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Journal ArticleDOI

Frequency Modulation During Song in a Suboscine Does Not Require Vocal Muscles

01 May 2008-Journal of Neurophysiology (American Physiological Society)-Vol. 99, Iss: 5, pp 2383-2389

TL;DR: This work investigates sound production and control of sound frequency in the Great Kiskadee by recording air sac pressure and vocalizations during spontaneously generated song and assumes a nonlinear restitution force for the oscillating membrane folds in a two mass model of sound production to reproduce the frequency modulations of the observed vocalizations.

AbstractThe physiology of sound production in suboscines is poorly investigated. Suboscines are thought to develop song innately unlike the closely related oscines. Comparing phonatory mechanisms might therefore provide interesting insight into the evolution of vocal learning. Here we investigate sound production and control of sound frequency in the Great Kiskadee (Pitangus sulfuratus) by recording air sac pressure and vocalizations during spontaneously generated song. In all the songs and calls recorded, the modulations of the fundamental frequency are highly correlated to air sac pressure. To test whether this relationship reflects frequency control by changing respiratory activity or indicates synchronized vocal control, we denervated the syringeal muscles by bilateral resection of the tracheosyringeal nerve. After denervation, the strong correlation between fundamental frequency and air sac pressure patterns remained unchanged. A single linear regression relates sound frequency to air sac pressure in the intact and denervated birds. This surprising lack of control by syringeal muscles of frequency in Kiskadees, in strong contrast to songbirds, poses the question of how air sac pressure regulates sound frequency. To explore this question theoretically, we assume a nonlinear restitution force for the oscillating membrane folds in a two mass model of sound production. This nonlinear restitution force is essential to reproduce the frequency modulations of the observed vocalizations.

Topics: Vocal muscle (54%)

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Citations
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Journal ArticleDOI
Abstract: A growing number of studies ask whether and how bird songs vary between areas with low versus high levels of anthropogenic noise. Across numerous species, birds are seen to sing at higher frequencies in urban versus rural populations, presumably because of selection for higher-pitched songs in the face of low-frequency urban noise, or because birds can avoid masking directly by shifting to higher-frequency sounds (Fernandez-Juricic et al. 2005; Slabbekoorn & den Boer-Visser 2006; Nemeth & Brumm 2009; Gross et al. 2010; Potvin et al. 2010). In addition to changing song frequency, birds are also reported to respond to increased background noise by singing at higher amplitudes (Brumm & Zollinger 2011). Nightingales, Luscinia megarhynchos, for example, sing with a higher sound pressure level in areas with intense traffic noise as compared to quieter locations (Brumm 2004). While frequencyand amplitude-based responses to ambient noise are often considered independently, the twomight also vary in tandem

180 citations


Cites background from "Frequency Modulation During Song in..."

  • ...A strong correlation between subsyringeal pressure and vocalization frequencywas also found in a suboscine bird, the great kiskadee, Pitangus sulphuratus (Amador et al. 2008), providing further evidence that driving pressure and frequency are biomechanically linked....

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Journal ArticleDOI
TL;DR: The results show that the syringeal skeleton is optimized for low weight driven by physiological constraints on song production, and a cartilaginous structure suited to play a crucial role in the uncoupling of sound frequency and amplitude control, which permits a novel explanation of the evolutionary success of songbirds.
Abstract: Like human infants, songbirds learn their species-specific vocalizations through imitation learning. The birdsong system has emerged as a widely used experimental animal model for understanding the underlying neural mechanisms responsible for vocal production learning. However, how neural impulses are translated into the precise motor behavior of the complex vocal organ (syrinx) to create song is poorly understood. First and foremost, we lack a detailed understanding of syringeal morphology. To fill this gap we combined non-invasive (high-field magnetic resonance imaging and micro-computed tomography) and invasive techniques (histology and micro-dissection) to construct the annotated high-resolution three-dimensional dataset, or morphome, of the zebra finch (Taeniopygia guttata) syrinx. We identified and annotated syringeal cartilage, bone and musculature in situ in unprecedented detail. We provide interactive three-dimensional models that greatly improve the communication of complex morphological data and our understanding of syringeal function in general. Our results show that the syringeal skeleton is optimized for low weight driven by physiological constraints on song production. The present refinement of muscle organization and identity elucidates how apposed muscles actuate different syringeal elements. Our dataset allows for more precise predictions about muscle co-activation and synergies and has important implications for muscle activity and stimulation experiments. We also demonstrate how the syrinx can be stabilized during song to reduce mechanical noise and, as such, enhance repetitive execution of stereotypic motor patterns. In addition, we identify a cartilaginous structure suited to play a crucial role in the uncoupling of sound frequency and amplitude control, which permits a novel explanation of the evolutionary success of songbirds.

179 citations


Journal ArticleDOI
07 Mar 2013-Nature
TL;DR: The HVC precisely encodes vocal motor output through activity at the times of extreme points of movement trajectories, and it is proposed that the sequential activity of HVC neurons is used as a 'forward' model, representing the sequence of gestures in song to make predictions on expected behaviour and evaluate feedback.
Abstract: Quantitative biomechanical models can identify control parameters that are used during movements, and movement parameters that are encoded by premotor neurons. We fit a mathematical dynamical systems model including subsyringeal pressure, syringeal biomechanics and upper-vocal-tract filtering to the songs of zebra finches. This reduces the dimensionality of singing dynamics, described as trajectories (motor 'gestures') in a space of syringeal pressure and tension. Here we assess model performance by characterizing the auditory response 'replay' of song premotor HVC neurons to the presentation of song variants in sleeping birds, and by examining HVC activity in singing birds. HVC projection neurons were excited and interneurons were suppressed within a few milliseconds of the extreme time points of the gesture trajectories. Thus, the HVC precisely encodes vocal motor output through activity at the times of extreme points of movement trajectories. We propose that the sequential activity of HVC neurons is used as a 'forward' model, representing the sequence of gestures in song to make predictions on expected behaviour and evaluate feedback.

153 citations


Journal ArticleDOI
TL;DR: Comparison of patterns of song adjustment to noise in oscines and suboscines in Brazil and Mexico City suggests that song learning and/or song plasticity allows adaptation to new habitats and that this selective advantage may be linked to the evolution ofsong learning and plasticity.
Abstract: Song learning has evolved within several avian groups. Although its evolutionary advantage is not clear, it has been proposed that song learning may be advantageous in allowing birds to adapt their songs to the local acoustic environment. To test this hypothesis, we analysed patterns of song adjustment to noisy environments and explored their possible link to song learning. Bird vocalizations can be masked by low-frequency noise, and birds respond to this by singing higher-pitched songs. Most reports of this strategy involve oscines, a group of birds with learning-based song variability, and it is doubtful whether species that lack song learning (e.g. suboscines) can adjust their songs to noisy environments. We address this question by comparing the degree of song adjustment to noise in a large sample of oscines (17 populations, 14 species) and suboscines (11 populations, 7 species), recorded in Brazil (Manaus, Brasilia and Curitiba) and Mexico City. We found a significantly stronger association between minimum song frequency and noise levels (effect size) in oscines than in suboscines, suggesting a tighter match in oscines between song transmission capacity and ambient acoustics. Suboscines may be more vulnerable to acoustic pollution than oscines and thus less capable of colonizing cities or acoustically novel habitats. Additionally, we found that species whose song frequency was more divergent between populations showed tighter noise-song frequency associations. Our results suggest that song learning and/or song plasticity allows adaptation to new habitats and that this selective advantage may be linked to the evolution of song learning and plasticity.

67 citations


Journal ArticleDOI
TL;DR: It is suggested that adjustments in song frequency and amplitude are largely independent and, thus, can be complementary rather than alternative vocal adjustments to noise.
Abstract: Songbirds often sing at higher frequency (pitch) in urban, noise-polluted areas, which reduces acoustic masking by low-frequency anthropogenic noise. Such frequency shifts, however, are less efficient at overcoming background noise than simply singing louder. Therefore, it was suggested that high-frequency singing might not be a functional adjustment to noise, but a physiological consequence of singing louder (also known as the Lombard effect). We tested for the first time the main tenet of this hypothesis, for birdsong whether increasing sound amplitude has a concomitant effect on song frequency, using a representative species with higher urban minimum frequency, the dark-eyed junco, Junco hyemalis. The frequency bandwidth of songs and syllables increased with amplitude, involving lower minimum frequency in louder songs and syllables. Therefore, louder singing does not explain the higher minimum frequency of urban dark-eyed juncos. Amplitude and peak frequency were weakly positively related across but not within songs, suggesting that increased frequency is not an obligatory outcome of singing louder. Instead, birds may adjust both amplitude and frequency in response to changing noise or motivation across songs. Our results suggest that adjustments in song frequency and amplitude are largely independent and, thus, can be complementary rather than alternative vocal adjustments to noise. We discuss oscine vocal physiology and details of the behaviour of urban birds, both of which we argue are consistent with the increased frequency of urban birdsong generally being a functional adjustment to noise, rather than a consequence of singing louder.

64 citations


References
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9,303 citations


Journal ArticleDOI
TL;DR: Human speech and birdsong have numerous parallels, with striking similarities in how sensory experience is internalized and used to shape vocal outputs, and how learning is enhanced during a critical period of development.
Abstract: Human speech and birdsong have numerous parallels. Both humans and songbirds learn their complex vocalizations early in life, exhibiting a strong dependence on hearing the adults they will imitate, as well as themselves as they practice, and a waning of this dependence as they mature. Innate predispositions for perceiving and learning the correct sounds exist in both groups, although more evidence of innate descriptions of species-specific signals exists in songbirds, where numerous species of vocal learners have been compared. Humans also share with songbirds an early phase of learning that is primarily perceptual, which then serves to guide later vocal production. Both humans and songbirds have evolved a complex hierarchy of specialized forebrain areas in which motor and auditory centers interact closely, and which control the lower vocal motor areas also found in nonlearners. In both these vocal learners, however, how auditory feedback of self is processed in these brain areas is surprisingly unclear. Finally, humans and songbirds have similar critical periods for vocal learning, with a much greater ability to learn early in life. In both groups, the capacity for late vocal learning may be decreased by the act of learning itself, as well as by biological factors such as the hormones of puberty. Although some features of birdsong and speech are clearly not analogous, such as the capacity of language for meaning, abstraction, and flexible associations, there are striking similarities in how sensory experience is internalized and used to shape vocal outputs, and how learning is enhanced during a critical period of development. Similar neural mechanisms may therefore be involved.

1,435 citations


Book
01 Jan 1974

1,404 citations


Journal ArticleDOI
TL;DR: It is concluded that Area X and LMAN contribute differently to song acquisition: the song variability that is typical of vocal development persists following early deafness or lesions of Area X but ends abruptly following removal of LMAN.
Abstract: Song production in song birds is controlled by an efferent pathway. Appended to this pathway is a “recursive loop” that is necessary for song acquisition but not for the production of learned song. Since zebra finches learn their song by imitating external models, we speculated that the importance of the recursive loop for learning might derive from its processing of auditory feedback during song acquisition. This hypothesis was tested by comparing the effects on song in birds deafened early in life and birds with early lesions in either of two nuclei--Area X and the lateral magnocellular nucleus of the anterior neostriatum (LMAN). These nuclei are part of the recursive loop. The three treatments affected song development differently, as reflected by various parameters of the adult song of these birds. Whereas LMAN lesions resulted in songs with monotonous repetitions of a single note complex, songs of Area X-lesioned birds consisted of rambling series of unusually long and variable notes. Furthermore, whereas song of LMAN lesioned birds stabilized early, song stability as seen in intact birds was never achieved in Area X-lesioned birds. Early deafness also resulted in poorly structured and unstable song. We conclude that Area X and LMAN contribute differently to song acquisition: the song variability that is typical of vocal development persists following early deafness or lesions of Area X but ends abruptly following removal of LMAN. Apparently, LMAN plays a crucial role in fostering the kinds of circuit plasticity necessary for learning.

891 citations


Journal ArticleDOI
25 May 1984-Science
TL;DR: Lesions in the magnocellular nucleus of the anterior neostriatum of passerine birds disrupted song development in juvenile male zebra finches but did not affect maintenance of stable song patterns by adult birds.
Abstract: The magnocellular nucleus of the anterior neostriatum is a forebrain nucleus of passerine birds that accumulates testosterone and makes monosynaptic connections with other telencephalic nuclei that control song production in adult birds. Lesions in the magnocellular nucleus disrupted song development in juvenile male zebra finches but did not affect maintenance of stable song patterns by adult birds. These results represent an instance in which lesions of a discrete brain region during only a restricted phase in the development of a learned behavior cause permanent impairment. Because cells of the magnocellular nucleus accumulate androgens these findings raise the possibility that this learning is mediated by hormones.

812 citations