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1.
The songs of Bengalese finches (Lonchura striata var. domestica) have complex syntax and provide an opportunity to investigate how complex sequential behaviour emerges via the evolutionary process. In this study, we suggest that a simple mechanism, i.e. many-to-one mapping from internal states onto syllables, may underlie the emergence of apparent complex syllable sequences that have higher order history dependencies. We analysed the songs of Bengalese finches and of their wild ancestor, the white-rumped munia (L. striata), whose songs are more stereotypical and simpler compared with those of Bengalese finches. The many-to-one mapping mechanism sufficiently accounted for the differences in the complexity of song syllable sequences of these two strains. 相似文献
2.
Songs of many songbird species consist of variable sequences of a finite number of syllables. A common approach for characterizing the syntax of these complex syllable sequences is to use transition probabilities between the syllables. This is equivalent to the Markov model, in which each syllable is associated with one state, and the transition probabilities between the states do not depend on the state transition history. Here we analyze the song syntax in Bengalese finch. We show that the Markov model fails to capture the statistical properties of the syllable sequences. Instead, a state transition model that accurately describes the statistics of the syllable sequences includes adaptation of the self-transition probabilities when states are revisited consecutively, and allows associations of more than one state to a given syllable. Such a model does not increase the model complexity significantly. Mathematically, the model is a partially observable Markov model with adaptation (POMMA). The success of the POMMA supports the branching chain network model of how syntax is controlled within the premotor song nucleus HVC, but also suggests that adaptation and many-to-one mapping from the syllable-encoding chain networks in HVC to syllables should be included in the network model. 相似文献
3.
Complex sequencing rules observed in birdsongs provide an opportunity to investigate the neural mechanism for generating complex sequential behaviors. To relate the findings from studying birdsongs to other sequential behaviors such as human speech and musical performance, it is crucial to characterize the statistical properties of the sequencing rules in birdsongs. However, the properties of the sequencing rules in birdsongs have not yet been fully addressed. In this study, we investigate the statistical properties of the complex birdsong of the Bengalese finch (Lonchura striata var. domestica). Based on manual-annotated syllable labeles, we first show that there are significant higher-order context dependencies in Bengalese finch songs, that is, which syllable appears next depends on more than one previous syllable. We then analyze acoustic features of the song and show that higher-order context dependencies can be explained using first-order hidden state transition dynamics with redundant hidden states. This model corresponds to hidden Markov models (HMMs), well known statistical models with a large range of application for time series modeling. The song annotation with these models with first-order hidden state dynamics agreed well with manual annotation, the score was comparable to that of a second-order HMM, and surpassed the zeroth-order model (the Gaussian mixture model; GMM), which does not use context information. Our results imply that the hierarchical representation with hidden state dynamics may underlie the neural implementation for generating complex behavioral sequences with higher-order dependencies. 相似文献
4.
Background
There has been plentiful evidence of kinesthetically induced rapid compensation for unanticipated perturbation in speech articulatory movements. However, the role of auditory information in stabilizing articulation has been little studied except for the control of voice fundamental frequency, voice amplitude and vowel formant frequencies. Although the influence of auditory information on the articulatory control process is evident in unintended speech errors caused by delayed auditory feedback, the direct and immediate effect of auditory alteration on the movements of articulators has not been clarified.Methodology/Principal Findings
This work examined whether temporal changes in the auditory feedback of bilabial plosives immediately affects the subsequent lip movement. We conducted experiments with an auditory feedback alteration system that enabled us to replace or block speech sounds in real time. Participants were asked to produce the syllable /pa/ repeatedly at a constant rate. During the repetition, normal auditory feedback was interrupted, and one of three pre-recorded syllables /pa/, /Φa/, or /pi/, spoken by the same participant, was presented once at a different timing from the anticipated production onset, while no feedback was presented for subsequent repetitions. Comparisons of the labial distance trajectories under altered and normal feedback conditions indicated that the movement quickened during the short period immediately after the alteration onset, when /pa/ was presented 50 ms before the expected timing. Such change was not significant under other feedback conditions we tested.Conclusions/Significance
The earlier articulation rapidly induced by the progressive auditory input suggests that a compensatory mechanism helps to maintain a constant speech rate by detecting errors between the internally predicted and actually provided auditory information associated with self movement. The timing- and context-dependent effects of feedback alteration suggest that the sensory error detection works in a temporally asymmetric window where acoustic features of the syllable to be produced may be coded. 相似文献5.
Background
We ordinarily perceive our voice sound as occurring simultaneously with vocal production, but the sense of simultaneity in vocalization can be easily interrupted by delayed auditory feedback (DAF). DAF causes normal people to have difficulty speaking fluently but helps people with stuttering to improve speech fluency. However, the underlying temporal mechanism for integrating the motor production of voice and the auditory perception of vocal sound remains unclear. In this study, we investigated the temporal tuning mechanism integrating vocal sensory and voice sounds under DAF with an adaptation technique.Methods and Findings
Participants produced a single voice sound repeatedly with specific delay times of DAF (0, 66, 133 ms) during three minutes to induce ‘Lag Adaptation’. They then judged the simultaneity between motor sensation and vocal sound given feedback. We found that lag adaptation induced a shift in simultaneity responses toward the adapted auditory delays. This indicates that the temporal tuning mechanism in vocalization can be temporally recalibrated after prolonged exposure to delayed vocal sounds. Furthermore, we found that the temporal recalibration in vocalization can be affected by averaging delay times in the adaptation phase.Conclusions
These findings suggest vocalization is finely tuned by the temporal recalibration mechanism, which acutely monitors the integration of temporal delays between motor sensation and vocal sound. 相似文献6.
Adult Bengalese finches generate a variable song that obeys a distinct and individual syntax. The syntax is gradually lost
over a period of days after deafening and is recovered when hearing is restored. We present a spiking neuronal network model
of the song syntax generation and its loss, based on the assumption that the syntax is stored in reafferent connections from
the auditory to the motor control area. Propagating synfire activity in the HVC codes for individual syllables of the song
and priming signals from the auditory network reduce the competition between syllables to allow only those transitions that
are permitted by the syntax. Both imprinting of song syntax within HVC and the interaction of the reafferent signal with an
efference copy of the motor command are sufficient to explain the gradual loss of syntax in the absence of auditory feedback.
The model also reproduces for the first time experimental findings on the influence of altered auditory feedback on the song
syntax generation, and predicts song- and species-specific low frequency components in the LFP. This study illustrates how
sequential compositionality following a defined syntax can be realized in networks of spiking neurons. 相似文献
7.
Accurate coordination of the sequencing and timing of motor gestures is important for the performance of complex and evolutionarily relevant behaviors. However, the degree to which motor sequencing and timing are related remains largely unknown. Birdsong is a communicative behavior that consists of discrete vocal motor elements (‘syllables’) that are sequenced and timed in a precise manner. To reveal the relationship between syllable sequencing and timing, we analyzed how variation in the probability of syllable transitions at branch points, nodes in song with variable sequencing across renditions, correlated with variation in the duration of silent gaps between syllable transitions (‘gap durations’) for adult Bengalese finch song. We observed a significant negative relationship between transition probability and gap duration: more prevalent transitions were produced with shorter gap durations. We then assessed the degree to which long-term age-dependent changes and acute context-dependent changes to syllable sequencing and timing followed this inverse relationship. Age- but not context-dependent changes to syllable sequencing and timing were inversely related. On average, gap durations at branch points decreased with age, and the magnitude of this decrease was greater for transitions that increased in prevalence than for transitions that decreased in prevalence. In contrast, there was no systematic relationship between acute context-dependent changes to syllable sequencing and timing. Gap durations at branch points decreased when birds produced female-directed courtship song compared to when they produced undirected song, and the magnitude of this decrease was not related to the direction and magnitude of changes to transition probabilities. These analyses suggest that neural mechanisms that regulate syllable sequencing could similarly control syllable timing but also highlight mechanisms that can independently regulate syllable sequencing and timing. 相似文献
8.
Animals modulate sensory processing in concert with motor actions. Parallel copies of motor signals, called corollary discharge (CD), prepare the nervous system to process the mixture of externally and self-generated (reafferent) feedback that arises during locomotion. Commonly, CD in the peripheral nervous system cancels reafference to protect sensors and the central nervous system from being fatigued and overwhelmed by self-generated feedback. However, cancellation also limits the feedback that contributes to an animal’s awareness of its body position and motion within the environment, the sense of proprioception. We propose that, rather than cancellation, CD to the fish lateral line organ restructures reafference to maximize proprioceptive information content. Fishes’ undulatory body motions induce reafferent feedback that can encode the body’s instantaneous configuration with respect to fluid flows. We combined experimental and computational analyses of swimming biomechanics and hair cell physiology to develop a neuromechanical model of how fish can track peak body curvature, a key signature of axial undulatory locomotion. Without CD, this computation would be challenged by sensory adaptation, typified by decaying sensitivity and phase distortions with respect to an input stimulus. We find that CD interacts synergistically with sensor polarization to sharpen sensitivity along sensors’ preferred axes. The sharpening of sensitivity regulates spiking to a narrow interval coinciding with peak reafferent stimulation, which prevents adaptation and homogenizes the otherwise variable sensor output. Our integrative model reveals a vital role of CD for ensuring precise proprioceptive feedback during undulatory locomotion, which we term external proprioception.Animals modulate sensory processing in concert with motor actions. A study of the corollary discharge in zebrafish reveals that it modulates the sensitivity of the lateral line during swimming to prevent sensor adaptation and maintain the high-quality feedback necessary for kinematic control. 相似文献
9.
When we speak, we provide ourselves with auditory speech input. Efficient monitoring of speech is often hypothesized to depend on matching the predicted sensory consequences from internal motor commands (forward model) with actual sensory feedback. In this paper we tested the forward model hypothesis using functional Magnetic Resonance Imaging. We administered an overt picture naming task in which we parametrically reduced the quality of verbal feedback by noise masking. Presentation of the same auditory input in the absence of overt speech served as listening control condition. Our results suggest that a match between predicted and actual sensory feedback results in inhibition of cancellation of auditory activity because speaking with normal unmasked feedback reduced activity in the auditory cortex compared to listening control conditions. Moreover, during self-generated speech, activation in auditory cortex increased as the feedback quality of the self-generated speech decreased. We conclude that during speaking early auditory cortex is involved in matching external signals with an internally generated model or prediction of sensory consequences, the locus of which may reside in auditory or higher order brain areas. Matching at early auditory cortex may provide a very sensitive monitoring mechanism that highlights speech production errors at very early levels of processing and may efficiently determine the self-agency of speech input. 相似文献
10.
Lukas A. Hoffmann Conor W. Kelly David A. Nicholson Samuel J. Sober 《Journal of visualized experiments : JoVE》2012,(69)
Experimental manipulations of sensory feedback during complex behavior have provided valuable insights into the computations underlying motor control and sensorimotor plasticity1. Consistent sensory perturbations result in compensatory changes in motor output, reflecting changes in feedforward motor control that reduce the experienced feedback error. By quantifying how different sensory feedback errors affect human behavior, prior studies have explored how visual signals are used to recalibrate arm movements2,3 and auditory feedback is used to modify speech production4-7. The strength of this approach rests on the ability to mimic naturalistic errors in behavior, allowing the experimenter to observe how experienced errors in production are used to recalibrate motor output.Songbirds provide an excellent animal model for investigating the neural basis of sensorimotor control and plasticity8,9. The songbird brain provides a well-defined circuit in which the areas necessary for song learning are spatially separated from those required for song production, and neural recording and lesion studies have made significant advances in understanding how different brain areas contribute to vocal behavior9-12. However, the lack of a naturalistic error-correction paradigm - in which a known acoustic parameter is perturbed by the experimenter and then corrected by the songbird - has made it difficult to understand the computations underlying vocal learning or how different elements of the neural circuit contribute to the correction of vocal errors13.The technique described here gives the experimenter precise control over auditory feedback errors in singing birds, allowing the introduction of arbitrary sensory errors that can be used to drive vocal learning. Online sound-processing equipment is used to introduce a known perturbation to the acoustics of song, and a miniaturized headphones apparatus is used to replace a songbird''s natural auditory feedback with the perturbed signal in real time. We have used this paradigm to perturb the fundamental frequency (pitch) of auditory feedback in adult songbirds, providing the first demonstration that adult birds maintain vocal performance using error correction14. The present protocol can be used to implement a wide range of sensory feedback perturbations (including but not limited to pitch shifts) to investigate the computational and neurophysiological basis of vocal learning. 相似文献
11.
Background
When one watches a sports game, one may feel her/his own muscles moving in synchrony with the player''s. Such parallels between observed actions of others and one''s own has been well supported in the latest progress in neuroscience, and coined “mirror system.” It is likely that due to such phenomena, we are able to learn motor skills just by observing an expert''s performance. Yet it is unknown whether such indirect learning occurs only at higher cognitive levels, or also at basic sensorimotor levels where sensorimotor delay is compensated and the timing of sensory feedback is constantly calibrated.Methodology/Principal Findings
Here, we show that the subject''s passive observation of an actor manipulating a computer mouse with delayed auditory feedback led to shifts in subjective simultaneity of self mouse manipulation and auditory stimulus in the observing subjects. Likewise, self adaptation to the delayed feedback modulated the simultaneity judgment of the other subjects manipulating a mouse and an auditory stimulus. Meanwhile, subjective simultaneity of a simple visual disc and the auditory stimulus (flash test) was not affected by observation of an actor nor self-adaptation.Conclusions/Significance
The lack of shift in the flash test for both conditions indicates that the recalibration transfer is specific to the action domain, and is not due to a general sensory adaptation. This points to the involvement of a system for the temporal monitoring of actions, one that processes both one''s own actions and those of others. 相似文献12.
Some characteristics of arm movements that humans exhibit during learning the dynamics of reaching are consistent with a
theoretical framework where training results in motor commands that are gradually modified to predict and compensate for novel
forces that may act on the hand. As a first approximation, the motor control system behaves as an adapting controller that
learns an internal model of the dynamics of the task. It approximates inverse dynamics and predicts motor commands that are
appropriate for a desired limb trajectory. However, we had previously noted that subtle motion characteristics observed during
changes in task dynamics challenged this simple model and raised the possibility that adaptation also involved sensory–motor
feedback pathways. These pathways reacted to sensory feedback during the course of the movement. Here we hypothesize that
adaptation to dynamics might also involve a modification of how the CNS responds to sensory feedback. We tested this through
experiments that quantified how the motor system's response to errors during voluntary movements changed as it adapted to
dynamics of a force field. We describe a nonlinear approach that approximates the impedance of the arm, i.e., force response
as a function of arm displacement trajectory. We observe that after adaptation, the impedance function changes in a way that
closely matches and counters the effect of the force field. This is particularly prominent in the long-latency (>100 ms) component
of response to perturbations. Therefore, it appears that practice not only modifies the internal model with which the brain
generates motor commands that initiate a movement, but also the internal model with which sensory feedback is integrated with
the ongoing descending commands in order to respond to error during the movement.
Received: 10 January 2001 / Accepted in revised form: 30 May 2001 相似文献
13.
Auditory feedback is necessary for adult song maintenance in both oscines and psittacines. Although belonging to phylogenically
separated orders, deafened adult oscine Bengalese finches and psittacine budgerigars exhibit similarities in certain aspects
of song changes. An interesting question is whether these birds share common mechanisms for song maintenance. Therefore, it
is important to compare the effects of deafening on adult song patterns among and within orders. Although zebra and Bengalese
finches are closely related oscine species, few studies have performed direct, long-term, quantitative comparisons of their
songs after deafening because suitable song characteristics have not been identified. Based on our previous findings for Bengalese
finch songs, we analyzed zebra finch songs over 9 months after deafening, focusing on changes in the number of syllables categorized
according to fundamental frequencies. Deafened zebra finches demonstrated a gradual but significant decrease in high-frequency
syllables and a tendency to increase low-frequency syllables, similar to deafened Bengalese finches. Although this change
took longer in zebra finches, the altered proportion of syllables eventually stabilized. Results indicated that adult songs
show similar aspects after auditory deprivation, and that neural mechanisms involved in the maintenance of high-frequency
song syllables, using auditory feedback, may be present in both finches despite species differences. 相似文献
14.
Recent studies have provided evidence for sensory-motor adaptive changes and action goal coding of visually guided manual action in premotor and posterior parietal cortices. To extend these results to orofacial actions, devoid of auditory and visual feedback, we used a repetition suppression paradigm while measuring neural activity with functional magnetic resonance imaging during repeated intransitive and silent lip, jaw and tongue movements. In the motor domain, this paradigm refers to decreased activity in specific neural populations due to repeated motor acts and has been proposed to reflect sensory-motor adaptation. Orofacial movements activated a set of largely overlapping, common brain areas forming a core neural network classically involved in orofacial motor control. Crucially, suppressed neural responses during repeated orofacial actions were specifically observed in the left ventral premotor cortex, the intraparietal sulcus, the inferior parietal lobule and the superior parietal lobule. Since no visual and auditory feedback were provided during orofacial actions, these results suggest somatosensory-motor adaptive control of intransitive and silent orofacial actions in these premotor and parietal regions. 相似文献
15.
Voluntary motor commands produce two kinds of consequences. Initially, a sensory consequence is observed in terms of activity in our primary sensory organs (e.g., vision, proprioception). Subsequently, the brain evaluates the sensory feedback and produces a subjective measure of utility or usefulness of the motor commands (e.g., reward). As a result, comparisons between predicted and observed consequences of motor commands produce two forms of prediction error. How do these errors contribute to changes in motor commands? Here, we considered a reach adaptation protocol and found that when high quality sensory feedback was available, adaptation of motor commands was driven almost exclusively by sensory prediction errors. This form of learning had a distinct signature: as motor commands adapted, the subjects altered their predictions regarding sensory consequences of motor commands, and generalized this learning broadly to neighboring motor commands. In contrast, as the quality of the sensory feedback degraded, adaptation of motor commands became more dependent on reward prediction errors. Reward prediction errors produced comparable changes in the motor commands, but produced no change in the predicted sensory consequences of motor commands, and generalized only locally. Because we found that there was a within subject correlation between generalization patterns and sensory remapping, it is plausible that during adaptation an individual''s relative reliance on sensory vs. reward prediction errors could be inferred. We suggest that while motor commands change because of sensory and reward prediction errors, only sensory prediction errors produce a change in the neural system that predicts sensory consequences of motor commands. 相似文献
16.
Yann S. Dufour Xiongfei Fu Luis Hernandez-Nunez Thierry Emonet 《PLoS computational biology》2014,10(6)
Inputs to signaling pathways can have complex statistics that depend on the environment and on the behavioral response to previous stimuli. Such behavioral feedback is particularly important in navigation. Successful navigation relies on proper coupling between sensors, which gather information during motion, and actuators, which control behavior. Because reorientation conditions future inputs, behavioral feedback can place sensors and actuators in an operational regime different from the resting state. How then can organisms maintain proper information transfer through the pathway while navigating diverse environments? In bacterial chemotaxis, robust performance is often attributed to the zero integral feedback control of the sensor, which guarantees that activity returns to resting state when the input remains constant. While this property provides sensitivity over a wide range of signal intensities, it remains unclear how other parameters such as adaptation rate and adapted activity affect chemotactic performance, especially when considering that the swimming behavior of the cell determines the input signal. We examine this issue using analytical models and simulations that incorporate recent experimental evidences about behavioral feedback and flagellar motor adaptation. By focusing on how sensory information carried by the response regulator is best utilized by the motor, we identify an operational regime that maximizes drift velocity along chemical concentration gradients for a wide range of environments and sensor adaptation rates. This optimal regime is outside the dynamic range of the motor response, but maximizes the contrast between run duration up and down gradients. In steep gradients, the feedback from chemotactic drift can push the system through a bifurcation. This creates a non-chemotactic state that traps cells unless the motor is allowed to adapt. Although motor adaptation helps, we find that as the strength of the feedback increases individual phenotypes cannot maintain the optimal operational regime in all environments, suggesting that diversity could be beneficial. 相似文献
17.
Duration estimation is known to be far from veridical and to differ for sensory estimates and motor reproduction. To investigate how these differential estimates are integrated for estimating or reproducing a duration and to examine sensorimotor biases in duration comparison and reproduction tasks, we compared estimation biases and variances among three different duration estimation tasks: perceptual comparison, motor reproduction, and auditory reproduction (i.e. a combined perceptual-motor task). We found consistent overestimation in both motor and perceptual-motor auditory reproduction tasks, and the least overestimation in the comparison task. More interestingly, compared to pure motor reproduction, the overestimation bias was reduced in the auditory reproduction task, due to the additional reproduced auditory signal. We further manipulated the signal-to-noise ratio (SNR) in the feedback/comparison tones to examine the changes in estimation biases and variances. Considering perceptual and motor biases as two independent components, we applied the reliability-based model, which successfully predicted the biases in auditory reproduction. Our findings thus provide behavioral evidence of how the brain combines motor and perceptual information together to reduce duration estimation biases and improve estimation reliability. 相似文献
18.
Songbirds develop their songs by imitating songs of adults. For song learning to proceed normally, the bird's hearing must remain intact throughout the song development process. In many species, song learning takes place during one period early in life, and no more new song elements are learned thereafter. In these so-called close-ended learners, it has long been assumed that once song development is complete, audition is no longer necessary to maintain the motor patterns of full song. However, many of these close-ended learners maintain plasticity in overall song organization; the number and the sequence of song elements included in a song of an individual vary from one utterance to another, although no new song elements are added or lost in adulthood. It is conceivable that these species rely on continued auditory feedback to produce normal song syntax. The Bengalese finch is a close-ended learner that produces considerably variable songs as an adult. In the present study, we found that Bengalese finches require real-time auditory feedback for motor control even after song learning is complete; deafening adult finches resulted in development of abnormal song syntax in as little as 5 days. We also found that there was considerable individual variation in the degree of song deterioration after deafening. The neural mechanisms underlying adult song production in different species of songbirds may be more diverse than has been traditionally considered. © 1997 John Wiley & Sons, Inc. J Neurobiol 33: 343–356, 1997 相似文献
19.
Comparison of human and humanoid robot control of upright stance 总被引:1,自引:0,他引:1
There is considerable recent interest in developing humanoid robots. An important substrate for many motor actions in both humans and biped robots is the ability to maintain a statically or dynamically stable posture. Given the success of the human design, one would expect there are lessons to be learned in formulating a postural control mechanism for robots. In this study we limit ourselves to considering the problem of maintaining upright stance. Human stance control is compared to a suggested method for robot stance control called zero moment point (ZMP) compensation. Results from experimental and modeling studies suggest there are two important subsystems that account for the low- and mid-frequency (DC to 1 Hz) dynamic characteristics of human stance control. These subsystems are (1) a “sensory integration” mechanism whereby orientation information from multiple sensory systems encoding body kinematics (i.e. position, velocity) is flexibly combined to provide an overall estimate of body orientation while allowing adjustments (sensory re-weighting) that compensate for changing environmental conditions and (2) an “effort control” mechanism that uses kinetic-related (i.e., force-related) sensory information to reduce the mean deviation of body orientation from upright. Functionally, ZMP compensation is directly analogous to how humans appear to use kinetic feedback to modify the main sensory integration feedback loop controlling body orientation. However, a flexible sensory integration mechanism is missing from robot control leaving the robot vulnerable to instability in conditions where humans are able to maintain stance. We suggest the addition of a simple form of sensory integration to improve robot stance control. We also investigate how the biological constraint of feedback time delay influences the human stance control design. The human system may serve as a guide for improved robot control, but should not be directly copied because the constraints on robot and human control are different. 相似文献
20.
Heron J Aaen-Stockdale C Hotchkiss J Roach NW McGraw PV Whitaker D 《Proceedings. Biological sciences / The Royal Society》2012,279(1729):690-698
The task of deciding how long sensory events seem to last is one that the human nervous system appears to perform rapidly and, for sub-second intervals, seemingly without conscious effort. That these estimates can be performed within and between multiple sensory and motor domains suggest time perception forms one of the core, fundamental processes of our perception of the world around us. Given this significance, the current paucity in our understanding of how this process operates is surprising. One candidate mechanism for duration perception posits that duration may be mediated via a system of duration-selective 'channels', which are differentially activated depending on the match between afferent duration information and the channels' 'preferred' duration. However, this model awaits experimental validation. In the current study, we use the technique of sensory adaptation, and we present data that are well described by banks of duration channels that are limited in their bandwidth, sensory-specific, and appear to operate at a relatively early stage of visual and auditory sensory processing. Our results suggest that many of the computational principles the nervous system applies to coding visual spatial and auditory spectral information are common to its processing of temporal extent. 相似文献