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T M Shackleton

Publications and source records attributed to T M Shackleton.

14 recordsLinked to original sources
PubMed2003

Morphology of physiologically characterised ventral cochlear nucleus stellate cells.

Stellate cells within the ventral cochlear nucleus (VCN) are a diverse cell group that have been classified according to their size and morphology. Some of these stellate cell types constitute major projection neurones into the brainstem and directly into the inferior colliculus, while others are implicated in more local processing. It is still not clear whether a specific physiological profile is uniquely associated with each distinct type of stellate cell. To investigate such associations, we have analysed 23 units with a battery of physiological stimuli in vivo and then examined their shape and outputs following juxtacellular labelling with biocytin. Five physiologically identified groups of cells were filled. These formed two major response classes: onset cells and chopper cells. The two classes could be separated purely on morphological grounds. The onset cells had large somata, large symmetrical dendritic trees and profuse axonal branches that were restricted to the cochlear nucleus on one (On-L) or both sides (On-C) of the brainstem. The chopper cells had smaller, asymmetric, dendritic trees, which were either planar or marginal, had smaller somata and an output axon that left via the trapezoid body. We have confirmed profuse projections into the dorsal cochlear nucleus from all onset cells, and more focal projections from some members of all three groups of chopper cells.

Acoustic Stimulation↗
PubMed2001

The ability of inferior colliculus neurons to signal differences in interaural delay.

Sound localization in humans depends largely on interaural time delay (ITD). The ability to discriminate differences in ITD is highly accurate. ITD discrimination (Delta ITD) thresholds, under some circumstances, are as low as 10-20 micros. It has been assumed that thresholds this low could only be obtained if the outputs from many neurons were combined. Here we use Receiver Operating Characteristic analysis to compute neuronal Delta ITD thresholds from 53 cells in the inferior colliculus in guinea pigs. The Delta ITD thresholds of single neurons range from several hundreds of micros down to 20-30 micros. The lowest single-cell thresholds are comparable to human thresholds determined with similar stimuli. This finding suggests that the highly accurate sound localization of human observers is consistent with the resolution of single cells and need not reflect the combined activity of many neurons.

Animals↗
PubMed2000

Phase-locked responses to pure tones in guinea pig auditory cortex.

Phase-locked responses to pure tones are a characteristic of most auditory cells at the level of the brain stem and allow sophisticated analyses based on coincidence detection. Phase-locking to tones has not previously been shown at the level of the auditory cortex in single unit studies. We have now identified phase-locked responses in 10% of low-frequency (< 1 kHz) units in the ventrorostral belt, a strip of cortex immediately ventral to the primary auditory area. All of these units showed phase-locking in their response to binaural tone pips of 60-200 Hz and showed narrow band pass characteristics within this range.

Acoustic Stimulation↗
PubMed2000

Organisation of binaural interactions in the primary and dorsocaudal fields of the guinea pig auditory cortex.

This study investigated the nature and topography of binaural interactions in the primary auditory field (AI) and dorsocaudal field (DC) of the urethane anaesthetised guinea pig auditory cortex. Single and multi-units were classified by their responses to monaural and binaural stimulation. In both AI and DC, units displayed binaural facilitation, binaural inhibition, or a level dependent mixture of facilitation and inhibition. There was a significant difference in the distribution of binaural response types between the two fields. Facilitated units predominated in DC (facilitated: 58%; inhibited: 24%; mixed: 6%; non-interacting: 12%), while inhibited units were the most common class in AI (facilitated: 15%; inhibited: 44%; mixed: 18%; non-interacting: 22%). It has previously been suggested that inhibited and facilitated units are concerned with processing different areas of space suggesting a possible separation of function between the two core fields. Topographically, the binaural response properties in AI and DC varied along isofrequency bands, with neurones displaying similar interactions aggregating in clusters. These clusters were similar in size for the two fields and often overlapped neighbouring isofrequency bands. However, their shape and position varied between different animals. This clustered organisation of binaural interactions is similar to that reported in recent studies of AI in other mammals.

Acoustic Stimulation↗
PubMed2000

Modelling convergent input onto interaural-delay-sensitive inferior colliculus neurones.

Convergent input from cells in the medial superior olive (MSO) and lateral superior olive (LSO) onto a single inferior colliculus (IC) cell explains many findings that are not compatible with a simple coincidence detector mechanism. Here this explanation is tested using a physiologically accurate computer model of the binaural pathway in which the input to the IC cell is either from two MSO cells or a MSO and a LSO cell. Auditory nerve (AN) spike trains are formed by a stochastic hair cell model following a basilar membrane simulation using a gammatone filter. In subsequent cells input spikes cause post-synaptic potentials (PSPs) which are summed causing the cell to fire when the sum crosses a threshold. The individual cells are matched to the physiology by varying the number of inputs, the magnitude and duration of the PSPs and the firing threshold. Non-linear best-phase-versus-frequency functions arise if the two IC inputs have different best frequencies and different characteristic delays. One input can be selectively suppressed by turning on an additional tone at the worst phase of that input. Non-zero characteristic phases arise if the characteristic frequencies of the AN fibres feeding into a single superior olive cell are mismatched.

Ear↗
PubMed2000

Responses of neurons in the inferior colliculus to dynamic interaural phase cues: evidence for a mechanism of binaural adaptation.

Responses to sound stimuli that humans perceive as moving were obtained for 89 neurons in the inferior colliculus (IC) of urethan-anesthetized guinea pigs. Triangular and sinusoidal interaural phase modulation (IPM), which produced dynamically varying interaural phase disparities (IPDs), was used to present stimuli with different depths, directions, centers, and rates of apparent motion. Many neurons appeared sensitive to dynamic IPDs, with responses at any given IPD depending strongly on the IPDs the stimulus had just passed through. However, it was the temporal pattern of the response, rather than the motion cues in the IPM, that determined sensitivity to features such as motion depth, direction, and center locus. IPM restricted only to the center of the IPD responsive area, evoked lower discharge rates than when the stimulus either moved through the IPD responsive area from outside, or up and down its flanks. When the stimulus was moved through the response area first in one direction and then back in the other, and the same IPDs evoked different responses, the response to the motion away from the center of the IPD responsive area was always lower than the response to the motion toward the center. When the IPD was closer at which the direction of motion reversed was to the center, the response to the following motion was lower. In no case did we find any evidence for neurons that under all conditions preferred one direction of motion to the other. We conclude that responses of IC neurons to IPM stimuli depend not on the history of stimulation, per se, but on the history of their response to stimulation, irrespective of the specific motion cues that evoke those responses. These data are consistent with the involvement of an adaptation mechanism that resides at or above the level of binaural integration. We conclude that our data provide no evidence for specialized motion detection involving dynamic IPD cues in the auditory midbrain of the mammal.

Acoustic Stimulation↗
PubMed1998

Convergent input from brainstem coincidence detectors onto delay-sensitive neurons in the inferior colliculus.

Responses of low-frequency neurons in the inferior colliculus (IC) of anesthetized guinea pigs were studied with binaural beats to assess their mean best interaural phase (BP) to a range of stimulating frequencies. Phase plots (stimulating frequency vs BP) were produced, from which measures of characteristic delay (CD) and characteristic phase (CP) for each neuron were obtained. The CD provides an estimate of the difference in travel time from each ear to coincidence-detector neurons in the brainstem. The CP indicates the mechanism underpinning the coincidence detector responses. A linear phase plot indicates a single, constant delay between the coincidence-detector inputs from the two ears. In more than half (54 of 90) of the neurons, the phase plot was not linear. We hypothesized that neurons with nonlinear phase plots received convergent input from brainstem coincidence detectors with different CDs. Presentation of a second tone with a fixed, unfavorable delay suppressed the response of one input, linearizing the phase plot and revealing other inputs to be relatively simple coincidence detectors. For some neurons with highly complex phase plots, the suppressor tone altered BP values, but did not resolve the nature of the inputs. For neurons with linear phase plots, the suppressor tone either completely abolished their responses or reduced their discharge rate with no change in BP. By selectively suppressing inputs with a second tone, we are able to reveal the nature of underlying binaural inputs to IC neurons, confirming the hypothesis that the complex phase plots of many IC neurons are a result of convergence from simple brainstem coincidence detectors.

Acoustic Stimulation↗
PubMed1994

Perceived contrast of gratings and plaids: non-linear summation across oriented filters.

Plaids composed of two orthogonal sine-wave gratings appeared to be of lower contrast than single gratings of the same Michelson luminance contrast. This effect for plaids was obtained at all spatial frequencies (1-16 c/deg) and contrast levels (2-32%). Contrast-matching data plotted as a function of the angle between plaid components (0-90 deg) and as a function of spatial frequency and standard contrast level were consistent with a model in which the response of each orientation-tuned spatial frequency channel is a threshold-corrected power function of contrast, and is followed by quadratic summation of responses across all channels. The best-fitting contrast-response exponent in the main experiment was 0.63. Analysis of several other data-sets suggested a slightly higher value, 0.80. The same model gave a good account of contrast-matching between simple and compound (two-component) one dimensional gratings, accounting in particular for the apparent increase in contrast summation exponent at low contrasts reported by Quick, Hamerly and Reichert [(1976) Vision Research, 16, 351-355]. The model can, with one further assumption, account for the finding that contrast-matching between sine-wave and square-wave gratings depended only on the amplitude at the fundamental frequency. Comparison with contrast discrimination studies suggests that internal noise (variance of a channel's contrast-response) is not constant, but increases approximately in proportion to the mean response.

Contrast Sensitivity↗
PubMed1994

The role of resolved and unresolved harmonics in pitch perception and frequency modulation discrimination.

A series of experiments investigated the influence of harmonic resolvability on the pitch of, and the discriminability of differences in fundamental frequency (F0) between, frequency-modulated (FM) harmonic complexes. Both F0 (62.5 to 250 Hz) and spectral region (LOW: 125-625 Hz, MID: 1375-1875 Hz, and HIGH: 3900-5400 Hz) were varied orthogonally. The harmonics that comprised each complex could be summed in either sine (0 degree) phase (SINE) or alternating sine-cosine (0 degree-90 degrees) phase (ALT). Stimuli were presented in a continuous pink-noise background. Pitch-matching experiments revealed that the pitch of ALT-phase stimuli, relative to SINE-phase stimuli, was increased by an octave in the HIGH region, for all F0's, but was the same as that of SINE-phase stimuli when presented in the LOW region. In the MID region, the pitch of ALT-phase relative to SINE-phase stimuli depended on F0, being an octave higher at low F0's, equal at high F0's, and unclear at intermediate F0's. The same stimuli were then used in three measures of discriminability: FM detection thresholds (FMTs), frequency difference limens (FDLs), and FM direction discrimination thresholds (FMDDTs, defined as the minimum FM depth necessary for listeners to discriminate between two complexes modulated 180 degrees out of phase with each other). For all three measures, at all F0's, thresholds were low (< 4% for FMTs, < 5% for FMDDTs, and < 1.5% for FDLs) when stimuli were presented in the LOW region, and high (> 10% for FMTs, > 7% for FMDDTs, and > 2.5% for FDLs) when presented in the HIGH region. When stimuli were presented in the MID region, thresholds were low for low F0's, and high for high F0's. Performance was not markedly affected by the phase relationship between the components of a complex, except for stimuli with intermediate F0's in the MID spectral region, where FDLs and FMDDTs were much higher for ALT-phase stimuli than for SINE-phase stimuli, consistent with their unclear pitch. This difference was much smaller when FMTs were measured. The interaction between F0 and spectral region for both sets of experiments can be accounted for by a single definition of resolvability.

Acoustic Stimulation↗
PubMed1992

No evidence for dichoptic motion sensing: a reply to Carney and Shadlen.

Georgeson and Shackleton (1989, Vision Research, 29, 1511-1523) confirmed the existence of dichoptic apparent motion (AM), but argued that its basis was the spatio-temporal correspondence of visible features ("feature tracking"), not early motion sensors. Direction selectivity in motion sensors was probably purely monocular. The key evidence came from AM of missing-fundamental (MF) gratings. Monocular MF motion was always reversed, implying motion sensors responding to the third harmonic (3f) component. Dichoptic MF motion was in the correct direction at the higher contrasts and lower drift rates, and overall was highly correlated with judgements of pattern structure, suggesting feature-tracking. Carney and Shadlen (1992, Vision Research, 32, 187-191) criticized some of our methodology and the theoretical interpretation. Their central argument was that dichoptic AM for sine-waves did reflect dichoptic motion sensing, but was less reliable at higher contrasts. Hence forward motion of the dichoptic MF pattern should occur only where sine-wave (3f) motion-sensing declines. We discuss their critique, and find little support for it. We also present additional data on dichoptic AM for 3f and 5f gratings, showing that Carney and Shadlen's predictions were not upheld. Feature tracking remains the most plausible account of dichoptic AM.

Contrast Sensitivity↗
PubMed1992

A computer model of a cochlear-nucleus stellate cell: responses to amplitude-modulated and pure-tone stimuli.

A computer model of a ventral-cochlear-nucleus (VCN) stellate cell with chop-S type response properties is presented and evaluated. The model is based on a simplified model of spike generation preceded by a stage that simulates dendritic low-pass filtering. Input to the model is in the form of simulated auditory-nerve spikes produced by a model of the auditory periphery [Meddis and Hewitt, J. Acoust. Soc. Am. 89, 2866-2882 (1991)]. Outputs from the stellate-cell model are shown to qualitatively replicate a wide range of typical in vivo responses. These include: (a) realistic onset and steady-state rate-level functions, (b) "chopper"-type post-stimulus time histogram responses; (c) typical "chop-S"-type neuron responses characterized by a low coefficient of variance (CV less than 0.3) of interspike intervals as a function of time; (d) level-dependent amplitude-modulation transfer functions; (e) intrinsic oscillations in responses to pure-tone stimuli; (f) amplitude-modulation encoding over a wide dynamic range; and (g) frequency-limited phase locking to pure tones. It is shown that these responses can be explained primarily by the membrane properties of the cells. More specifically, how the model encodes signal amplitude modulation was studied and an explanation was suggested for the generation of the bandpass modulation transfer functions. Such functions are observed neurally in response to amplitude-modulated stimuli presented at moderate to high signal levels.

Animals↗
PubMed1991

Lateralization of very-short-duration tone pulses of low and high frequencies.

The position and image-width of the simultaneous images produced by very short tone pulses were measured as a function of interaural time difference (ITD) at both low- (250 and 800 Hz) and high- (2500 and 8000 Hz) frequencies using a direct-estimation technique. Primary images are lateralized towards the ear receiving the leading stimulus. At low frequencies image position is proportional to interaural phase-difference (IPD) below 90 degrees and remains at the lead-ear for larger values. At high frequencies images are reported is proportional to ITD up to 500-1000 microseconds. Secondary images are reported on the opposite side of the head for IPDs greater than 180 degrees at low frequencies, and at ITDs greater than 500 microseconds at high frequencies. Image width is approximately constant for all ITDs and both images at a given frequency, but becomes more compact as frequency increases. The data are discussed in terms of onset cues and stimulus fine-structure cues. The best explanation is in terms of an onset mechanism, but one that is calibrated in terms of IPD at low frequencies. The existence of double images is explained in terms of a breakdown in the mechanism determining fusion.

Adult↗
PubMed1989

Monocular motion sensing, binocular motion perception.

The two-process account of motion perception and its binocular organization were addressed in experiments on apparent movement (AM) with three types of grating: sinusoidal; random bar width; and square-wave with missing fundamental (MF). Monocular MF gratings sampled four times per cycle of drift always appeared to move backwards. Here AM was unrelated to the spatial appearance of the pattern, and followed the motion of the dominant spatial frequency component (the third harmonic). We take this reversed AM to be characteristic of "short-range" motion sensors. It did not occur dichoptically, implying that the direction-selective mechanism of motion sensors is purely monocular. AM was seen with dichoptic presentation for all three types of grating. Performance improved with the length of the stimulus sequence, as predicted by probability summation. This result reconciles previous positive and negative findings on dichoptic AM. The perceived direction of dichoptic AM was consistent with polarity-selective matching of features over time (the "long-range process"). The most telling effect supporting feature-matching in dichoptic motion was that dichoptic MF motion reversed direction with a change in the visible features of the pattern (induced by changes in contrast and pulse duration); monocular apparent motion did not. Two routes from spatial frequency channels to the perception of object motion are discussed.

Contrast Sensitivity↗