Song System Research Articles

Parallel executive pallio-motor loops in the pigeon brain.

A core component of the avian pallial cognitive network is the multimodal nidopallium caudolaterale (NCL) that is considered to be analogous to the mammalian prefrontal cortex (PFC). The NCL plays a key role in a multitude of executive tasks such as working memory, decision-making during navigation, and extinction learning in complex learning environments. Like the PFC, the NCL is positioned at the transition from ascending sensory to descending motor systems. For the latter, it sends descending premotor projections to the intermediate arcopallium (AI) and the medial striatum (MSt). To gain detailed insight into the organization of these projections, we conducted several retrograde and anterograde tracing experiments. First, we tested whether NCL neurons projecting to AI (NCLarco neurons) and MSt (NCLMSt neurons) are constituted by a single neuronal population with bifurcating neurons, or whether they form two distinct populations. Here, we found two distinct projection patterns to both target areas that were associated with different morphologies. Second, we revealed a weak topographic projection toward the medial and lateral striatum and a strong topographic projection toward AI with clearly distinguishable sensory termination fields. Third, we investigated the relationship between the descending NCL pathways to the arcopallium with those from the hyperpallium apicale, which harbors a second major descending pathway of the avian pallium. We embed our findings within a system of parallel pallio-motor loops that carry information from separate sensory modalities to different subpallial systems. Our results also provide insights into the evolution of the avian motor system from which, possibly, the song system has emerged.

Hemispheric dominance in HVC is experience-dependent in juvenile male zebra finches

Juvenile male zebra finches (Taeniopygia guttata) must be exposed to an adult tutor during a sensitive period to develop normal adult song. The pre-motor nucleus HVC (acronym used as a proper name), plays a critical role in song learning and production (cf. Broca’s area in humans). In the human brain, left-side hemispheric dominance in some language regions is positively correlated with proficiency in linguistic skills. However, it is unclear whether this pattern depends upon language learning, develops with normal maturation of the brain, or is the result of pre-existing functional asymmetries. In juvenile zebra finches, even though both left and right HVC contribute to song production, baseline molecular activity in HVC is left-dominant. To test if HVC exhibits hemispheric dominance prior to song learning, we raised juvenile males in isolation from adult song and measured neuronal activity in the left and right HVC upon first exposure to an auditory stimulus. Activity in the HVC was measured using the immediate early gene (IEG) zenk (acronym for zif-268, egr-1, NGFI-a, and krox-24) as a marker for neuronal activity. We found that neuronal activity in the HVC of juvenile male zebra finches is not lateralized when raised in the absence of adult song, while normally-reared juvenile birds are left-dominant. These findings show that there is no pre-existing asymmetry in the HVC prior to song exposure, suggesting that lateralization of the song system depends on learning through early exposure to adult song and subsequent song-imitation practice.

Retracted: Research on the Architecture of Digital Song and Dance Costume Design System Based on Intelligent Deep Learning Algorithm

Retracted: Research on the Architecture of Digital Song and Dance Costume Design System Based on Intelligent Deep Learning Algorithm

A neural circuit for vocal production responds to viscerosensory input in the songbird.

Motor performance is monitored continuously by specialized brain circuits and used adaptively to modify behavior on a moment-to-moment basis and over longer time periods. During vocal behaviors, such as singing in songbirds, internal evaluation of motor performance relies on sensory input from the auditory and vocal-respiratory systems. Sensory input from the auditory system to the motor system, often referred to as auditory feedback, has been well studied in singing zebra finches (Taeniopygia guttata), but little is known about how and where nonauditory sensory feedback is evaluated. Here we show that brief perturbations in air sac pressure cause short-latency neural responses in the higher-order song control nucleus HVC (used as proper name), an area necessary for song learning and song production. Air sacs were briefly pressurized through a cannula in anesthetized or sedated adult male zebra finches, and neural responses were recorded in both nucleus parambigualis (PAm), a brainstem inspiratory center, and HVC, a cortical premotor nucleus. These findings show that song control nuclei in the avian song system are sensitive to perturbations directly targeted to vocal-respiratory, or viscerosensory, afferents and support a role for multimodal sensory feedback integration in modifying and controlling vocal control circuits.NEW & NOTEWORTHY This study presents the first evidence of sensory input from the vocal-respiratory periphery directly activating neurons in a motor circuit for vocal production in songbirds. It was previously thought that this circuit relies exclusively on sensory input from the auditory system, but we provide groundbreaking evidence for nonauditory sensory input reaching the higher-order premotor nucleus HVC, expanding our understanding of what sensory feedback may be available for vocal control.

Anterior forebrain pathway in parrots is necessary for producing learned vocalizations with individual signatures

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.

Songbird mesostriatal dopamine pathways are spatially segregated before the onset of vocal learning.

Diverse dopamine (DA) pathways send distinct reinforcement signals to different striatal regions. In adult songbirds, a DA pathway from the ventral tegmental area (VTA) to Area X, the striatal nucleus of the song system, carries singing-related performance error signals important for learning. Meanwhile, a parallel DA pathway to a medial striatal area (MST) arises from a distinct group of neighboring DA neurons that lack connectivity to song circuits and do not encode song error. To test if the structural and functional segregation of these two pathways depends on singing experience, we carried out anatomical studies early in development before the onset of song learning. We find that distinct VTA neurons project to either Area X or MST in juvenile birds before the onset of substantial vocal practice. Quantitative comparisons of early juveniles (30-35 days post hatch), late juveniles (60-65 dph), and adult (>90 dph) brains revealed an outsized expansion of Area X-projecting neurons relative to MST-projecting neurons in VTA over development. These results show that a mesostriatal DA system dedicated to social communication can exist and be spatially segregated before the onset of vocal practice and associated sensorimotor experience.

Uncoordinated sleep replay across hemispheres in the zebra finch

Bilaterally organized brain regions are often simultaneously active in both humans1,2,3 and animal models,4,5,6,7,8,9 but the extent to which the temporal progression of internally generated dynamics is coordinated across hemispheres and how this coordination changes with brain state remain poorly understood. To address these issues, we investigated the zebra finch courtship song (duration: 0.5-1.0 s), a highly stereotyped complex behavior10,11 produced by a set of bilaterally organized nuclei.12,13,14 Unilateral lesions to these structures can eliminate or degrade singing,13,15,16,17 indicating that both hemispheres are required for song production.18 Additionally, previous work demonstrated broadly coherent and symmetric bilateral premotor signals during song.9 To precisely track the temporal evolution of activity in each hemisphere, we recorded bilaterally in the song production pathway. We targeted the robust nucleus of the arcopallium (RA) in the zebra finch, where population activity reflects the moment-to-moment progression of the courtship song during awake vocalizations19,20,21,22,23,24 and sleep, where song-related network dynamics reemerge in "replay" events.24,25 We found that activity in the left and right RA is synchronized within a fraction of a millisecond throughout song. In stark contrast, the two hemispheres displayed largely independent replay activity during sleep, despite shared interhemispheric arousal levels. These findings demonstrate that the degree of bilateral coordination in the zebra finch song system is dynamically modulated by behavioral state.

Song system neuroanatomy, and immediate early gene expression in a finch species with extensive male and female song.

Birdsong is a relatively well-studied behavior, both due to its importance as a model for vocal production learning and as an intriguing complex social behavior. Until the last few decades, work on birdsong focused almost exclusively on males. However, it is now widely accepted that female song not only exists, but is fairly common throughout the oscine passerines. Despite this, and the large number of researchers who have begun exploring female song in the field, researchers in the lab have been slow to adopt model species with female song. Studying female song in the lab is critical for our understanding of sex-specific factors in the physiology controlling this fascinating behavior. Additionally, as a model for vocal production learning in humans, understanding the mechanistic and neuroendocrine control of female song is clearly important. In this study, we examined the red-cheeked cordon bleu (RCCB), an Estrildid finch species with extensive female song. Specifically, we found that there were no significant sex differences in circulating levels of testosterone and progesterone, nor in song production rate. There were no significant differences in cell densities in the three nuclei of the song control system we examined. Additionally, the volume of the robust nucleus of the arcopallium was not significantly different and we report the smallest sex difference in HVC yet published in a songbird. Finally, we demonstrated similar levels of motor driven immediate early gene expression in both males and females after song production.

The effects of Wnt, BMP, and Notch signaling pathways on cell proliferation and neural differentiation in a song control nucleus (HVC) of Lonchura striata.

There is obvious sexual dimorphism in the song control system of songbirds. In the higher vocal center (HVC), cell proliferation and neuronal differentiation contribute to the net addition of neurons. However, the mechanism underlying these changes is unclear. Given that Wnt, Bmp, and Notch pathways are involved in cell proliferation and neuronal differentiation, no reports are available to study the role of the three pathways in the song control system. To address the issue, we studied cell proliferation in the ventricle zone overlying the developing HVC and neural differentiation within the HVC of Bengalese finches (Lonchura striata) at posthatching day 15 when HVC progenitor cells are generated on a large scale and differentiate into neurons, after Wnt and Bmp pathways were activated by using a pharmacological agonist (LiCl) or Bmp4, respectively, and the Notch pathway was inhibited by an inhibitor (N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester), DAPT). The results indicated that both cell proliferation and neural differentiation toward HVC neurons increased significantly after activation of the Wnt signaling pathway or inhibition of the Notch signaling pathway. Although cell proliferation was increased, neural differentiation was inhibited after treatment with Bmp4. There was obvious synergetic enhancement in the number of proliferating cells after the coregulation of two or three signaling pathways. In addition, synergetic enhancement was also found in the Wnt and Notch pathways in neural differentiation toward neurons within HVC. These results suggest that the three signaling pathways are involved in cell proliferation and neural differentiation of HVC.

The National Collective Management Institute (LMKN) as an Appraiser of Economic Value in the Binding of Music and Song Copyrights as Objects of Fiduciary Guarantees

One of the creations protected by copyright is songs and/or music. Legal protection for them is a necessity in order to create a conducive climate to the development of music industry in Indonesia. In order to facilitate song and/or music writers with better control over their works, they can appoint representatives to regulate the management of their royalty, in this case LMKN (The National Collective Management Institute). Law number 28 of 2014 concerning copyrights regulates that copyright can be used as an object of fiduciary guarantee. However, its use is still hampered by the fact that the economic value of music and song copyrights cannot be determined easily, especially if it is used for fiduciary guarantee. Most banks do not accept it for loan collateral. The objective of this research was to analyze the roles of LMKN as the determiner of the economic values of song and music copyrights for the purpose of binding them for objects of fiduciary guarantee. This normative study uses statutory approach, conceptual approach, analytical approach, and futuristic approach. The determination of the economic values of song and music copyrights can be done by LMKN, a one- stop service institution that collects the royalty of music and song in Indonesia. The National Collective Management Institute (hereinafter briefly referred to as LMKN) as an institution that can provide an appraiser of the economic value of Music and Song Copyrights. The use of SILM (a music and song information system and data center) can help the estimation of music and song’s economic value based on the royalty accepted by the art’s creator or copyright holder. The amount of royalty can determine the binding of music and song copyrights for the objects of fiduciary guarantee.

Divergent projections from locus coeruleus to the corticobasal ganglia system and ventral tegmental area of the adult male zebra finch.

The locus coeruleus (LC) is a small noradrenergic brainstem nucleus that plays a central role in regulating arousal, attention, and performance. In the mammalian brain, individual LC neurons make divergent axonal projections to different brain regions, which are distinguished in part by which noradrenaline (NA) receptor subtypes they express. Here, we sought to determine whether similar organizational features characterize LC projections to corticobasal ganglia (CBG) circuitry in the zebra finch song system, with a focus on the basal ganglia nucleus Area X, the thalamic nucleus DLM, as well as the cortical nuclei HVC, LMAN, and RA. Single and dual retrograde tracer injections reveal that single LC-NA neurons make divergent projections to LMAN and Area X, as well as to the dopaminergic VTA/SNc complex that innervates this CBG circuit. Moreover, in situ hybridization revealed that differential expression of mRNA encoding α2A and α2C adrenoreceptors distinguishes LC-recipient CBG song nuclei. Therefore, LC-NA signaling in the zebra finch CBG circuit employs a similar strategy as in mammals, which could allow a relatively small number of LC neurons to exert widespread yet distinct effects across multiple brain regions.

Neural correlates of cognitively controlled vocalizations in a corvid songbird

The neuronal basis of the songbird's song system is well understood. However, little is known about the neuronal correlates of the executive control of songbird vocalizations. Here, we record single-unit activity from the pallial endbrain region "nidopallium caudolaterale" (NCL) of crows that vocalize to the presentation of a visual go-cue but refrain from vocalizing during trials without a go-cue. We find that the preparatory activity of single vocalization-correlated neurons, but also of the entire population of NCL neurons, before vocal onset predicts whether or not the crows will produce an instructed vocalization. Fluctuations in baseline neuronal activity prior to the go-cue influence the premotor activity of such vocalization-correlated neurons and seemingly bias the crows' decision to vocalize. Neuronal response modulation significantly differs between volitional and task-unrelated vocalizations. This suggests that the NCL can take control over the vocal motor network during the production of volitional vocalizations in a corvid songbird.

Local and range-wide distribution of song types suggest Ovenbirds (Seiurus aurocapilla) have song neighborhoods but not macro-dialects

The song systems of oscine passerines (songbirds) are complex and diverse. Because songs are used for both mate attraction and territory defense and are therefore important signals for survival and reproduction, comprehensive knowledge of within and among species song structure and distribution is informative for understanding the evolution of song repertoires and vocal behaviour. In this study, we explored variation in the song structure of the Ovenbird (Seiurus aurocapilla), a widespread warbler (Family Parulidae) found in North American forests. We analyzed recordings from the 2021 breeding season to assess song type variation at a local (n ​= ​158 birds; Sault Ste. Marie, ON) and breeding range scale (n ​= ​512 birds; eBird). We characterized the local song types and tested whether Ovenbirds share song types with their neighbors more often than expected by chance. We then characterized song types of Ovenbirds across the breeding range to determine whether any geographic pattern of song clustering exists (i.e., macro-dialects). We found 10 distinct song types and some evidence for song type clustering at the local study site (i.e., song neighborhoods). We found 7 of those 10 song types throughout the breeding range and identified an additional 24 types that were not recorded in our local population. We found no evidence for song dialects across the Ovenbird breeding range. This study contributes to our understanding of Ovenbird song while simultaneously adding to our understanding of geographic structuring of warbler repertoires. Our work contributes to delineating a more comprehensive understanding of factors affecting dialect development for this diverse group of songbirds.

Discrimination in Silapathikaram

Silapathikaram is considered the best epic in Tamil literature. This epic is about the three kings who has been called Moovaendars and the three modes of entertainment such as literature, music and drama. The ancient people’s life style, social structure, standard of living of the people, caste discrimination, and folk song system have been highlighted in this text. Varuna (division of people based on caste) plays a major role in people's lives. Author IIangovadigal points out that this situation came from Silapathikaram period. The people are divided into four categories as priests, kings, merchants and servants. Though the text speaks well about merchants, it gave importance to priests. The protagonist Kovalan is highlighted in the text through the priests. Through this, we can feel their primacy in the society. Even though the kings and servants have material possessions, they are next to the Antananars (priests). In Silapathikaram, you can see a society where caste discrimination is established. It is not known whether this was the case at that time or whether the Ilangovadigal was trying to record such racial discrimination. Thus, the purpose of this article is to highlight the caste discrimination in Silapathikaram

Forebrain nuclei linked to woodpecker territorial drum displays mirror those that enable vocal learning in songbirds.

Vocal learning is thought to have evolved in 3 orders of birds (songbirds, parrots, and hummingbirds), with each showing similar brain regions that have comparable gene expression specializations relative to the surrounding forebrain motor circuitry. Here, we searched for signatures of these same gene expression specializations in previously uncharacterized brains of 7 assumed vocal non-learning bird lineages across the early branches of the avian family tree. Our findings using a conserved marker for the song system found little evidence of specializations in these taxa, except for woodpeckers. Instead, woodpeckers possessed forebrain regions that were anatomically similar to the pallial song nuclei of vocal learning birds. Field studies of free-living downy woodpeckers revealed that these brain nuclei showed increased expression of immediate early genes (IEGs) when males produce their iconic drum displays, the elaborate bill-hammering behavior that individuals use to compete for territories, much like birdsong. However, these specialized areas did not show increased IEG expression with vocalization or flight. We further confirmed that other woodpecker species contain these brain nuclei, suggesting that these brain regions are a common feature of the woodpecker brain. We therefore hypothesize that ancient forebrain nuclei for refined motor control may have given rise to not only the song control systems of vocal learning birds, but also the drumming system of woodpeckers.

A brain for all seasons: An in vivo MRI perspective on songbirds.

Seasonality in songbirds includes not only reproduction but also seasonal changes in singing behavior and its neural substrate, the song control system (SCS). Prior research mainly focused on the role of sex steroids on this seasonal SCS neuroplasticity in males. In this review, we summarize the advances made in the field of seasonal neuroplasticity by applying in vivo magnetic resonance imaging (MRI) in male and female starlings, analyzing the entire brain, monitoring birds longitudinally and determining the neuronal correlates of seasonal variations in plasma hormone levels and song behavior. The first MRI studies in songbirds used manganese enhanced MRI to visualize the SCS in a living bird and validated previously described brain volume changes related to different seasons and testosterone. MRI studies with testosterone implantation established how the consequential boost in singing was correlated to structural changes in the SCS, indicating activity-induced neuroplasticity as song proficiency increased. Next, diffusion tensor MRI explored seasonal neuroplasticity in the entire brain, focusing on networks beyond the SCS, revealing that other sensory systems and even the cerebellum, which is important for the integration of sensory perception and song behavior, experience neuroplasticity starting in the photosensitive period. Functional MRI showed that olfactory, and auditory processing was modulated by the seasons. The convergence of seasonal variations in so many sensory and sensorimotor systems resembles multisensory neuroplasticity during the critical period early in life. This sheds new light on seasonal songbirds as a model for unlocking the brain by recreating seasonally the permissive circumstances for heightened neuroplasticity.

Sex differences in seasonal brain plasticity and the neuroendocrine regulation of vocal behavior in songbirds

Birdsong is controlled in part by a discrete network of interconnected brain nuclei regulated in turn by steroid hormones and environmental stimuli. This complex interaction results in neural changes that occur seasonally as the environment varies (e.g., photoperiod, food/water availability, etc.). Variation in environment, vocal behavior, and neuroendocrine control has been primarily studied in male songbirds in both laboratory studies of captive birds and field studies of wild caught birds. The bias toward studying seasonality in the neuroendocrine regulation of song in male birds comes from a historic focus on sexually selected male behaviors. In fact, given that male song is often loud and accompanied by somewhat extravagant courtship behaviors, female song has long been overlooked. To compound this bias, the primary model songbird species for studies in the lab, zebra finches (Taeniopygia guttata) and canaries (Serinus canaria), exhibit little or no female song. Therefore, understanding the degree of variation and neuroendocrine control of seasonality in female songbirds is a major gap in our knowledge. In this review, we discuss the importance of studying sex differences in seasonal plasticity and the song control system. Specifically, we discuss sex differences in 1) the neuroanatomy of the song control system, 2) the distribution of receptors for androgens and estrogens and 3) the seasonal neuroplasticity of the hypothalamo-pituitary-gonadal axis as well as in the neural and cellular mechanisms mediating song system changes. We also discuss how these neuroendocrine mechanisms drive sex differences in seasonal behavior. Finally, we highlight specific gaps in our knowledge and suggest experiments critical for filling these gaps.

Induction of an immortalized songbird cell line allows for gene characterization and knockout by CRISPR-Cas9

The zebra finch is one of the most commonly studied songbirds in biology, particularly in genomics, neuroscience and vocal communication. However, this species lacks a robust cell line for molecular biology research and reagent optimization. We generated a cell line, designated CFS414, from zebra finch embryonic fibroblasts using the SV40 large and small T antigens. This cell line demonstrates an improvement over previous songbird cell lines through continuous and density-independent growth, allowing for indefinite culture and monoclonal line derivation. Cytogenetic, genomic, and transcriptomic profiling established the provenance of this cell line and identified the expression of genes relevant to ongoing songbird research. Using this cell line, we disrupted endogenous gene sequences using S.aureus Cas9 and confirmed a stress-dependent localization response of a song system specialized gene, SAP30L. The utility of CFS414 cells enhances the comprehensive molecular potential of the zebra finch and validates cell immortalization strategies in a songbird species.

Research on the Architecture of Digital Song and Dance Costume Design System Based on Intelligent Deep Learning Algorithm

In order to improve the intelligence and reliability of song and dance costume design, this article combines the intelligent deep learning algorithm to improve the digital costume design process. In order to solve the problem of minimizing the energy of image deformation, this paper proposes to transform it into a sparse matrix to solve the problem, which is solved by constructing a coefficient matrix and multiple iterations. After the algorithm is improved, this article combines the digital intelligence method to construct the digital song and dance costume design system structure based on the intelligent deep learning algorithm and analyzes its work flow. Finally, this paper verifies the method in this paper through feasibility analysis and system performance test. From the experimental results, it can be seen that the digital song and dance costume design system based on the intelligent deep learning algorithm proposed in this article has a good effect in the song and dance costume design.

Ancient Beliefs and Traditional Singing of Ukrainians: Gender Differentiation

Gender differentiation in the Ukrainian song tradition is most consistently manifested in the ritual system, and partly – in the epic tradition and lyrical song tradition. The primary attention in the article is paid to the ritual sphere, first of all to the calendar cycle. The paper highlights in detail the traditional distribution of functions between men and women in pre-Christian rituals, during which, according to ancient ideas and beliefs, there was some contact between «that» (sacred) and «this» («profane») worlds.
 Males, or more precisely, boys who were members of the so-called «parubotchi gromady» (young men communities) took an active part in the rites only once a year – at the beginning of the calendar-time cycle that is, in winter (in ancient times this happened in the spring) during the rituals of the yards circumambulation. The main purpose of these actions was to wish good for each family from the dead ancestors for the coming year (verbal magic), and in return, the ancestors received gifts – sacrificial food from representatives of the living world to appease them for the next year.
 Women, as representatives of «this» world, maintained contact with otherworldly forces throughout the entire agrarian period from sowing to harvest, as well as in ceremonies associated with the birth of a child, a wedding, or escorting the deceased to the afterlife. In times of crisis in the development of nature and human life, they turned to their deceased ancestors for help. The magical instrument of this connection was the voice, which filled the ritual texts with specific ritual timbre-intonation.
 The gender distribution in other genres of Ukrainian traditional song is somewhat different. Thus, if in the epic songs the prerogative belongs to men, then the lyric song system is characterized by the joint and almost equal participation of men and women. However, it should be noted that the performers of social songs were predominantly men, and women sang family lyric songs. But the most common was a mixed lineup of singing groups. Even more this property is inherent in the late layer of lyrical song performance.
 The author also draws attention to the age aspect of the performance of ritual and non-ritual songs in the Ukrainian tradition