Vocal learning

声乐学习
  • 文章类型: Journal Article
    背景声乐学习是一种罕见的,融合的特征,是人类语言和鸟鸣的基础。叉头盒P2(FoxP2)转录因子似乎对于两种类型的学习信号都是必需的,因为人类FoxP2的突变会导致言语障碍,并破坏其在斑马雀中的表达会损害男性特定的歌曲学习。在幼年和成年雄性雀类中,在歌唱过程中,纹状体FoxP2mRNA和蛋白质在歌曲专用神经元内急剧下降,表明其转录靶标也是行为调控的。鸣鸟中这些目标的身份,以及它们是否因性别而异,发育和/或行为条件,基本上是未知的。结果在这里,我们使用染色质免疫沉淀,然后测序(ChIP-Seq)来鉴定男性和女性中FoxP2结合的基因组位点,少年和成人,唱歌和不唱歌的鸟。我们的结果表明,强大的FoxP2结合集中在基因的推定启动子区域。可能被FoxP2结合的基因数量因条件而异,建议与性别相关的候选人目标的专门角色,年龄,和行为状态。我们通过生物信息验证了这些结合靶标,与以前的研究和生物化学的比较,使用推定靶基因的抗体进行免疫组织化学。基因本体论分析显示,人类语言和语言相关功能仅在男性中丰富,与该物种中歌曲学习的性二态一致。相对于成年人,在青少年中发现的此类目标较少,这表明这个监管网络随着成熟而扩展。在歌唱条件下发现最少的与言语有关的目标,与有据可查的歌唱驱动的FoxP2在鸣禽纹状体中的下调一致。总体结论,这些数据提供了FoxP2在鸟类发声学习者中的监管格局的初始目录,为未来的研究提供几十个目标基因,并提供对声乐学习的分子基础的见解。
    UNASSIGNED: Vocal learning is a rare, convergent trait that is fundamental to both human speech and birdsong. The Forkhead Box P2 (FoxP2) transcription factor appears necessary for both types of learned signals, as human mutations in FoxP2 result in speech deficits, and disrupting its expression in zebra finches impairs male-specific song learning. In juvenile and adult male finches, striatal FoxP2 mRNA and protein decline acutely within song-dedicated neurons during singing, indicating that its transcriptional targets are also behaviorally regulated. The identities of these targets in songbirds, and whether they differ across sex, development and/or behavioral conditions, are largely unknown.
    UNASSIGNED: Here we used chromatin immunoprecipitation followed by sequencing (ChIP-Seq) to identify genomic sites bound by FoxP2 in male and female, juvenile and adult, and singing and non-singing birds. Our results suggest robust FoxP2 binding concentrated in putative promoter regions of genes. The number of genes likely to be bound by FoxP2 varied across conditions, suggesting specialized roles of the candidate targets related to sex, age, and behavioral state. We validated these binding targets both bioinformatically, with comparisons to previous studies and biochemically, with immunohistochemistry using an antibody for a putative target gene. Gene ontology analyses revealed enrichment for human speech- and language-related functions in males only, consistent with the sexual dimorphism of song learning in this species. Fewer such targets were found in juveniles relative to adults, suggesting an expansion of this regulatory network with maturation. The fewest speech-related targets were found in the singing condition, consistent with the well-documented singing-driven down-regulation of FoxP2 in the songbird striatum.
    UNASSIGNED: Overall, these data provide an initial catalog of the regulatory landscape of FoxP2 in an avian vocal learner, offering dozens of target genes for future study and providing insight into the molecular underpinnings of vocal learning.
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  • 文章类型: Journal Article
    背景:大多数声乐学习物种表现出早期的关键时期,在此期间,它们的声乐控制神经回路有助于获得新的发声。一些分类单元,最著名的是人类和鹦鹉,在整个成年期保留一定程度的神经行为可塑性,但是这种可塑性的程度和潜在的神经遗传学机制仍不清楚。转录因子FoxP2在鸣鸟和鹦鹉声音控制核中的差异表达先前已被确定为促进声音学习的关键模式。我们假设,开放式学习者的声乐学习对认知能力下降的韧性将反映在神经FoxP2表达中没有与年龄相关的变化。我们在鹦鹉(Melopsittacusundulatus)中检验了这一假设,一种群居的小型鹦鹉,成年人在响应组成员的变化时聚集在共享呼叫类型上。我们形成了4个以前不熟悉的男性属于同一年龄段的新群体,“年轻成年人”(6个月-1年)或“年长成年人”(≥3年),然后在20天的学习期内收集录音以评估声乐学习能力。在行为记录之后,对收集的神经组织进行免疫组织化学,以测量鹦鹉发声学习中心的FoxP2蛋白表达,内侧纹状体的大细胞核(MMSt),和它相邻的纹状体。
    结果:尽管与年轻人相比,老年人在MMST中表现出较低的声音多样性(即曲目大小)和较高的FoxP2绝对水平,我们发现在这两个年龄组中FoxP2的持续下调和等效的声带可塑性和声带趋同.未检测到声乐学习措施的个体差异与FoxP2表达之间的关系。
    结论:我们发现神经证据支持鹦鹉持续的声乐学习,在该物种的开放式学习计划中表明对衰老的抵抗力。FoxP2表达与声乐学习表现的个体差异之间缺乏显着关系,这表明其他神经遗传学机制也可以调节这种复杂的行为。
    BACKGROUND: Most vocal learning species exhibit an early critical period during which their vocal control neural circuitry facilitates the acquisition of new vocalizations. Some taxa, most notably humans and parrots, retain some degree of neurobehavioral plasticity throughout adulthood, but both the extent of this plasticity and the neurogenetic mechanisms underlying it remain unclear. Differential expression of the transcription factor FoxP2 in both songbird and parrot vocal control nuclei has been identified previously as a key pattern facilitating vocal learning. We hypothesize that the resilience of vocal learning to cognitive decline in open-ended learners will be reflected in an absence of age-related changes in neural FoxP2 expression. We tested this hypothesis in the budgerigar (Melopsittacus undulatus), a small gregarious parrot in which adults converge on shared call types in response to shifts in group membership. We formed novel flocks of 4 previously unfamiliar males belonging to the same age class, either \"young adult\" (6 mo - 1 year) or \"older adult\" (≥ 3 year), and then collected audio-recordings over a 20-day learning period to assess vocal learning ability. Following behavioral recording, immunohistochemistry was performed on collected neural tissue to measure FoxP2 protein expression in a parrot vocal learning center, the magnocellular nucleus of the medial striatum (MMSt), and its adjacent striatum.
    RESULTS: Although older adults show lower vocal diversity (i.e. repertoire size) and higher absolute levels of FoxP2 in the MMSt than young adults, we find similarly persistent downregulation of FoxP2 and equivalent vocal plasticity and vocal convergence in the two age cohorts. No relationship between individual variation in vocal learning measures and FoxP2 expression was detected.
    CONCLUSIONS: We find neural evidence to support persistent vocal learning in the budgerigar, suggesting resilience to aging in the open-ended learning program of this species. The lack of a significant relationship between FoxP2 expression and individual variability in vocal learning performance suggests that other neurogenetic mechanisms could also regulate this complex behavior.
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  • 文章类型: Journal Article
    鸣鸟的声乐掌握令人印象深刻,但这在多大程度上是实践的结果?他们能吗,基于与已知目标的不匹配,planthenecessarychangestorecoverthetargetinapractice-freemannersinging?Inadultzebrainches,我们将歌曲音节的音高从其稳定(基线)变体中移开,然后我们撤回增援,随后通过静音或震耳欲聋来剥夺他们的歌唱经验。在这个被剥夺的国家,鸟不恢复他们的基线歌曲。然而,他们将他们的歌曲回复到目标,大约是他们最近练习的1个标准差,在后者期间提供感觉反馈,表明音调与目标不匹配。因此,有针对性的声带可塑性不需要立即的感官体验,表明斑马雀能够进行目标导向的声音规划。
    Songbirds\' vocal mastery is impressive, but to what extent is it a result of practice? Can they, based on experienced mismatch with a known target, plan the necessary changes to recover the target in a practice-free manner without intermittently singing? In adult zebra finches, we drive the pitch of a song syllable away from its stable (baseline) variant acquired from a tutor, then we withdraw reinforcement and subsequently deprive them of singing experience by muting or deafening. In this deprived state, birds do not recover their baseline song. However, they revert their songs toward the target by about 1 standard deviation of their recent practice, provided the sensory feedback during the latter signaled a pitch mismatch with the target. Thus, targeted vocal plasticity does not require immediate sensory experience, showing that zebra finches are capable of goal-directed vocal planning.
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  • 文章类型: Journal Article
    发声通过传达有关发件人的个人或群体身份的信息来协调特定物种之间的社交互动。社会适应是声音学习的一种形式,通过使声音与特定声音融合或不同来表明社会亲和力。调查社会住宿是否与发件人的社会生活方式有关,我们调查了分散的灵长类动物在社会适应方面的性别差异,灰鼠狐猴(Microcebusmurinus),女性形成稳定的睡眠群体,而男性独自生活。我们使用了来自圈养繁殖群体的36个个体的482个颤音来比较具有遗传相关性的个体之间的声学差异,社交时间和体重。我们的结果表明,女性花在一起的时间越多,女性的颤音就越相似,独立于遗传关系,暗示声音收敛。相比之下,男性颤音受遗传因素的影响大于社会因素。然而,只关注社会化的男性,随着笼子伴侣的时间增加,男性的颤音出现了更大的分歧。因此,灰鼠狐猴显示出社会住宿的能力,女性聚集他们的颤音来表明与沉睡的团体伙伴的社交亲密关系,而男性不适应或分散他们的颤音来表明个人的独特性。本文是“声音的力量:揭示声音交流如何塑造群体动力学”主题的一部分。
    Vocalizations coordinate social interactions between conspecifics by conveying information concerning the individual or group identity of the sender. Social accommodation is a form of vocal learning where social affinity is signalled by converging or diverging vocalizations with those of conspecifics. To investigate whether social accommodation is linked to the social lifestyle of the sender, we investigated sex-specific differences in social accommodation in a dispersed living primate, the grey mouse lemur (Microcebus murinus), where females form stable sleeping groups whereas males live solitarily. We used 482 trill calls of 36 individuals from our captive breeding colony to compare acoustic dissimilarity between individuals with genetic relatedness, social contact time and body weight. Our results showed that female trills become more similar the more time females spend with each other, independent of genetic relationship, suggesting vocal convergence. In contrast, male trills were affected more by genetic than social factors. However, focusing only on socialized males, increasing time as cage partners caused greater divergence in males\' trills. Thus, grey mouse lemurs show the capacity for social accommodation, with females converging their trills to signal social closeness to sleeping group partners, whereas males do not adapt or diverge their trills to signal individual distinctiveness. This article is part of the theme issue \'The power of sound: unravelling how acoustic communication shapes group dynamics\'.
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  • 文章类型: Journal Article
    转录因子FoxP1的罕见破坏涉及人类神经发育障碍,其特征是自闭症和/或智力障碍,在言语和语言能力方面存在突出问题。该转录因子的禽类直系同源物在进化上是保守的,并在鸣禽大脑的特定区域高度表达。包括与声乐制作学习和听觉感知相关的领域。这里,我们调查了FoxP1对幼年和成年雌性斑马雀的歌曲辨别和听觉感知的可能贡献。他们在参与听觉刺激处理的两个大脑区域之一中接受了FoxP1的慢病毒敲除,HVC(专有名称)或CMM(尾内侧中镓)。96名女性,训练分布在不同的实验组和对照组中,以有效的Go/Nogo范式区分两首刺激歌曲,随后使用一系列刺激进行测试。这使得可以评估他们对训练刺激的改变版本的识别和分类程度,以及局部FoxP1击倒是否影响了歌曲辨别和分类过程中不同特征的作用。尽管FoxP1表达通过敲除显著降低,既不区分刺激歌曲,也不对音高修改的歌曲进行分类,音节的顺序或反向播放受到影响。随后,我们分析了完整的数据集,以评估不同刺激操作对歌曲辨别中线索称重的影响。我们的发现表明,斑马雀依赖于多个参数来区分歌曲,但是在频谱参数和音节排序方面具有相对更重要的作用,可以作为歌曲辨别的线索。
    Rare disruptions of the transcription factor FOXP1 are implicated in a human neurodevelopmental disorder characterized by autism and/or intellectual disability with prominent problems in speech and language abilities. Avian orthologues of this transcription factor are evolutionarily conserved and highly expressed in specific regions of songbird brains, including areas associated with vocal production learning and auditory perception. Here, we investigated possible contributions of FoxP1 to song discrimination and auditory perception in juvenile and adult female zebra finches. They received lentiviral knockdowns of FoxP1 in one of two brain areas involved in auditory stimulus processing, HVC (proper name) or CMM (caudomedial mesopallium). Ninety-six females, distributed over different experimental and control groups were trained to discriminate between two stimulus songs in an operant Go/Nogo paradigm and subsequently tested with an array of stimuli. This made it possible to assess how well they recognized and categorized altered versions of training stimuli and whether localized FoxP1 knockdowns affected the role of different features during discrimination and categorization of song. Although FoxP1 expression was significantly reduced by the knockdowns, neither discrimination of the stimulus songs nor categorization of songs modified in pitch, sequential order of syllables or by reversed playback were affected. Subsequently, we analyzed the full dataset to assess the impact of the different stimulus manipulations for cue weighing in song discrimination. Our findings show that zebra finches rely on multiple parameters for song discrimination, but with relatively more prominent roles for spectral parameters and syllable sequencing as cues for song discrimination.NEW & NOTEWORTHY In humans, mutations of the transcription factor FoxP1 are implicated in speech and language problems. In songbirds, FoxP1 has been linked to male song learning and female preference strength. We found that FoxP1 knockdowns in female HVC and caudomedial mesopallium (CMM) did not alter song discrimination or categorization based on spectral and temporal information. However, this large dataset allowed to validate different cue weights for spectral over temporal information for song recognition.
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  • 文章类型: Journal Article
    由于不同的进化过程,动物歌曲可以在种群内部和种群之间发生变化。当这些过程包括文化传播时,从特定物种中获得信息或行为的社会学习,歌曲可以经历快速发展,因为文化新奇比基因突变更频繁地出现。了解这些歌曲在大的时间和空间尺度上的变化可以提供对模式的见解,歌曲进化的驱动因素和限制,最终可以告知物种适应快速变化的声学环境的能力。这里,我们分析了北大西洋中部和东部二十年来记录的长鳍鲸(Balaenopteraphysalus)歌曲的变化。我们记录了仅在四个歌唱季节中快速替换歌曲INI(音符间隔),与混合歌曲(与两个INI)共同发生,以及在过渡期间不同歌曲INI的发生具有明显的地理梯度。我们还发现了十多年来INI和音符频率的逐渐变化,长须鲸采用了歌曲变化。这些结果为长须鲸的声音学习提供了证据,并揭示了歌曲进化的模式,这引起了对该物种歌曲变异极限的质疑。
    Animal songs can change within and between populations as the result of different evolutionary processes. When these processes include cultural transmission, the social learning of information or behaviours from conspecifics, songs can undergo rapid evolutions because cultural novelties can emerge more frequently than genetic mutations. Understanding these song variations over large temporal and spatial scales can provide insights into the patterns, drivers and limits of song evolution that can ultimately inform on the species\' capacity to adapt to rapidly changing acoustic environments. Here, we analysed changes in fin whale (Balaenoptera physalus) songs recorded over two decades across the central and eastern North Atlantic Ocean. We document a rapid replacement of song INIs (inter-note intervals) over just four singing seasons, that co-occurred with hybrid songs (with both INIs), and a clear geographic gradient in the occurrence of different song INIs during the transition period. We also found gradual changes in INIs and note frequencies over more than a decade with fin whales adopting song changes. These results provide evidence of vocal learning in fin whales and reveal patterns of song evolution that raise questions on the limits of song variation in this species.
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  • 文章类型: Journal Article
    在这篇文章中,我正在使用我在催产素方面的工作作为指南针来完成我的科学和个人旅程。我讲述了这些年来我的科学问题是如何形成的,以及我如何通过不同领域的镜头研究它们,从语言学和神经科学到各种脊椎动物物种的比较和人口基因组学。我解释了我的进化发现和关于催产素-血管生成素配体和受体家族中通用基因命名法的提议如何影响相关领域,以及我对鸣禽催产素和血管催产素系统的研究,人类和非人类灵长类动物使我现在开始测试鼻内催产素作为言语缺陷的候选治疗方法。我还讨论了我关于舞蹈神经生物学的项目,以及催产素在平行学习演讲和舞蹈中的位置。最后,我简要交流了我作为一名女性和国际科学和学术界学者所面临的挑战,以及我个人克服它们的方法。
    In this article, I am going through my scientific and personal journey using my work on oxytocin as a compass. I recount how my scientific questions were shaped over the years, and how I studied them through the lens of different fields ranging from linguistics and neuroscience to comparative and population genomics in a wide range of vertebrate species. I explain how my evolutionary findings and proposal for a universal gene nomenclature in the oxytocin-vasotocin ligand and receptor families have impacted relevant fields, and how my studies in the oxytocin and vasotocin system in songbirds, humans and non-human primates have led me to now be testing intranasal oxytocin as a candidate treatment for speech deficits. I also discuss my projects on the neurobiology of dance and where oxytocin fits in the picture of studying speech and dance in parallel. Lastly, I briefly communicate the challenges I have been facing as a woman and an international scholar in science and academia, and my personal ways to overcome them.
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  • 文章类型: Journal Article
    鹦鹉具有巨大的人声模仿能力,并产生独特的人声签名。像鸣鸟一样,鹦鹉具有有核的神经歌曲系统,具有不同的前(AFP)和后前脑通路(PFP)。为了测试鹦鹉和鸣鸟的歌曲系统,这在5000万年前发生了分歧,有类似的职能组织,我们首先建立了与神经科学兼容的呼叫和响应行为范式,以引起鹦鹉(Melopsittacusundulatus)中学习的联系呼叫。使用基于变分自动编码器的机器学习方法,我们表明,附属群体内的联系呼叫会收敛,但个人保持独特的声学特征,或者声音签名,即使在呼叫收敛之后。接下来,我们暂时使AFP的输出失活,以测试是否可以单独通过PFP产生已学习的发声。就像鸣鸟一样,AFP失活对发声有立竿见影的影响,与运动前的作用一致。但与鸣鸟相反,孤立的PFP足以产生定型和声学上正常的发声,鹦鹉PFP的隔离导致呼叫声学结构的退化,刻板印象,和个人的独特性。因此,AFP的贡献和孤立的PFP产生学习发声的能力在鸣禽和鹦鹉之间大相径庭,可能是由他们独特的行为生态学和神经连通性驱动的。
    Parrots have enormous vocal imitation capacities and produce individually unique vocal signatures. Like songbirds, parrots have a nucleated neural song system with distinct anterior (AFP) and posterior forebrain pathways (PFP). To test if song systems of parrots and songbirds, which diverged over 50 million years ago, have a similar functional organization, we first established a neuroscience-compatible call-and-response behavioral paradigm to elicit learned contact calls in budgerigars (Melopsittacus undulatus). Using variational autoencoder-based machine learning methods, we show that contact calls within affiliated groups converge but that individuals maintain unique acoustic features, or vocal signatures, even after call convergence. Next, we transiently inactivated the outputs of AFP to test if learned vocalizations can be produced by the PFP alone. As in songbirds, AFP inactivation had an immediate effect on vocalizations, consistent with a premotor role. But in contrast to songbirds, where the isolated PFP is sufficient to produce stereotyped and acoustically normal vocalizations, isolation of the budgerigar PFP caused a degradation of call acoustic structure, stereotypy, and individual uniqueness. Thus, the contribution of AFP and the capacity of isolated PFP to produce learned vocalizations have diverged substantially between songbirds and parrots, likely driven by their distinct behavioral ecology and neural connectivity.
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  • 文章类型: Journal Article
    声乐制作学习和节拍感知和同步(BPS)有一些共同的特点,这使声乐学习和节奏同步假设(VLH)对节奏同步能力的演变做出了合理的解释。然而,即使在声乐学习者中,很少看到非人类动物向人类音乐展示BPS。因此,本文的第一个目的是提出一些可能的原因,为什么我们没有在鹦鹉中看到BPS,一个优秀的声乐学习物种,同时提出我自己的一些发现。本文的第二个目标是提出一个无缝的桥梁,以连接语音学习和运动中的BPS能力。为此,我提出我自己的发现,其中花花公子自发地与人类音乐的旋律同步唱歌。这种行为可以被认为是BPS的声音版本。因此,它可以在这两种能力之间建立联系。本文同意除声乐学习系统以外的其他机制可能启用BPS的可能性,与VLH的最初想法相反。然而,将声乐学习的能力和BPS的能力联系起来仍然是合理的。至少,声乐学习能力可能有助于BPS的进化。从这些论点中,本文还提出了一种方案,其中包括同步发声作为BPS发展的驱动力和音乐制作的能力。
    Vocal production learning and beat perception and synchronization (BPS) share some common characteristics, which makes the vocal learning and rhythmic synchronization hypothesis (VLH) a reasonable explanation for the evolution of the capability for rhythmic synchronization. However, even in vocal learners, it is rare to see non-human animals demonstrate BPS to human music. Therefore, the first objective of this article is to propose some possible reasons why we do not see BPS in budgerigars, an excellent vocal learning species, while presenting some of my own findings. The second objective of this article is to propose a seamless bridge to connect the capability for vocal learning and BPS in locomotion. For this purpose, I present my own findings, wherein cockatiels spontaneously sang in synchrony with a melody of human music. This behavior can be considered a vocal version of BPS. Therefore, it can establish a connection between these two capabilities. This article agrees with the possibility that some mechanisms other than the vocal learning system may enable BPS, contrary to the original idea of VLH. Nevertheless, it is still reasonable to connect the capability for vocal learning and that for BPS. At the very least, the capability for vocal learning may contribute to the evolution of BPS. From these arguments, this article also proposes a scenario which includes vocalizing in synchrony as a driving force for the evolution of BPS and the capability for music production.
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  • 文章类型: Editorial
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