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Jan W H Schnupp

Publications and source records attributed to Jan W H Schnupp.

12 recordsLinked to original sources

Plasticity of temporal pattern codes for vocalization stimuli in primary auditory cortex.

It has been suggested that "call-selective" neurons may play an important role in the encoding of vocalizations in primary auditory cortex (A1). For example, marmoset A1 neurons often respond more vigorously to natural than to time-reversed twitter calls, although the spectral energy distribution in the natural and time-reversed signals is the same. Neurons recorded in cat A1, in contrast, showed no such selectivity for natural marmoset calls. To investigate whether call selectivity in A1 can arise purely as a result of auditory experience, we recorded responses to marmoset calls in A1 of naive ferrets, as well as in ferrets that had been trained to recognize these natural marmoset calls. We found that training did not induce call selectivity for the trained vocalizations in A1. However, although ferret A1 neurons were not call selective, they efficiently represented the vocalizations through temporal pattern codes, and trained animals recognized marmoset twitters with a high degree of accuracy. These temporal patterns needed to be analyzed at timescales of 10-50 ms to ensure efficient decoding. Training led to a substantial increase in the amount of information transmitted by these temporal discharge patterns, but the fundamental nature of the temporal pattern code remained unaltered. These results emphasize the importance of temporal discharge patterns and cast doubt on the functional significance of call-selective neurons in the processing of animal communication sounds at the level of A1.

Acoustic Stimulation↗

Response linearity in primary auditory cortex of the ferret.

The responses of neurons within the primary auditory cortex (A1) of the ferret elicited by broadband dynamic spectral ripple stimuli were examined over a range of ripple spectral densities and ripple velocities. The large majority of neurons showed modulated responses to these stimuli and responded most strongly at low ripple densities and velocities. The period histograms of their responses were subjected to Fourier analysis, and the ratio of the magnitudes of the f1 and fo (DC) components of these responses were calculated to give a quantitative index of response linearity. For 82 out of 396 neurons tested (20.7%) this ratio remained above 1.0 over the entire range of ripple densities and velocities. These neurons were classified as 'consistently linear'. A further 134/396 (33.8%) of neurons maintained an f1/f0 ratio above 1.0 for either a range of ripple densities at a fixed ripple velocity, or over a range of ripple velocities at a specific ripple density, and were classified as 'locally linear'. Interestingly, for the superficial layers of the primary auditory cortex, consistently linear and locally linear neurons outnumbered nonlinear neurons by a 2:1 ratio. The converse was true for the deep layers. Unlike in primary visual cortex, where f1/f0 ratios have been reported to exhibit a bimodal distribution with a minimum at f1/f0 = 1, f1/f0 ratios for A1 are unimodally distributed with a peak at f1/f0 = 1.

Acoustic Stimulation↗

Binaural-level functions in ferret auditory cortex: evidence for a continuous distribution of response properties.

Many previous studies have subdivided auditory neurons into a number of physiological classes according to various criteria applied to their binaural response properties. However, it is often unclear whether such classifications represent discrete classes of neurons or whether they merely reflect a potentially convenient but ultimately arbitrary partitioning of a continuous underlying distribution of response properties. In this study we recorded the binaural response properties of 310 units in the auditory cortex of anesthetized ferrets, using an extensive range of interaural level differences (ILDs) and average binaural levels (ABLs). Most recordings were from primary auditory fields on the middle ectosylvian gyrus and from neurons with characteristic frequencies >5 kHz. We used simple multivariate statistics to quantify a fundamental coding feature: the shapes of the binaural response functions. The shapes of all 310 binaural response surfaces were represented as points in a five-dimensional principal component space. This space captured the underlying shape of all the binaural response surfaces. The distribution of binaural level functions was not homogeneous because some shapes were more common than others. Despite this, clustering validation techniques revealed no evidence for the existence of discrete, or partially overlapping, clusters that could serve as a basis for an objective classification of binaural-level functions. We also examined the gradients of the response functions for the population of units; these gradients were greatest near the midline, which is consistent with free-field data showing that cortical neurons are most sensitive to changes in stimulus location in this region of space.

Acoustic Stimulation↗

Interaural timing cues do not contribute to the map of space in the ferret superior colliculus: a virtual acoustic space study.

In this study, we used individualized virtual acoustic space (VAS) stimuli to investigate the representation of auditory space in the superior colliculus (SC) of anesthetized ferrets. The VAS stimuli were generated by convolving broadband noise bursts with each animal's own head-related transfer function and presented over earphones. Comparison of the amplitude spectra of the free-field and VAS signals and of the spatial receptive fields of neurons recorded in the inferior colliculus with each form of stimulation confirmed that the VAS provided an accurate simulation of sounds presented in the free field. Units recorded in the deeper layers of the SC responded predominantly to virtual sound directions within the contralateral hemifield. In most cases, increasing the sound level resulted in stronger spike discharges and broader spatial receptive fields. However, the preferred sound directions, as defined by the direction of the centroid vector, remained largely unchanged across different levels and, as observed in previous free-field studies, varied topographically in azimuth along the rostrocaudal axis of the SC. We also examined the contribution of interaural time differences (ITDs) to map topography by digitally manipulating the VAS stimuli so that ITDs were held constant while allowing other spatial cues to vary naturally. The response properties of the majority of units, including centroid direction, remained unchanged with fixed ITDs, indicating that sensitivity to this cue is not responsible for tuning to different sound directions. These results are consistent with previous data suggesting that sensitivity to interaural level differences and spectral cues provides the basis for the map of auditory space in the mammalian SC.

Acoustic Stimulation↗

Encoding of virtual acoustic space stimuli by neurons in ferret primary auditory cortex.

Recent studies from our laboratory have indicated that the spatial response fields (SRFs) of neurons in the ferret primary auditory cortex (A1) with best frequencies > or =4 kHz may arise from a largely linear processing of binaural level and spectral localization cues. Here we extend this analysis to investigate how well the linear model can predict the SRFs of neurons with different binaural response properties and the manner in which SRFs change with increases in sound level. We also consider whether temporal features of the response (e.g., response latency) vary with sound direction and whether such variations can be explained by linear processing. In keeping with previous studies, we show that A1 SRFs, which we measured with individualized virtual acoustic space stimuli, expand and shift in direction with increasing sound level. We found that these changes are, in most cases, in good agreement with predictions from a linear threshold model. However, changes in spatial tuning with increasing sound level were generally less well predicted for neurons whose binaural frequency-time receptive field (FTRF) exhibited strong excitatory inputs from both ears than for those in which the binaural FTRF revealed either a predominantly inhibitory effect or no clear contribution from the ipsilateral ear. Finally, we found (in agreement with other authors) that many A1 neurons exhibit systematic response latency shifts as a function of sound-source direction, although these temporal details could usually not be predicted from the neuron's binaural FTRF.

Acoustic Stimulation↗

Encoding stimulus information by spike numbers and mean response time in primary auditory cortex.

Neurons can transmit information about sensory stimuli via their firing rate, spike latency, or by the occurrence of complex spike patterns. Identifying which aspects of the neural responses actually encode sensory information remains a fundamental question in neuroscience. Here we compared various approaches for estimating the information transmitted by neurons in auditory cortex in two very different experimental paradigms, one measuring spatial tuning and the other responses to complex natural stimuli. We demonstrate that, in both cases, spike counts and mean response times jointly carry essentially all the available information about the stimuli. Thus, in auditory cortex, whereas spike counts carry only partial information about stimulus identity or location, the additional availability of relatively coarse temporal information is sufficient in order to extract essentially all the sensory information available in the spike discharge pattern, at least for the relatively short stimuli (<approximately 100 ms) commonly used in auditory research.

Acoustic Stimulation↗

The detection of multisensory stimuli in an orthogonal sensory space.

The detection of a stimulus can be considerably facilitated if the stimulus engages two or more sensory modalities simultaneously. This phenomenon, commonly referred to as multisensory (or cross-modal) facilitation, has been demonstrated behaviorally in cats and humans. A number of rules are thought to govern this phenomenon. These rules state that strong facilitation is to be expected only if the two sensory modalities are stimulated simultaneously and at the same place, and if the stimuli themselves are weak. However, these rules are not sufficient to allow accurate predictions of multimodal stimulus detection probabilities directly from physical stimulus parameters. Here we show that such predictions are possible on the basis of a simple and biologically plausible psychophysical model, which relates the detection of audio-visual, audio-tactile or visual-tactile stimuli to the Euclidean distance that these stimuli span in an orthogonal sensory space.

Acoustic Stimulation↗

Large-scale organization of ferret auditory cortex revealed using continuous acquisition of intrinsic optical signals.

We have adapted a new approach for intrinsic optical imaging, in which images were acquired continuously while stimuli were delivered in a series of continually repeated sequences, to provide the first demonstration of the large-scale tonotopic organization of both primary and nonprimary areas of the ferret auditory cortex. Optical responses were collected during continuous stimulation by repeated sequences of sounds with varying frequency. The optical signal was averaged as a function of time during the sequence, to produce reflectance modulation functions (RMFs). We examined the stability and properties of the RMFs and show that their zero-crossing points provide the best temporal reference points for quantifying the relationship between the stimulus parameter values and optical responses. Sequences of different duration and direction of frequency change gave rise to comparable results, although in some cases discrepancies were observed, mostly between upward- and downward-frequency sequences. We demonstrated frequency maps, consistent with previous data, in primary auditory cortex and in the anterior auditory field, which were verified with electrophysiological recordings. In addition to these tonotopic gradients, we demonstrated at least 2 new acoustically responsive areas on the anterior and posterior ectosylvian gyri, which have not previously been described. Although responsive to pure tones, these areas exhibit less tonotopic order than the primary fields.

Acoustic Stimulation↗

Acoustic factors govern developmental sharpening of spatial tuning in the auditory cortex.

Auditory localization relies on the detection and interpretation of acoustic cues that change in value as the head and external ears grow. Here we show that the maturation of these structures is an important determinant for the development of spatial selectivity in the ferret auditory cortex. Spatial response fields (SRFs) of high-frequency cortical neurons recorded at postnatal days (P) 33-39 were broader, and transmitted less information about stimulus direction, than in older ferrets. They also exhibited slightly broader frequency tuning than neurons recorded in adult animals. However, when infant neurons were stimulated through virtual ears of adults, SRFs sharpened significantly and the amount of transmitted information increased. This improvement was predicted by a model that generates SRF shape from the localization cue values and the neurons' binaural spectrotemporal response properties. The maturation of spatial response characteristics in auditory cortex therefore seems to be limited by peripheral rather than by central factors.

Acoustic Stimulation↗

Modeling individual differences in ferret external ear transfer functions.

Individual variations in head and outer ear size, as well as growth of these structures during development, can markedly alter the values of the binaural and monaural cues which form the basis for auditory localization. This study investigated individual differences in the directional component of the head-related transfer function of both adult and juvenile ferrets. In line with previous studies in humans and cats, intersubject spectral differences were found to be reduced by scaling one of the directional transfer functions on a log-frequency axis. The optimal scale factor correlated most highly with pinna cavity height. Optimal frequency scaling reduced interear spectral difference equally well for adult-juvenile comparisons as for comparisons between pairs of adult ears. This illustrates that the developmental changes in localization cue values should be at least partly predictable on the basis of the expected growth rate of the outer ear structures. Predictions of interaural time differences (ITDs) were also derived from the physical dimensions of the head. ITDs were found to be poorly fitted by the spherical head model, while much better predictions could be derived from a model based on von Mises spherical basis functions. Together, these findings show how more accurate estimates of spatial cue values can be made from knowledge of the dimensions of the head and outer ears, and may facilitate the generation of virtual acoustic space stimuli in the absence of acoustical measurements from individual subjects.

Age Factors↗

Cortical plasticity: learning from cortical reorganisation.

Neocortical circuits can undergo dynamic rearrangements, not only in response to injury, but also when new skills are acquired. But although training can lead to functional rewiring of the cortex, we are far from being able to reprogram an animal by manipulating its cortical circuitry directly.

Acoustic Stimulation↗

Spectrotemporal receptive field properties of single units in the primary, dorsocaudal and ventrorostral auditory cortex of the guinea pig.

We report the spectrotemporal response properties of single units in the primary (A1) and dorsocaudal (DC) fields, and the ventrorostral belt of the urethane-anaesthetised guinea pig auditory cortex. Using reverse correlation analysis, spectrotemporal receptive fields (STRFs) were constructed and subsequently classified according to a novel qualitative scheme that was based on the duration and bandwidth of excitatory and inhibitory regions within the STRF. The STRFs of units in both A1 and DC showed either broad-band (> or = 1 octave) or narrow-band (< 1 octave) excitatory and inhibitory regions occurring either alone or together. The excitatory regions were of short duration (lasting for <50 ms) or more sustained (up to about 100 ms) and inhibitory areas either followed excitation or were located as inhibitory sidebands along the high- and low-frequency edges of the excitatory regions. Inhibitory areas that followed excitatory regions were found to be either short lasting (10-20 ms) or longer lasting (up to 200 ms or more). The STRFs recorded from each cortical area indicated temporal response properties consistent with those shown by traditional peristimulus time histogram analysis. Overall, fields A1 and DC showed no significant differences in the distribution of STRF types. Thus, it appears that both fields display similar spectrotemporal sensitivities to auditory stimuli and therefore, appear to process such stimuli in a parallel fashion. Single units recorded in the ventrorostral belt area showed STRF types similar to those recorded in A1 and DC. However, the proportions of STRF types were significantly different, suggesting a difference in spectrotemporal processing between the ventrorostral belt and the core areas.

Animals↗