Biomedical subjects

Alan R Palmer

Publications and source records attributed to Alan R Palmer.

11 recordsLinked to original sources

PubMed2003

Interaural time difference discrimination thresholds for single neurons in the inferior colliculus of Guinea pigs.

Sensitivity to changes in the interaural time difference (ITD) of 50 msec tones was measured in single units in the inferior colliculus of urethane-anesthetized guinea pigs. ITD functions were measured with 100 repeats and fine spacing (100 points per cycle). The just noticeable difference (jnd) for ITD was determined using receiver operating characteristic (ROC) analysis of the spike-count distribution at each ITD. The jnd became progressively smaller as the signal frequency increased from 50 to 800 Hz but became unmeasurable above 1 kHz. The lowest jnds (30 microsec) were comparable with human jnds, indicating that there is sufficient information in the firings of individual neurons to permit discrimination without obligatory pooling. ROC analysis requires the choice of a reference ITD from which the jnd may be found by stepping the target ITD through the ITD function. For each neuron the reference was chosen to minimize the jnd. The lowest jnd was usually for ipsilateral leading references, near the minimum of the ITD function where the variance was also low, but where the slope was nearing its steepest. This was despite the peak of the ITD function occurring for contralateral leading stimuli. When the reference ITD was on midline, a jnd could be obtained by looking for firing rates either greater or smaller than the firing rate at midline. The lower jnd was usually obtained by looking for a decrease in firing rate. As duration increased, jnds either decreased or increased, depending on unit type, whereas when level increased, jnds generally increased.

Animals↗

PubMed2003

The sound-level-dependent growth in the extent of fMRI activation in Heschl's gyrus is different for low- and high-frequency tones.

fMRI (functional magnetic resonance imaging) was used to investigate whether the growth in activation of the human auditory cortex, with increasing sound level, is discernibly different for high- and low-frequency tones. Ten volunteers were scanned whilst listening to sequences of low-frequency (0.30-kHz) tones at sound levels between 42 and 96 dB sound pressure level (SPL), and 10 whilst listening to high-frequency (4.75-kHz) tones at the same sound levels. Activation was measured in Heschl's gyrus (including primary auditory cortex) which has been shown to be most sensitive to changes in sound level. For the 0.30-kHz tone, the extent of activation was flat up to 66 dB and then showed a rapid growth which continued up to the highest level studied (96 dB SPL). In contrast, increasing the level of 4.75-kHz tones produced a steady growth in the extent of activation across the range of levels studied. These results are consistent with physiological evidence suggesting that recruitment of primary auditory cortical neurones may be different at high and low frequencies.

Acoustic Stimulation↗

PubMed2003

Amplitude and frequency-modulated stimuli activate common regions of human auditory cortex.

Hall et al. (Hall et al., 2002, Cerebral Cortex 12:140-149) recently showed that pulsed frequency-modulated tones generate considerably higher activation than their unmodulated counterparts in non-primary auditory regions immediately posterior and lateral to Heschl's gyrus (HG). Here, we use fMRI to explore the type of modulation necessary to evoke such differential activation. Carrier signals were a single tone and a harmonic-complex tone, with a 300 Hz fundamental, that were modulated at a rate of 5 Hz either in frequency, or in amplitude, to create six stimulus conditions (unmodulated, FM, AM). Relative to the silent baseline, the modulated tones, in particular, activated widespread regions of the auditory cortex bilaterally along the supra-temporal plane. When compared with the unmodulated tones, both AM and FM tones generated significantly greater activation in lateral HG and the planum temporale, replicating the previous findings. These activation patterns were largely overlapping, indicating a common sensitivity to both AM and FM. Direct comparisons between AM and FM revealed a higher magnitude of activation in response to the variation in amplitude than in frequency, plus a small part of the posterolateral region in the right hemisphere whose response was specifically AM-, and not FM-, dependent. The dominant pattern of activation was that of co-localized activation by AM and FM, which is consistent with a common neural code for AM and FM within these brain regions.

Acoustic Stimulation↗

PubMed2002

Histochemical identification of cortical areas in the auditory region of the human brain.

Despite numerous studies stretching over the last 100 years there is still no general agreement on the number of auditory areas in the human cortex or even how to define them by histological methods. Full definition of these areas will require a combination of functional and histological methods but, by using six complementary histological methods, of which most have been used in the monkey, we provide a clearer description of these areas. The primary auditory area was located on the posteromedial two-thirds of the first transverse temporal (Heschl's) gyrus and was distinguished by a dense band of cytochrome oxidase activity in layer IV and the base of layer III, as well as a relatively thick, pale layer V and VI. Layers V and VI together made up 40% of the cortical thickness. Acetylcholinesterase (AChE)-containing pyramidal cells were sparsely distributed within the primary auditory area. The anterolateral third of Heschl's gyrus did not have a clear band of high cytochrome oxidase activity but contained a moderately high density of AChE-containing pyramidal cells and thus appeared to be part of the auditory belt. Within Heschl's sulcus there was a third area, which had a band of high cytochrome oxidase activity and bands of high parvalbumin immunoreactivity and AChE activity in layer IV. This area appeared to be part of the auditory core. Thus the use of staining methods for cytochrome oxidase, AChE and parvalbumin provided additional information which allowed a clearer definition of auditory areas than Nissl or myelin staining alone. Our results suggest that there are two core areas surrounded by at least six belt areas in the human auditory region.

Acetylcholinesterase↗

PubMed2002

Blocking GABAergic inhibition increases sensitivity to sound motion cues in the inferior colliculus.

Responses of low-frequency neurons in the inferior colliculus (IC) of anesthetized guinea pigs were recorded to interaural phase modulation (IPM) before, during, and after iontophoresis of bicuculline, an antagonist to the inhibitory neurotransmitter GABA. Sensitivity to the direction of virtual motion resulting from IPM is an emergent property of neurons at the level of the IC. One model to account for this emergent sensitivity depends on GABAergic inhibition. Blocking GABAergic inhibition with bicuculline substantially increased neuronal discharge rates and increased the extent to which neurons were sensitive to the apparent-motion cues of IPM. The effect of GABA blockade is consistent with the hypothesis that sensitivity to the motion cues of IPM results from a process of adaptation-of-excitation whereby the magnitude of the recent response history influences subsequent neuronal responsiveness. These results indicate that GABAergic inhibition strongly influences the context-dependent processing of low-frequency binaural signals in the IC.

Acoustic Stimulation↗

PubMed2002

Interconnections of auditory areas in the guinea pig neocortex.

By studying the efferent projections of five auditory areas in the guinea pig cortex, we sought evidence that the larger fields can be divided into subareas based on unique patterns of cortical connections. Small extracellular injections of biocytin were made in combination with evoked potential mapping or single-unit analysis and histochemical determination of cortical landmarks. The two core fields, primary (AI) and dorsocaudal (DC), are partially surrounded by six adjacent belt areas, leaving two gaps: one at the rostral edge of AI and the other at the dorsal edge. All of the areas studied projected to their nearest neighbors, but AI was the only area to project to all seven of the other auditory areas. The caudal, high-frequency (more than 4 kHz) end of AI had different projections from the rostral, low-frequency (less than 1.5 kHz) end, and there was no evidence of connections between the two ends. Each end had separate dorsal and ventral projections. The two ends of AI may be working independently. By contrast, area DC had strong connections between its high- and low-frequency ends and it may be involved in auditory/visual integration. The dorsorostral belt (DRB) was subdivided into two zones on the basis of its projections: the more rostral part appears to overlap the second somatosensory area and be bimodal, while the caudal part has stronger auditory connections. The small belt area (area S) had separate physiological and anatomical properties from the rest of the rostral belt.

Animals↗

PubMed2002

Heschl's gyrus is more sensitive to tone level than non-primary auditory cortex.

Previous neuroimaging studies generally demonstrate a growth in the cortical response with an increase in sound level. However, the details of the shape and topographic location of such growth remain largely unknown. One limiting methodological factor has been the relatively sparse sampling of sound intensities. Additionally, most studies have either analysed the entire auditory cortex without differentiating primary and non-primary regions or have limited their analyses to Heschl's gyrus (HG). Here, we characterise the pattern of responses to a 300-Hz tone presented in 6-dB steps from 42 to 96 dB sound pressure level as a function of its sound level, within three anatomically defined auditory areas; the primary area, on HG, and two non-primary areas, consisting of a small area lateral to the axis of HG (the anterior lateral area, ALA) and the posterior part of auditory cortex (the planum temporale, PT). Extent and magnitude of auditory activation increased non-linearly with sound level. In HG, the extent and magnitude were more sensitive to increasing level than in ALA and PT. Thus, HG appears to have a larger involvement in sound-level processing than does ALA or PT.

Acoustic Stimulation↗

PubMed2002

Phase-locked responses to pure tones in the primary auditory cortex.

At the level of the brainstem, precise temporal information is essential for some aspects of binaural processing, while at the level of the cortex, rate and place mechanisms for neural coding seem to predominate. However, we now show that precise timing of steady-state responses to pure tones occurs in the primary auditory cortex (AI). Recordings were made from 163 multi-units in guinea pig AI. All units increased their firing rate in response to pure tones at 100 Hz and 46 (28%) gave sustained responses which were synchronised with the stimulus waveform (phase-locking). The phase-locking units were clustered together in columns. Phase-locking was generally strongest in layers III and IV but was also recorded in layers I, II and V. Good phase-locking was observed over a range of 60-250 Hz: some units (30%) were narrow band while others (37%) were low-pass (33% were not determined). Phase-locking strength was also influenced by sound level: some units showed monotonic increases in strength with level and others were non-monotonic. Ten of the units provided a good temporal representation of the fundamental frequency (270 Hz) of a guinea pig vocalisation (rumble) and may be involved in analysing communication calls.

Acoustic Stimulation↗

PubMed2002

Microelectrode and neuroimaging studies of central auditory function.

Imaging studies in humans are revealing parallels with the functional organisation of the auditory brain discovered in microelectrode studies in animals: the rate of amplitude modulation generating the strongest response declines systematically from the lower brain stem to the cortex; an increase in sound level induces a higher level and a greater extent of activity; spectra are represented tonotopically in multiple cortical areas. There are also differences: evidence of organisation reflecting the sound level of the stimulus is absent in animals, but has been found in humans. Additionally, imaging has revealed functional specialisations which have not (yet) been located in animals: areas that respond more strongly to sounds with stronger pitches and to sounds that move in space. Microelectrode studies suggest that vocalisations are represented by spatially distributed populations of neurones in secondary auditory areas. In humans, likewise, activation progressively more specific to speech is found as the search moves from primary to secondary to accessory areas.

Animals↗

PubMed2002

Spectral and temporal processing in human auditory cortex.

Hierarchical processing suggests that spectrally and temporally complex stimuli will evoke more activation than do simple stimuli, particularly in non-primary auditory fields. This hypothesis was tested using two tones, a single frequency tone and a harmonic tone, that were either static or frequency modulated to create four stimuli. We interpret the location of differences in activation by drawing comparisons between fMRI and human cytoarchitectonic data, reported in the same brain space. Harmonic tones produced more activation than single tones in right Heschl's gyrus (HG) and bilaterally in the lateral supratemporal plane (STP). Activation was also greater to frequency-modulated tones than to static tones in these areas, plus in left HG and bilaterally in an anterolateral part of the STP and the superior temporal sulcus. An elevated response magnitude to both frequency-modulated tones was found in the lateral portion of the primary area, and putatively in three surrounding non-primary regions on the lateral STP (one anterior and two posterior to HG). A focal site on the posterolateral STP showed an especially high response to the frequency-modulated harmonic tone. Our data highlight the involvement of both primary and lateral non-primary auditory regions.

Acoustic Stimulation↗

PubMedDate not supplied

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↗