furthermore, birds that showed stronger post-sleep deterioration during development achieved a better final imitation. The effect diminished with age. Our experiments showed that these oscillations were not a result of sleep inertia or lack of practice, indicating the possible involvement of an active process, perhaps neural song-replay during sleep. We suggest that these oscillations correspond to competing demands of plasticity and consolidation during learning, creating repeated opportunities to reshape previously learned motor skills.     

Projection neurons within a vocal motor pathway are born during song learning in zebra finches

Many birds learn song during a restricted 'sensitive' period. Juveniles memorize a song model, and then learn the pattern of muscle contractions necessary to reproduce the song. Of the neural changes accompanying avian song learning, perhaps the most remarkable is the production of new neurons which are inserted into the hyperstriatum ventralis pars caudalis (HVc), a region critical for song production. We report here that in young male zebra finches many of the new neurons incorporated into the HVc innervate the robust nucleus of the archistriatum (RA) which projects to motor neurons controlling the vocal musculature. Furthermore, far fewer of these new neurons are incorporated into the HVc of either adult males that are beyond the sensitive learning period, or young females (who do not develop song). Thus, a major portion of the vocal motor pathway is actually created during song learning. This may enable early sensory experience and vocal practice to not only modify existing neuronal circuits, but also shape the insertion and initial synaptic contacts of neurons controlling adult song.