Michael S Brainard's research while affiliated with University of California, San Francisco and other places
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Publications (80)
Sensory feedback is required for stable execution of learned motor skills, and its loss can severely disrupt motor performance. The neural mechanisms that mediate sensorimotor stability have been extensively studied at systems and physiological levels, yet relatively little is known about how disruptions to sensory input alter the molecular propert...
Songbirds and humans share the ability to adaptively modify their vocalizations based on sensory feedback. Prior studies have focused primarily on the role that auditory feedback plays in shaping vocal output throughout life. In contrast, it is unclear how non-auditory information drives vocal plasticity. Here, we first used a reinforcement learnin...
Complex behaviors depend on the coordinated activity of neural ensembles in interconnected brain areas. The behavioral function of such coordination, often measured as co-fluctuations in neural activity across areas, is poorly understood. One hypothesis is that rapidly varying co-fluctuations may be a signature of moment-by-moment task-relevant inf...
Songbirds and humans share the ability to adaptively modify their vocalizations based on sensory feedback. Prior studies have focused primarily on the role that auditory feedback plays in shaping vocal output throughout life. In contrast, it is unclear whether and how non-auditory information drives vocal plasticity. Here, we first used a reinforce...
The flexible control of sequential behavior is a fundamental aspect of speech, enabling endless reordering of a limited set of learned vocal elements (syllables or words). Songbirds are phylogenetically distant from humans but share both the capacity for vocal learning and neural circuitry for vocal control that includes direct pallial-brainstem pr...
Complex learned behaviors exhibit striking variation within populations, yet how heritable factors contribute to such inter-individual differences remains largely unknown. Here, we used behavioral-genetic analysis within a Bengalese finch population ( Lonchura striata domestica ) to investigate molecular and circuit mechanisms underlying heritable...
The cells of songbird motor circuits
Birds have complex motor and cognitive abilities that rival or exceed the performance of many mammals, but their brains are organized in a notably different way. Parts of the bird brain have been functionally compared to the mammalian neocortex. However, it is still controversial to what extent these regions are...
The flexible control of sequential behavior is a fundamental aspect of speech, enabling endless reordering of a limited set of learned vocal elements (i.e. syllables or words). Songbirds are phylogenetically distant from humans, but share the capacity for vocal learning as well as neural circuitry for vocal control that includes direct cortical-bra...
Acetylcholine is well-understood to enhance cortical sensory responses and perceptual sensitivity in aroused or attentive states. Yet little is known about cholinergic influences on motor cortical regions. Here we use the quantifiable nature of birdsong to investigate how acetylcholine modulates the cortical (pallial) premotor nucleus HVC and shape...
Acetylcholine is well-understood to enhance cortical sensory responses and perceptual sensitivity in aroused or attentive states. Yet little is known about cholinergic influences on motor cortical regions. Here we use the quantifiable nature of birdsong to investigate how acetylcholine modulates the cortical (pallial) premotor nucleus HVC and shape...
Acetylcholine is well-understood to enhance cortical sensory responses and perceptual sensitivity in aroused or attentive states. Yet little is known about cholinergic influences on motor cortical regions. Here we use the quantifiable nature of birdsong to investigate how acetylcholine modulates the cortical (pallial) premotor nucleus HVC and shape...
It is widely argued that personalized instruction based on individual differences in learning styles or genetic predispositions could improve learning outcomes. However, this proposition has resisted clear demonstration in human studies, where it is difficult to control experience and quantify outcomes. Here, we take advantage of the tractable natu...
The basal ganglia (BG) participate in aspects of reinforcement learning that require evaluation and selection of motor programs associated with improved performance. However, whether the BG additionally contribute to behavioral variation (“motor exploration”) that forms the substrate for such learning remains unclear. In songbirds, a tractable syst...
This script implements the Gaussian mixture model selection process detailed in the paper.
It provides an estimate of the number of syllables present in a specific set of songs. It is intended to be used to estimate the number of syllables present in a given set of song data prior to calculating the Song DKL. Details and requirements for usage are...
Song DKL is consistent when greater than 1000 input syllables are used.
(A-D) Correlation between DKL calculated with 3000 syllables of input data vs. 100, 500, 1000, and 2000 syllables of input data. DKL values calculated from 44 song comparisons are plotted. Unity line is shown in red.
(TIFF)
Studies of learning mechanisms critically depend on the ability to accurately assess learning outcomes. This assessment can be impeded by the often complex, multidimensional nature of behavior. We present a novel, automated approach to evaluating imitative learning. Conceptually, our approach estimates how much of the content present in a reference...
The influence of increasing temporal sampling density of syllable spectral content on Song DKL.
(A-C) Correlation between DKL derived from syllables represented with 10 PSDs, evenly distributed across syllable duration, and syllables represented with one, two and five PSDs. DKL values calculated from 44 song comparisons are plotted. Unity line is s...
Song DKL is consistent when greater than 40 basis syllables are used.
(A-E) Correlation between Song DKL calculated for 160 basis syllables vs. 5, 10, 20, 40, and 80 basis syllables. DKL values calculated from 44 song comparisons are plotted. Unity line is shown in red.
(TIFF)
Distributions of handwritten numerals in numeral similarity-space are separated based on numeral identity and are distributed approximately normally in any given dimension.
Handwritten numerals (zero and one) for this analysis were drawn from the MNIST handwritten digit data set. The data set contains examples of each numeral from many individuals....
Song DKL is robust to changes in the number of Gaussian mixture components.
(A-H) Correlation between DKL calculated for models with the number of mixture components indicated by the BIC (nBIC) and models with deviations from this value ranging from nBIC-4 to nBIC+4. DKL values calculated from 44 song comparisons are plotted. Unity line is shown in...
This script implements the Song DKL calculations detailed in the paper.
It calculates the Song DKL between two sets of song data. Details and requirements for usage are in the script.
(PY)
Discrimination between spoken words composed of overlapping elements, such as “captain” and “captive,” relies on sensitivity to unique combinations of prefix and suffix elements that span a “uniqueness point” where the word candidates diverge. To model such combinatorial processing, adult female zebra finches were trained to discriminate between ta...
Background
Vocal learning in songbirds has emerged as a powerful model for sensorimotor learning. Neuro-behavioral studies of Bengalese finch (Lonchura striata domestica) song, naturally more variable and plastic than songs of other finch species, have demonstrated the importance of behavioral variability for initial learning, maintenance, and plas...
Significance
Learning reflects the influence of experience on genetically determined circuitry, but little is known about how experience and genetics interact to determine learned phenotypes. Here, we use vocal learning in songbirds to study genetic influences on learned behavior. We first show that the tempo of learned song is strongly influenced...
Skill learning is instantiated by changes to functional connectivity within premotor circuits, but whether the specificity of learning depends on structured changes to inhibitory circuitry remains unclear. We used slice electrophysiology to measure connectivity changes associated with song learning in the avian analog of primary motor cortex (robus...
Motor skills depend on the reuse of individual gestures in multiple sequential contexts (e.g., a single phoneme in different words). Yet optimal performance requires that a given gesture be modified appropriately depending on the sequence in which it occurs. To investigate the neural architecture underlying such context-dependent modifications, we...
Studies of learning mechanisms critically depend on the ability to accurately assess learning outcomes. This assessment can be impeded by the often complex, multidimensional nature of behavior. We present a novel, automated approach to evaluating imitative learning that is founded in information theory. Conceptually, our approach estimates the amou...
To investigate mechanisms of action sequencing, we examined the relationship between timing and sequencing of syllables in Bengalese finch song. An individual's song comprises acoustically distinct syllables organized into probabilistic sequences: a given syllable potentially can transition to several different syllables (divergence points) and sev...
Significance
The neural basis of prediction is poorly understood. Here, utilizing the probabilistic song sequences of the Bengalese finch, we recorded neural activity following the termination of auditory playback of an individual’s produced song sequences. We demonstrate that induced neural activity is predictive of the timing and identity of indi...
The majority of distinct sensory and motor events occur as temporally ordered sequences with rich probabilistic structure. Sequences can be characterized by the probability of transitioning from the current state to upcoming states (forward probability), as well as the probability of having transitioned to the current state from previous states (ba...
Consecutive repetition of actions is common in behavioral sequences. Although
integration of sensory feedback with internal motor programs is important for
sequence generation, if and how feedback contributes to repetitive actions is
poorly understood. Here we study how auditory feedback contributes to
generating repetitive syllable sequences in so...
Many complex behaviors, such as human speech and birdsong, reflect a set of categorical actions that can be flexibly organized into variable sequences. However, little is known about how the brain encodes the probabilities of such sequences. Behavioral sequences are typically characterized by the probability of transitioning from a given action to...
Many complex behaviors are supported by neurons with both sensory and motor properties. During behavior such sensory-motor neurons experience the probabilistic associations of pre-motor activity for current actions with feedback from previous actions. Consequently, these associations might become encoded through Hebbian mechanisms. To investigate t...
Songbirds, long of interest to basic neuroscience, have great potential as a model system for translational neuroscience. Songbirds learn their complex vocal behavior in a manner that exemplifies general processes of perceptual and motor skill learning and, more specifically, resembles human speech learning. Song is subserved by circuitry that is s...
The brain uses sensory feedback to correct behavioral errors. Larger errors by definition require greater corrections, and many models of learning assume that larger sensory feedback errors drive larger motor changes. However, an alternative perspective is that larger errors drive learning less effectively because such errors fall outside the range...
Variation in sequencing of actions occurs in many natural behaviors, yet how such variation is maintained is poorly understood. We investigated maintenance of sequence variation in adult Bengalese finch song, a learned skill with rendition-to-rendition variation in the sequencing of discrete syllables (i.e., syllable "b" might transition to "c" wit...
Vocalizations used by birds for territory defense, mate attraction, or both are often referred to as a given species' song. Birdsong refers to the often complex vocalizations produced most frequently by males of species that are members of the songbird order (passeriformes). Unlike most species-typical vocalizations produced by nonhuman animals, so...
Songbirds have emerged as a premier model system for studying how brain circuits learn and produce complex action sequences. The adult song of the most widely studied songbird, the zebra finch (ZF), consists of repeats of a stereotyped sequence of vocal gestures known as syllables. These songs are incredibly precise, with individual syllables and i...
We learn complex skills such as speech and dance through a gradual process of trial and error. Cortical-basal ganglia circuits have an important yet unresolved function in this trial-and-error skill learning; influential 'actor-critic' models propose that basal ganglia circuits generate a variety of behaviours during training and learn to implement...
In songbirds, the basal ganglia outflow nucleus LMAN is a cortical analog that is required for several forms of song plasticity and learning. Moreover, in adults, inactivating LMAN can reverse the initial expression of learning driven via aversive reinforcement. In the present study, we investigated how LMAN contributes to both reinforcement-driven...
Reinforcement signals indicating success or failure are known to alter the probability of selecting between distinct actions. However, successful performance of many motor skills, such as speech articulation, also requires learning behavioral trajectories that vary continuously over time. Here, we investigated how temporally discrete reinforcement...
The control of sequenced behaviors, including human speech, requires that the brain coordinate the production of discrete motor elements with their concatenation into complex patterns. In birdsong, another sequential vocal behavior, the acoustic structure (phonology) of individual song elements, or "syllables," must be coordinated with the sequenci...
Sensory feedback is important for the learning and control of a variety of behaviors. Vocal motor production in songbirds is a powerful model system to study sensory influences on behavior because the learning, maintenance, and control of song are critically dependent on auditory feedback. Based on previous behavioral and neural experiments, it has...
Humans learn to speak by a process of vocal imitation that requires the availability of auditory feedback. Similarly, young birds rely on auditory feedback when learning to imitate the songs of adult birds, providing one of the few examples of nonhuman vocal learning. However, although humans continue to use auditory feedback to correct vocal error...
Behavioral variability is important for motor skill learning but continues to be present and actively regulated even in well-learned behaviors. In adult songbirds, two types of song variability can persist and are modulated by social context: variability in syllable structure and variability in syllable sequencing. The degree to which the control o...
Birdsong, like human speech, relies critically on auditory feedback to provide information about the quality of vocalizations. Although the importance of auditory feedback to vocal learning is well established, whether and how feedback signals influence vocal premotor circuitry has remained obscure. Previous studies in singing birds have not detect...
Birdsong is a learned behavior remarkable for its high degree of stereotypy. Nevertheless, adult birds display substantial rendition-by-rendition variation in the structure of individual song elements or "syllables." Previous work suggests that some of this variation is actively generated by the avian basal ganglia circuitry for purposes of motor e...
Birdsong is a learned motor skill that is performed with a high degree of stereotypy in adult birds. Nevertheless, even in species where song "crystallizes" in a form that remains stable over time, there is residual variability. Such variability in well-learned skills is often construed as uncontrolled and irrelevant biological "noise." However, st...
Significant trial-by-trial variation persists even in the most practiced skills. One prevalent view is that such variation is simply 'noise' that the nervous system is unable to control or that remains below threshold for behavioural relevance. An alternative hypothesis is that such variation enables trial-and-error learning, in which the motor sys...
Trial-by-trial variability is important in feedback-based motor learning. Variation in motor output enables evaluation mechanisms to differentially reinforce patterns of motor activity that produce desired behaviors. Here, we studied neural substrates of variability in the performance of adult birdsong, a complex, learned motor skill used for court...
Songbirds and humans both rely critically on hearing for learning and maintaining accurate vocalizations. Evidence strongly indicates that auditory feedback contributes in real time to human speech, but similar contributions of feedback to birdsong remain unclear. Here, we assessed real-time influences of auditory feedback on Bengalese finch song u...
Cortical-basal ganglia circuits have a critical role in motor control and motor learning. In songbirds, the anterior forebrain pathway (AFP) is a basal ganglia-forebrain circuit required for song learning and adult vocal plasticity but not for production of learned song. Here, we investigate functional contributions of this circuit to the control o...
The anterior forebrain pathway (AFP) is a basal ganglia-dorsal forebrain circuit that is prominent specifically in birds that learn to sing. This circuit is interconnected with the song motor pathway, is active during song production, and contains neurons that are selective for the sound of the bird's own song, suggesting an important role for the...
Bird fanciers have known for centuries that songbirds learn their songs. This learning has striking parallels to speech acquisition: like humans, birds must hear the sounds of adults during a sensitive period, and must hear their own voice while learning to vocalize. With the discovery and investigation of discrete brain structures required for sin...
Birdsong is a learned, sequenced motor skill. For the zebra finch, learned song normally remains unchanging beyond early adulthood. However, stable adult song will gradually deteriorate after deafening (Nordeen and Nordeen, 1992), indicating an ongoing influence of auditory feedback on learned song. This plasticity of adult song in response to deaf...
Bird song, like human speech, is a learned vocal behavior that requires auditory feedback. Both as juveniles, while they learn to sing, and as adults, songbirds use auditory feedback to compare their own vocalizations with an internal model of a target song. Here we describe experiments that explore a role for the songbird anterior forebrain pathwa...
Songbirds are one of the best-studied examples of vocal learners. Learning of both human speech and birdsong depends on hearing. Once learned, adult song in many species remains unchanging, suggesting a reduced influence of sensory experience. Recent studies have revealed, however, that adult song is not always stable, extending our understanding o...
Birdsong, like speech, is a learned vocal behaviour that relies greatly on hearing; in both songbirds and humans the removal of auditory feedback by deafening leads to a gradual deterioration of adult vocal production. Here we investigate the neural mechanisms that contribute to the processing of auditory feedback during the maintenance of song in...
Previous studies have identified sensitive periods for the developing barn owl during which visual experience has a powerful influence on the calibration of sound localization behavior. Here we investigated neural correlates of these sensitive periods by assessing developmental changes in the capacity of visual experience to alter the map of audito...
Alignment of auditory and visual receptive fields in the optic tectum of the barn owl (Tyto alba) is maintained through experience-dependent modification of auditory responses in the external nucleus of the inferior colliculus
(ICX), which provides auditory input to the tectum. Newly learned tectal auditory responses, induced by altered visual expe...
1. In the optic tectum of normal barn owls, bimodal (auditory-visual) neurons are tuned to the values of interaural time difference (ITD) that are produced by sounds at the locations of their visual receptive fields (VRFs). The auditory tuning of tectal neurons is actively guided by visual experience during development: in the tectum of adult owls...
Recent behavioral experiments suggest that parallel pathways in the forebrain and midbrain are capable of mediating sound localization. Neurophysiological studies have advanced our understanding of how binaural and monaural localization cues contribute to the representations of auditory space within these pathways. Experience-dependent modification...
The optic tectum (homolog of the superior colliculus) con-tains mutually aligned neural maps of auditory and visual space. During development, the organization of the auditory map is guided by spatial information provided by vision: barn owls raised wearing prismatic spectacles, which optically shift the visual field and the visual map in the optic...
Cues for sound localization are inherently spatially ambiguous. Nevertheless, most neurons in the barn owl's optic tectum (superior colliculus) have receptive fields for broadband noise stimuli that are restricted to a single region of space. This study characterizes the spatial ambiguities associated with two important sets of localization cues, i...
Neural maps of visual and auditory space are aligned in the adult optic tectum. In barn owls, this alignment of sensory maps
was found to be controlled during ontogeny by visual instruction of the auditory spatial tuning of neurons. Large adaptive
changes in auditory spatial tuning were induced by raising owls with displacing prisms mounted in spec...
Citations
... This widely conserved feature appears to have been exapted for both the modification of vocal circuits essential for language learning in humans and song learning in birds. A recent paper that employed cutaneous sensory stimuli to shape vocal production rather than auditory feedback, also identified a role for dopamine in vocal learning (McGregor et al., 2022). A role for dopamine in reproductive-state dependent odor preferences has recently been identified in Drosophila as well (Boehm et al., 2022). ...
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... Recent studies have examined whether non-auditory feedback can substitute for the white noise. They provided the first evidence that a visual (a light-off or a light-on) as well as a tactile stimulus (applied to the scalp), respectively, can guide changes in the pitch value (McGregor et al., 2021;Zai et al., 2020). These results support the idea that the ability to carefully adjust one's own vocalizations, shared by humans and songbirds, can efficiently rely on information from several different modalities. ...
... Conversely, rapid adaptation in response to changing environmental contexts has been demonstrated in both birdsong [67][68][69] and human speech [70]. This apparent plasticity of a well learned, crystalized behavior suggests that vocalization may be controlled by a "malleable template," in which trial-by-trial variability is used to adapt learned behaviors [71]. ...
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... Circuit organization characterized by parallel processing and functional "switches" between parallel streams allows for a distributed but interconnected circuit structure. Such a structure permits variability in specific components that comprise larger behavioral routines in response to genetic or environmental factors, and has been suggested as a mechanism by which the nervous system achieves behavioral flexibility (Bargmann, 2012;Chan et al., 2002;Falkner et al., 2014;Hashikawa et al., 2016;Mets et al., 2021;Ragozzino, 2007). Across such distributed networks, neuromodulators and hormones coordinate circuit engagement, i.e., levels or patterns of activity within local circuits or the influence of their output (Bargmann, 2012;Cohn et al., 2015;Marder, 2012). ...
... Recent advances in single-cell RNA sequencing (scRNA-seq) have had significant effects on the study of complex tissues, leading to the discovery of novel cell types, cell states, and biomarkers [Stuart and Satija, 2019;Luecken and Theis, 2019;Alfieri et al., 2022;Zeng, 2022]. The scRNA-seq technology has opened up a plethora of opportunities to perform novel studies using new and classic model animals, including chickens (Gallus gallus) [Yamagata, 2022] and other birds Colquitt et al., 2021]. A chicken cell atlas project (aka, Tabula Gallus) has been proposed to create a cell atlas of all tissues in the mature and developing chicken [Yamagata, 2022]. ...
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... Song similarity analysis. To quantify the similarity between tutored or untutored songs and the tutor song ( Supplementary Fig. 3), we applied a method for automatically classifying vocalizations in an unbiased and unsupervised fashion 40 based on their acoustic content. Briefly, this method extracts syllables from a test song (e.g., from a tutored or untutored bird) and assembles a statistical model based on the syllables' acoustic content. ...
... 7,27 ACh is also linked to cognitive processes, including attention and memory, 28,29 signaling transition in movement state, and invigorating volitional movements. [30][31][32][33] Despite the diverse functions of the cholinergic system, a common theme is potentiating action in response to environmental stimuli. 29,34 This is supported by emerging evidence that ACh can rapidly and selectively modulate activity in specific brain areas. ...
... However, such modulation of song structure is often described as affectively controlled (Berwick et al., 2011;Nieder and Mooney, 2020). For example, the presence of potential mates or rivals elicits a global and unlearned modulation of song intensity (James et al., 2018) related to the singer's level of arousal or aggression (Alcami et al., 2021;Heinig et al., 2014;Jaffe and Brainard, 2020). Hence, while prior observations suggest that a variety of ethologically relevant factors can be integrated to influence song production in natural settings, it remains unclear whether song can be modified more flexibly by learned or cognitive factors. ...
... Several authors showed that genetic factors have a strong influence on song learning but a recent study by Mets & Brainard (2019) highlighted that genetic effects can be balanced by exposing pupils Bengalese finches (Lonchura striata domestica) to songs specifically created to corresponding to their learning styles. This observation could be very useful for future experiments as we could use the zebra finch robot to develop experimental procedures to balance genetic predispositions. ...
... On the other hand, several studies have shown the importance of within-trial variability, or f o fluctuations in one vocal element, for vocal adaptations 15,17 . Our study here measured the variability as the fluctuation in each prolonged vowel production and the adjustment as the compensatory response observed within each trial in human vocalization, while the relationship between the trial-by-trial variability and adaptive learnings over trials should be tested in future studies. ...
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