Biomedical subjects

Sarah M N Woolley

Publications and source records attributed to Sarah M N Woolley.

10 recordsLinked to original sources

PubMed2006

Sound representation methods for spectro-temporal receptive field estimation.

The spectro-temporal receptive field (STRF) of an auditory neuron describes the linear relationship between the sound stimulus in a time-frequency representation and the neural response. Time-frequency representations of a sound in turn require a nonlinear operation on the sound pressure waveform and many different forms for this non-linear transformation are possible. Here, we systematically investigated the effects of four factors in the non-linear step in the STRF model: the choice of logarithmic or linear filter frequency spacing, the time-frequency scale, stimulus amplitude compression and adaptive gain control. We quantified the goodness of fit of these different STRF models on data obtained from auditory neurons in the songbird midbrain and forebrain. We found that adaptive gain control and the correct stimulus amplitude compression scheme are paramount to correctly modelling neurons. The time-frequency scale and frequency spacing also affected the goodness of fit of the model but to a lesser extent and the optimal values were stimulus dependent.

Acoustic Stimulation↗

PubMed2006

Stimulus-dependent auditory tuning results in synchronous population coding of vocalizations in the songbird midbrain.

Physiological studies in vocal animals such as songbirds indicate that vocalizations drive auditory neurons particularly well. But the neural mechanisms whereby vocalizations are encoded differently from other sounds in the auditory system are unknown. We used spectrotemporal receptive fields (STRFs) to study the neural encoding of song versus the encoding of a generic sound, modulation-limited noise, by single neurons and the neuronal population in the zebra finch auditory midbrain. The noise was designed to match song in frequency, spectrotemporal modulation boundaries, and power. STRF calculations were balanced between the two stimulus types by forcing a common stimulus subspace. We found that 91% of midbrain neurons showed significant differences in spectral and temporal tuning properties when birds heard song and when birds heard modulation-limited noise. During the processing of noise, spectrotemporal tuning was highly variable across cells. During song processing, the tuning of individual cells became more similar; frequency tuning bandwidth increased, best temporal modulation frequency increased, and spike timing became more precise. The outcome was a population response to song that encoded rapidly changing sounds with power and precision, resulting in a faithful neural representation of the temporal pattern of a song. Modeling responses to song using the tuning to modulation-limited noise showed that the population response would not encode song as precisely or robustly. We conclude that stimulus-dependent changes in auditory tuning during song processing facilitate the high-fidelity encoding of the temporal pattern of a song.

Acoustic Stimulation↗

PubMed2005

Tuning for spectro-temporal modulations as a mechanism for auditory discrimination of natural sounds.

Vocal communicators discriminate conspecific vocalizations from other sounds and recognize the vocalizations of individuals. To identify neural mechanisms for the discrimination of such natural sounds, we compared the linear spectro-temporal tuning properties of auditory midbrain and forebrain neurons in zebra finches with the statistics of natural sounds, including song. Here, we demonstrate that ensembles of auditory neurons are tuned to auditory features that enhance the acoustic differences between classes of natural sounds, and among the songs of individual birds. Tuning specifically avoids the spectro-temporal modulations that are redundant across natural sounds and therefore provide little information; rather, it overlaps with the temporal modulations that differ most across sounds. By comparing the real tuning and a less selective model of spectro-temporal tuning, we found that the real modulation tuning increases the neural discrimination of different sounds. Additionally, auditory neurons discriminate among zebra finch song segments better than among synthetic sound segments.

Acoustic Stimulation↗

PubMed2005

Processing of modulated sounds in the zebra finch auditory midbrain: responses to noise, frequency sweeps, and sinusoidal amplitude modulations.

The avian auditory midbrain nucleus, the mesencephalicus lateralis, dorsalis (MLd), is the first auditory processing stage in which multiple parallel inputs converge, and it provides the input to the auditory thalamus. We studied the responses of single MLd neurons to four types of modulated sounds: 1) white noise; 2) band-limited noise; 3) frequency modulated (FM) sweeps, and 4) sinusoidally amplitude-modulated tones (SAM) in adult male zebra finches. Responses were compared with the responses of the same neurons to pure tones in terms of temporal response patterns, thresholds, characteristic frequencies, frequency tuning bandwidths, tuning sharpness, and spike rate/intensity relationships. Most neurons responded well to noise. More than one-half of the neurons responded selectively to particular portions of the noise, suggesting that, unlike forebrain neurons, many MLd neurons can encode specific acoustic components of highly modulated sounds such as noise. Selectivity for FM sweep direction was found in only 13% of cells that responded to sweeps. Those cells also showed asymmetric tuning curves, suggesting that asymmetric inhibition plays a role in FM directional selectivity. Responses to SAM showed that MLd neurons code temporal modulation rates using both spike rate and synchronization. Nearly all cells showed low-pass or band-pass filtering properties for SAM. Best modulation frequencies matched the temporal modulations in zebra finch song. Results suggest that auditory midbrain neurons are well suited for encoding a wide range of complex sounds with a high degree of temporal accuracy rather than selectively responding to only some sounds.

Acoustic Stimulation↗

PubMed2004

Modulation power and phase spectrum of natural sounds enhance neural encoding performed by single auditory neurons.

We examined the neural encoding of synthetic and natural sounds by single neurons in the auditory system of male zebra finches by estimating the mutual information in the time-varying mean firing rate of the neuronal response. Using a novel parametric method for estimating mutual information with limited data, we tested the hypothesis that song and song-like synthetic sounds would be preferentially encoded relative to other complex, but non-song-like synthetic sounds. To test this hypothesis, we designed two synthetic stimuli: synthetic songs that matched the power of spectral-temporal modulations but lacked the modulation phase structure of zebra finch song and noise with uniform band-limited spectral-temporal modulations. By defining neural selectivity as relative mutual information, we found that the auditory system of songbirds showed selectivity for song-like sounds. This selectivity increased in a hierarchical manner along ascending processing stages in the auditory system. Midbrain neurons responded with highest information rates and efficiency to synthetic songs and thus were selective for the spectral-temporal modulations of song. Primary forebrain neurons showed increased information to zebra finch song and synthetic song equally over noise stimuli. Secondary forebrain neurons responded with the highest information to zebra finch song relative to other stimuli and thus were selective for its specific modulation phase relationships. We also assessed the relative contribution of three response properties to this selectivity: (1) spiking reliability, (2) rate distribution entropy, and (3) bandwidth. We found that rate distribution and bandwidth but not reliability were responsible for the higher average information rates found for song-like sounds.

Acoustic Stimulation↗

PubMed2004

Auditory experience and adult song plasticity.

Adults Bengalese finches normally sing stereotyped songs, which do not change under the influence of auditory experience. If deafened, however, adult birds sing significantly degraded songs that are characterized by a lack of stereotypy of syllable order and the deterioration of syllable structure. We studied the importance of auditory feedback for maintenance of normal adult song. Auditory feedback can be partially or completely removed by eliminating auditory hair cells, the sensory receptor cells for hearing. The effects are reversible because birds regenerate new hair cells when original cells are lost. Limiting the frequency range of auditory feedback available to birds using partial hair cell lesions indicated that low-frequency information (1500 Hz) is necessary and sufficient for the maintenance of normal adult song. Reversible deafening experiments using hair cell loss and regeneration indicated that adult Bengalese finches store memories of their own songs, which can be used to guide vocal behavior. Additionally, destabilizing song behavior by removing auditory feedback leads to renewed song plasticity, and some adults can be induced to learn new song. These studies suggest that Bengalese finches maintain normal song by comparing ongoing vocal output to stored models of their own stable songs and that neural circuitry for song learning persists beyond the sensitive period for song learning.

Animals↗

PubMed2004

Methods for the analysis of auditory processing in the brain.

Understanding song perception and singing behavior in birds requires the study of auditory processing of complex sounds throughout the avian brain. We can divide the basics of auditory perception into two general processes: (1) encoding, the process whereby sound is transformed into neural activity and (2) decoding, the process whereby patterns of neural activity take on perceptual meaning and therefore guide behavioral responses to sounds. In birdsong research, most studies have focused on the decoding process: What are the responses of the specialized auditory neurons in the song control system? and What do they mean for the bird? Recently, new techniques addressing both encoding and decoding have been developed for use in songbirds. Here, we first describe some powerful methods for analyzing what acoustical aspects of complex sounds like songs are encoded by auditory processing neurons in songbird brain. These methods include the estimation and analysis of spectro-temporal receptive fields (STRFs) for auditory neurons. Then we discuss the decoding methods that have been used to understand how songbird neurons may discriminate among different songs and other sounds based on mean spike-count rates.

Animals↗

PubMed2004

Song selectivity in the song system and in the auditory forebrain.

The sensorimotor neurons found in the song-system nuclei are responsive to the sounds of the bird's own song. This selectivity emerges during vocal learning and appears to follow the development of the bird's song vocalization in two ways: at each stage, the neurons are most selective for the bird's current vocalizations and this selectivity increases as the bird learns to produce a stable adult song. Also, because of their location in the sensori-vocal pathway and because their physiological properties are correlated with the motor program, it is postulated that these neurons play a crucial role in interpreting the auditory feedback during song to preserve a desirable vocal output. The neurons found in presynaptic auditory areas lack this selectivity for the bird's own song. Auditory neurons in the secondary auditory areas caudal nidopallium and caudal mesopallium show specific responses to familiar songs or behaviorally relevant songs. These auditory areas might therefore be involved in perceptual tasks. Neurons in the primary forebrain auditory area are selective for the spectrotemporal modulations that are common in song, yielding an efficient neural representation of those sounds. Neurons that are particularly selective for the tutor song at the end of the sensory period have not yet been described in any areas. Although these three levels of selectivity found in the primary auditory forebrain areas, the secondary auditory forebrain areas, and the song system suggest a form of hierarchical sensory processing, the functional connectivity between these areas and the mechanisms generating the specific selectivity for songs that are behaviorally relevant or crucial in song learning and production have yet to be revealed.

Animals↗

PubMed2003

Response properties of single neurons in the zebra finch auditory midbrain: response patterns, frequency coding, intensity coding, and spike latencies.

The avian mesencephalicus lateralis, dorsalis (MLd) is the auditory midbrain nucleus in which multiple parallel inputs from lower brain stem converge and through which most auditory information passes to reach the forebrain. Auditory processing in the MLd has not been investigated in songbirds. We studied the tuning properties of single MLd neurons in adult male zebra finches. Pure tones were used to examine tonotopy, temporal response patterns, frequency coding, intensity coding, spike latencies, and duration tuning. Most neurons had no spontaneous activity. The tonotopy of MLd is like that of other birds and mammals; characteristic frequencies (CFs) increase in a dorsal to ventral direction. Four major response patterns were found: 1) onset (49% of cells); 2) primary-like (20%); 3) sustained (19%); and 4) primary-like with notch (12%). CFs ranged between 0.9 and 6.1 kHz, matching the zebra finch hearing range and the power spectrum of song. Tuning curves were generally V-shaped, but complex curves, with multiple peaks or noncontiguous excitatory regions, were observed in 22% of cells. Rate-level functions indicated that 51% of nononset cells showed monotonic relationships between spike rate and sound level. Other cells showed low saturation or nonmonotonic responses. Spike latencies ranged from 4 to 40 ms, measured at CF. Spike latencies generally decreased with increasing sound pressure level (SPL), although paradoxical latency shifts were observed in 16% of units. For onset cells, changes in SPL produced smaller latency changes than for cells showing other response types. Results suggest that auditory midbrain neurons may be particularly suited for processing temporally complex signals with a high degree of precision.

Acoustic Stimulation↗

PubMed2002

Vocal memory and learning in adult Bengalese Finches with regenerated hair cells.

Critical learning periods are common in vertebrate development. In many birds, song learning is limited by a critical period; juveniles copy songs from adult birds by forming memories of those songs during a restricted developmental period and then using auditory feedback to practice their own vocalizations. Adult songs are stable over time regardless of exposure to other birds, but auditory feedback is required for the maintenance of stable adult song. A technique was developed to reversibly deafen Bengalese Finches by destruction and regeneration of inner ear auditory hair cells. With this approach, we asked two questions about the plasticity of song information stored in the adult brain. First, do adult birds store memories or "templates" of their songs that exist independent of auditory reinforcement? Such memories could be used to control vocal output by acting as fixed models of song to which ongoing vocalizations are matched. Second, can adult song learning, which does not normally occur in this species, be induced by removing and then restoring hearing? Studying changes in adult song behavior during hair cell loss and regeneration revealed two findings: (1) adult birds store memories or templates of their songs that exist independent of auditory input and can be used to restore normal vocal behavior when hearing is restored; (2) under experimental circumstances, adult birds can be induced to acquire song material from other birds. Results suggest that, in Bengalese Finches, the degree of behavioral and neural plasticity in juvenile and adult birds may be less distinct that previously thought.

Animals↗