Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. I. Evidence for monaural directional cues

1993 ◽  
Vol 70 (2) ◽  
pp. 492-511 ◽  
Author(s):  
F. K. Samson ◽  
J. C. Clarey ◽  
P. Barone ◽  
T. J. Imig

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Neurons, sensitive to sound direction in the horizontal plane (azimuth), were identified by their responses to noise bursts, presented in the free field, that varied in azimuth and sound pressure level (SPL). SPLs typically varied between 0 and 80 dB and were presented at each azimuth that was tested. Each azimuth-sensitive neuron responded well to some SPLs at certain azimuths and did not respond well to any SPL at other azimuths. This report describes AI neurons that were sensitive to the azimuth of monaurally presented noise bursts. 2. Unilateral ear plugging was used to test each azimuth-sensitive neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the concha and ear canal, attenuated sound reaching the tympanic membrane by 25-70 dB. Binaural interactions were inferred by comparing responses obtained under binaural (no plug) and monaural (ear plug) conditions. 3. Of the total sample of 131 azimuth-sensitive cells whose responses to ear plugging were studied, 27 were sensitive to the azimuth of monaurally presented noise bursts. We refer to these as monaural directional (MD) cells, and this report describes their properties. The remainder of the sample consisted of cells that either required binaural stimulation for azimuth sensitivity (63/131), because they were insensitive to azimuth under unilateral ear plug conditions or responded too unreliably to permit detailed conclusions regarding the effect of ear plugging (41/131). 4. Most (25/27) MD cells received either monaural input (MD-E0) or binaural excitatory/inhibitory input (MD-EI), as inferred from ear plugging. Two MD cells showed other characteristics. The contralateral ear was excitatory for 25/27 MD cells. 5. MD-E0 cells (22%, 6/27) were monaural. They were unaffected by unilateral ear plugging, showing that they received excitatory input from one ear, and that stimulation of the other ear was without apparent effect. On the other hand, some monaural cells in AI were insensitive to the azimuth of noise bursts, showing that sensitivity to monaural directional cues is not a property of all monaural cells in AI. 6. MD-EI cells (70%, 19/27) exhibited an increase in responsiveness on the side of the plugged ear, showing that they received excitatory drive from one ear and inhibitory drive from the other. MD-EI cells remained azimuth sensitive with the inhibitory ear plugged, showing that they were sensitive to monaural directional cues at the excitatory ear.(ABSTRACT TRUNCATED AT 400 WORDS)

1994 ◽  
Vol 71 (6) ◽  
pp. 2194-2216 ◽  
Author(s):  
F. K. Samson ◽  
P. Barone ◽  
J. C. Clarey ◽  
T. J. Imig

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Observations were based on a sample of 131 high-best-frequency (> 5 kHz), azimuth-sensitive neurons. These were identified by their responses to a set of noise bursts, presented in the free field, that varied in azimuth and sound-pressure level (SPL). Each azimuth-sensitive neuron responded well to some levels at certain azimuths, but did not respond well to any level at other azimuths. 2. Unilateral ear plugging was used to infer each neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the external ear, attenuated sound reaching the tympanic membrane by 25–70 dB. The azimuth tuning of a large proportion of the sample (62/131), referred to as binaural directional (BD), was completely dependent upon binaural stimulation because with one ear plugged, these cells were insensitive to azimuth (either responded well at all azimuths or failed to respond at any azimuth) or in a few cases exhibited striking changes in location of azimuth function peaks. This report describes patterns of monaural responses and binaural interactions exhibited by BD neurons and relates them to each cell's azimuth and level tuning. The response of BD cells to ear plugging is consistent with the hypothesis that they derive azimuth tuning from interaural level differences present in noise bursts. Another component of the sample consisted of monaural directional (27/131) cells that derived azimuth tuning in part or entirely from monaural spectral cues. Cells in the remaining portion of the sample (42/131) responded too unreliably to permit specific conclusions. 3. Binaural interactions were inferred by statistical comparison of a cell's responses to monaural (unilateral plug) and binaural (no plug) stimulation. A larger binaural response than either monaural response was taken as evidence for binaural facilitation. A smaller binaural than monaural response was taken as evidence for binaural inhibition. Binaural facilitation was exhibited by 65% (40/62) of the BD sample (facilitatory cells). Many of these exhibited mixed interactions, i.e., binaural facilitation occurred in response to some azimuth-level combinations, and binaural inhibition to others. Binaural inhibition in the absence of binaural facilitation occurred in 35% (22/62) of the BD sample, a majority of which were EI cells, so called because they received excitatory (E) input from one ear (excitatory ear) and inhibitory (I) input from the other (inhibitory ear). One cell that exhibited binaural inhibition received excitatory input from each ear.(ABSTRACT TRUNCATED AT 400 WORDS)


1990 ◽  
Vol 63 (6) ◽  
pp. 1448-1466 ◽  
Author(s):  
T. J. Imig ◽  
W. A. Irons ◽  
F. R. Samson

1. The azimuth and sound pressure level (SPL) selectivities of single-unit responses recorded in primary auditory cortex of barbiturate-anesthetized cats were studied by the use of broadband noise bursts delivered in the free field from a moveable loud-speaker. The experiments were carried out with cats located inside a quasianechoic sound-isolation chamber. We studied 71 units with relatively stable response properties. All units were located in the frequency representation between 5.8 and 31 kHz. The data obtained for each unit were displayed as an azimuth-level response area, a contour plot that displays the distribution of response magnitude as a joint function of SPL and azimuth at 0 degrees elevation. From these, azimuth and level functions were obtained to derive descriptors of azimuth and level selectivity. 2. Sensitivity to sound-source azimuth was assessed from the modulation of the average azimuth function (average of azimuth functions obtained to each SPL of noise that was presented) for each unit. The sample was arbitrarily divided into a high-directionality (HD) group (66%) whose average azimuth functions had modulation values of greater than or equal to 75% and a low-directionality (LD) group (34%). The distinction between HD and LD groups was made so that we could analyze the characteristics of units likely to be involved in the representation of sound-source azimuth. 3. There is an overrepresentation of the contralateral sound field and the midline in the sample of HD units. The preferred sector for each unit was defined as the range of azimuths within the frontal sound field throughout which unit response was greater than or equal to 75% of maximum. Each unit was classified as either midline preferring (17%, the midpoint of the preferred sector, i.e., best azimuth, was located within 5 degrees of the midline), contralateral preferring (60%), or ipsilateral preferring (23%). The ratio of contralateral- to ipsilateral-preferring units was 2.5:1. A higher proportion of units had best azimuths located in the 10 degrees sector centered on the midline than in any other 10 degrees sector of the frontal sound field. 4. In one animal, recordings were obtained at seven closely spaced sites in layer IV from single- and multiunit responses, which were narrowly tuned to both azimuth and SPL. The units located along a 1-mm length of an isofrequency strip were tuned to similar frequencies and SPLs but had five distinctly different directional preferences distributed throughout the entire frontal sound field.(ABSTRACT TRUNCATED AT 400 WORDS)


1996 ◽  
Vol 75 (1) ◽  
pp. 75-96 ◽  
Author(s):  
D. R. Irvine ◽  
R. Rajan ◽  
L. M. Aitkin

1. Interaural intensity differences (IIDs) provide the major cue to the azimuthal location of high-frequency narrowband sounds. In recent studies of the azimuthal sensitivity of high-frequency neurons in the primary auditory cortex (field AI) of the cat, a number of different types of azimuthal sensitivity have been described and the azimuthal sensitivity of many neurons was found to vary as a function of changes in stimulus intensity. The extent to which the shape and the intensity dependence of the azimuthal sensitivity of AI neurons reflects features of their IID sensitivity was investigated by obtaining data on IID sensitivity from a large sample of neurons with a characteristic frequency (CF) > 5.5 kHz in AI of anesthetized cats. IID sensitivity functions were classified in a manner that facilitated comparison with previously obtained data on azimuthal sensitivity, and the effects of changes in the base intensity at which IIDs were introduced were examined. 2. IID sensitivity functions for CF tonal stimuli were obtained at one or more intensities for a total of 294 neurons, in most cases by a method of generating IIDs that kept the average binaural intensity (ABI) of the stimuli at the two ears constant. In the standard ABI range at which a function was obtained for each unit, five types of IID sensitivity were distinguished. Contra-max neurons (50% of the sample) had maximum response (a peak or a plateau) at IIDs corresponding to contralateral azimuths, whereas ipsi-max neurons (17%) had the mirror-image form of sensitivity. Near-zero-max neurons (18%) had a clearly defined maximum response (peak) in the range of +/- 10 dB IID, whereas a small group of tough neurons (2%) had a restricted range of minimal responsiveness with near-maximal responses at IIDs on either side. A final 18% of AI neurons were classified as insensitive to IIDs. The proportions of neurons exhibiting the various types of sensitivity corresponded closely to the proportions found to exhibit corresponding types of azimuthal sensitivity in a previous study. 3. There was a strong correlation between a neuron's binaural interaction characteristics and the form of its IID sensitivity function. Thus, neurons excited by monaural stimulation of only one ear but with either inhibitory, facilitatory, or mixed facilitatory-inhibitory effects of stimulation of the other ear had predominantly contra-max IID sensitivity (if contralateral monaural stimulation was excitatory) or ipsi-max sensitivity (if ipsilateral monaural stimulation was excitatory). Neurons driven weakly or not at all by monaural stimulation but facilitated binaurally almost all exhibited near-zero-max IID sensitivity. The exception to this tight association between binaural input and IID sensitivity was provided by neurons excited by monaural stimulation of either ear (EE neurons). Although EE neurons have frequently been considered to be insensitive to IIDs, our data were in agreement with two recent reports indicating that they can exhibit various forms of IID sensitivity: only 23 of 75 EE neurons were classified as insensitive and the remainder exhibited diverse types of sensitivity. 4. IID sensitivity was examined at two or more intensities (3-5 in most cases) for 84 neurons. The form of the IID sensitivity function (defined in terms of both shape and position along the IID axis) was invariant with changes in ABI for only a small proportion of IID-sensitive neurons (approximately 15% if a strict criterion of invariance was employed), and for many of these neurons the spike counts associated with a given IID varied with ABI, particularly at near-threshold levels. When the patterns of variation in the form of IID sensitivity produced by changes in ABI were classified in a manner equivalent to that used previously to classify the effects of intensity on azimuthal sensitivity, there was a close correspondence between the effects of intensity on corresponding types of azimuthal and IID sensitivity


2019 ◽  
Vol 9 (1) ◽  
Author(s):  
Hiroaki Tsukano ◽  
Xubin Hou ◽  
Masao Horie ◽  
Hiroki Kitaura ◽  
Nana Nishio ◽  
...  

AbstractRecent studies have examined the feedback pathway from the amygdala to the auditory cortex in conjunction with the feedforward pathway from the auditory cortex to the amygdala. However, these connections have not been fully characterized. Here, to visualize the comprehensive connectivity between the auditory cortex and amygdala, we injected cholera toxin subunit b (CTB), a bidirectional tracer, into multiple subfields in the mouse auditory cortex after identifying the location of these subfields using flavoprotein fluorescence imaging. After injecting CTB into the secondary auditory field (A2), we found densely innervated CTB-positive axon terminals that were mainly located in the lateral amygdala (La), and slight innervations in other divisions such as the basal amygdala. Moreover, we found a large number of retrogradely-stained CTB-positive neurons in La after injecting CTB into A2. When injecting CTB into the primary auditory cortex (A1), a small number of CTB-positive neurons and axons were visualized in the amygdala. Finally, we found a near complete absence of connections between the other auditory cortical fields and the amygdala. These data suggest that reciprocal connections between A2 and La are main conduits for communication between the auditory cortex and amygdala in mice.


2013 ◽  
Vol 110 (9) ◽  
pp. 2140-2151 ◽  
Author(s):  
Justin D. Yao ◽  
Peter Bremen ◽  
John C. Middlebrooks

The rat is a widely used species for study of the auditory system. Psychophysical results from rats have shown an inability to discriminate sound source locations within a lateral hemifield, despite showing fairly sharp near-midline acuity. We tested the hypothesis that those characteristics of the rat's sound localization psychophysics are evident in the characteristics of spatial sensitivity of its cortical neurons. In addition, we sought quantitative descriptions of in vivo spatial sensitivity of cortical neurons that would support development of an in vitro experimental model to study cortical mechanisms of spatial hearing. We assessed the spatial sensitivity of single- and multiple-neuron responses in the primary auditory cortex (A1) of urethane-anesthetized rats. Free-field noise bursts were varied throughout 360° of azimuth in the horizontal plane at sound levels from 10 to 40 dB above neural thresholds. All neurons encountered in A1 displayed contralateral-hemifield spatial tuning in that they responded strongly to contralateral sound source locations, their responses cut off sharply for locations near the frontal midline, and they showed weak or no responses to ipsilateral sources. Spatial tuning was quite stable across a 30-dB range of sound levels. Consistent with rat psychophysical results, a linear discriminator analysis of spike counts exhibited high spatial acuity for near-midline sounds and poor discrimination for off-midline locations. Hemifield spatial tuning is the most common pattern across all mammals tested previously. The homogeneous population of neurons in rat area A1 will make an excellent system for study of the mechanisms underlying that pattern.


2019 ◽  
Author(s):  
Jong Hoon Lee ◽  
Xiaoqin Wang ◽  
Daniel Bendor

AbstractIn primary auditory cortex, slowly repeated acoustic events are represented temporally by phase-locked activity of single neurons. Single-unit studies in awake marmosets (Callithrix jacchus) have shown that a sub-population of these neurons also monotonically increase or decrease their average discharge rate during stimulus presentation for higher repetition rates. Building on a computational single-neuron model that generates phase-locked responses with stimulus evoked excitation followed by strong inhibition, we find that stimulus-evoked short-term depression is sufficient to produce synchronized monotonic positive and negative responses to slowly repeated stimuli. By exploring model robustness and comparing it to other models for adaptation to such stimuli, we conclude that short-term depression best explains our observations in single-unit recordings in awake marmosets. Using this model, we emulated how single neurons could encode and decode multiple aspects of an acoustic stimuli with the monotonic positive and negative encoding of a given stimulus feature. Together, our results show that a simple biophysical mechanism in single neurons can allow a more complex encoding and decoding of acoustic stimuli.


1983 ◽  
Vol 50 (5) ◽  
pp. 1182-1196 ◽  
Author(s):  
A. Asanuma ◽  
D. Wong ◽  
N. Suga

The orientation sound emitted by the Panamanian mustached bat, Pteronotus parnellii rubiginosus, consists of four harmonics. The third harmonic is 6-12 dB weaker than the predominant second harmonic and consists of a long constant-frequency component (CF3) at about 92 kHz and a short frequency-modulated component (FM3) sweeping from about 92 to 74 kHz. Our primary aim is to examine how CF3 and FM3 are represented in a region of the primary auditory cortex anterior to the Doppler-shifted constant-frequency (DSCF) area. Extracellular recordings of neuronal responses from the unanesthetized animal were obtained during free-field stimulation of the ears with pure tones. FM sounds, and signals simulating their orientation sounds and echoes. Response properties of neurons and tonotopic and amplitopic representations were examined in the primary and the anteroventral nonprimary auditory cortex. In the anterior primary auditory cortex, neurons responded strongly to single pure tones but showed no facilitative responses to paired stimuli. Neurons with best frequencies from 110 to 90 kHz were tonotopically organized rostrocaudally, with higher frequencies located more rostrally. Neurons tuned to 92-94 kHz were overpresented, whereas neurons tuned to sound between 64 and 91 kHz were rarely found. Consequently a striking discontinuity in frequency representation from 91 to 64 kHz was found across the anterior DSCF border. Most neurons exhibited monotonic impulse-count functions and responded maximally to sound pressure level (SPL). There were also neurons that responded best to weak sounds but unlike the DSCF area, amplitopic representation was not found. Thus, the DSCF area is quite unique not only in its extensive representation of frequencies in the second harmonic CF component but also in its amplitopic representation. The anteroventral nonprimary auditory cortex consisted of neurons broadly tuned to pure tones between 88 and 99 kHz. Neither tonotopic nor amplitopic representation was observed. Caudal to this area and near the anteroventral border of the DSCF area, a small cluster of FM-FM neurons sensitive to particular echo delays was identified. The responses of these neurons fluctuated significantly during repetitive stimulation.


2018 ◽  
Vol 29 (9) ◽  
pp. 3796-3812 ◽  
Author(s):  
Feixue Liang ◽  
Haifu Li ◽  
Xiao-lin Chou ◽  
Mu Zhou ◽  
Nicole K Zhang ◽  
...  

Abstract Sparse representation is considered an important coding strategy for cortical processing in various sensory modalities. It remains unclear how cortical sparseness arises and is being regulated. Here, unbiased recordings from primary auditory cortex of awake adult mice revealed salient sparseness in layer (L)2/3, with a majority of excitatory neurons exhibiting no increased spiking in response to each of sound types tested. Sparse representation was not observed in parvalbumin (PV) inhibitory neurons. The nonresponding neurons did receive auditory-evoked synaptic inputs, marked by weaker excitation and lower excitation/inhibition (E/I) ratios than responding cells. Sparse representation arises during development in an experience-dependent manner, accompanied by differential changes of excitatory input strength and a transition from unimodal to bimodal distribution of E/I ratios. Sparseness level could be reduced by suppressing PV or L1 inhibitory neurons. Thus, sparse representation may be dynamically regulated via modulating E/I balance, optimizing cortical representation of the external sensory world.


1995 ◽  
Vol 74 (5) ◽  
pp. 1841-1855 ◽  
Author(s):  
D. M. Bowman ◽  
J. J. Eggermont ◽  
G. M. Smith

1. Neural activity was recorded extracellularly with two independent microelectrodes aligned in parallel and advanced perpendicular to isofrequency sheets in cat primary auditory cortex. Multiunit activity was separated into single-unit spike trains using a maximum variance spike sorting algorithm. Only units that demonstrated a high quality of sorting and a minimum spontaneous firing rate of 0.2 spikes/s were considered for analysis. The primary aim of this study was to describe the effect of periodic click train and broadband noise stimulation on short-time-scale (< or = 50 ms) bursts in the spike trains of single auditory cortical units and to determine whether stimulation influenced the occurrence, spike count, and/or temporal structure of burst firing relative to a spontaneous baseline. 2. Extracellular recordings were made in 20 juvenile and adult cats from 69 single auditory cortical units during click train stimulation and silence, and from 30 single units during noise stimulation and in silence. In an additional 15 single units the effect of both click train and noise stimulation was investigated. The incidence, spike count, and temporal structure of short-time-scale burst firing in the first 100 ms following stimulus presentation was compared with burst firing in the period starting 500 ms after stimulus presentation and with spontaneous burst firing. In addition, the serial dependence of interspike intervals within a burst was tested during periods of stimulation. 3. Burst firing was present in the stimulation, poststimulation, and spontaneous conditions. Longer bursts (consisting of > or = 3 spikes) were more commonly observed in the poststimulation and spontaneous conditions than in the stimulation condition. This effect was most pronounced during click stimulation. A period of elevated firing activity was present in a subset of units 0.5-1.5 s after stimulus presentation, indicating prolonged effects of stimulation on single-unit firing behavior. 4. For both stimuli, the proportion of single-unit responses composed of bursts was significantly greater in poststimulation and spontaneous periods than during stimulation. Burst rate was higher in post-click-train stimulation and spontaneous periods than during periods of click stimulation. The isolated spike rate was significantly higher during periods of noise and click stimulation than in the poststimulation and spontaneous periods. 5. An examination of the autocorrelograms and higher-order interspike interval histograms of single-unit responses during click train stimulation indicated that 25% of single-unit spike trains contained an excess of brief first-order intervals and 14% of spike trains contained a shortage of long higher-order interspike intervals relative to a spontaneous baseline. During noise stimulation, 10% of single-unit responses contained an excess of short intervals relative to baseline. Interspike intervals of short-duration bursts were not serially dependent during periods of stimulation. 6. A comparison of the autocorrelograms and higher-order interval histograms of single-unit responses in the poststimulation and spontaneous conditions indicated that 20% of single-unit spike trains contained an excess of short first-, second-, and third-order intervals following stimulation. This subgroups of single units could not be distinguished on the basis of the age of the animal or the depth at which the recording was made. 7. The low incidence of burst firing during stimulation opposes the view that bursts serve as a mechanism to emphasize or amplify particular stimulus-related responses in the presence of ongoing spontaneous activity in the primary auditory cortex. Moreover, there is little evidence to support the notion that brief bursts represent neural codes, because intraburst intervals are not serially dependent. It is suggested that pyramidal burst firing may be an effective way to evoke postsynaptic firing in inhibitory interneurons and subsequ


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