Binaural masking and sensitivity to interaural delay in the inferior colliculus

1992 ◽  
Vol 336 (1278) ◽  
pp. 415-422 ◽  

The binaural masking level difference (BMLD) is a psychophysical effect whereby signals masked by a noise at one ear become unmasked by sounds reaching the other, BMLD effects are largest at low frequencies where they depend on signal phase, suggesting that part of the physiological mechanism responsible for the BMLD resides in cells that are sensitive to interaural time disparities. We have investigated a physiological basis for unmasking in the responses of delay-sensitive cells in the central nucleus of the inferior colliculus in anaesthetized guinea pigs. The masking effects of a binaurally presented noise, as a function of the masker delay, were quantified by measuring the number of discharges synchronized to the signal, and by measuring the masked threshold. The noise level for masking was lowest at the best delay for the noise. The mean magnitude of the unmasking across our neural population was similar to the human psychophysical BMLD under the same signal and masker conditions.

1979 ◽  
Vol 42 (6) ◽  
pp. 1626-1639 ◽  
Author(s):  
M. N. Semple ◽  
L. M. Aitkin

1. The discharges of 632 units were isolated extracellularly during 42 penetrations of the central nucleus of the inferior colliculus (ICC) in 21 adult cats lightly anesthetized with pentobarbital and ketamine. Microelectrode penetrations were directed from caudal to rostral through ICC, parallel to the Horsley-Clarke (H-C) horizontal and sagittal planes. 2. The threshold best frequency (BF) and binaural response properties were examined for each unit, with the aim of elucidating the organization of these discharge characteristics within ICC. 3. Binaural unit classes consisted of monaural (contralateral) (EO), binaurally phase-sensitive (delay), contralateral excitatory/ipsilateral inhibitory (EI), binaurally excitatory (EE), and other more heterogeneous interaction patterns (other). 4. Detailed histological reconstruction of electrode tracks allowed the recording site for each unit to be related to the three dimensions of ICC. This structure was divided into three lateromedial and three rostrocaudal blocks such that each block contained a similar number of units, enabling meaningful statistical comparisons. Low (3.2 kHz greater than BF) and high (3.2 kHz less than BF) best-frequency classes provided a correlate of dorsoventral location. 5. The arrangements of BFs within ICC were found to be compatible with a model of this structure in which units having similar BFs are organized into layers lying in the H-C horizontal plane medially and gradually tilting in both a ventrolateral and ventrorostral direction. Low frequencies are concentrated dorsally and laterally; high frequencies, ventrally and medially. A rostrocaudal BF difference arises only in lateral aspects of the ICC, where lower frequencies are encountered rostrally. 6. Binaural response classes were distributed differentially throughout ICC. Thus, EO units were concentrated caudally, ventrally, and laterally, while delay units were in greatest numbers rostrally, dorsally, and laterally--almost totally segregated from EO and EI units. The latter populations overlapped ventrally and laterally, but EI units were in greatest density rostrally. The EE class occurred throughout the nucleus, but was most common medially. 7. It is suggested that the differential distributions of binaural responses reflect a partial segregation of the afferents, arising in the superior olive and cochlear nucleus, which terminate in ICC. The central nucleus of the inferior colliculus thus may be composed of several functionally segregated subregions contained within a common tonotopic organization.


1994 ◽  
Vol 72 (3) ◽  
pp. 1061-1079 ◽  
Author(s):  
Z. M. Fuzessery

1. While hunting, the pallid bat uses passive sound localization at low frequencies to find terrestrial prey, and echolocation for general orientation. It must therefore process two different types of acoustic input at the same time. The pallid bat's echolocation pulse is a downward frequency-modulated (FM) sweep from 60 to 30 kHz. This study examined the response selectivity of single neurons in the pallid bat's central nucleus of the inferior colliculus (ICC) for FM sweeps, comparing the response properties of the high-frequency population, tuned to the biosonar pulse, with the low-frequency population, tuned below the pulse. The working hypothesis was that the high-frequency population would exhibit a response selectivity for downward FM sweeps that was not present in the low-frequency population. 2. Neurons were tested for their selectivity for FM sweep direction, duration, frequency range and bandwidth, and rate of frequency change. The extent to which they responded exclusively to tones, noise, and FM sweeps was also examined. Significant differences in the response properties of neurons in the two populations were found. In the low-frequency population, all neurons responded to tones, but only 50% responded to FM sweeps. Only 23% were selective for sweep direction. In the high-frequency population, all neurons responded to FM sweeps, but 31% did not respond to tones. Over one-half of this population was selective for sweep direction, and of those that were selective, all preferred the downward sweep direction of the biosonar pulse. A large percentage (31%) responded exclusively to downward sweeps, and not to tones or upward sweeps. None of the cells in either population responded to noise, or did so only at very high relative thresholds. 3. Both populations contained neurons that were selective for short stimulus durations that approximated the duration of the biosonar pulse, although the percentage was greater in the high-frequency population (58% vs. 20%). In the high-frequency population, 31% of the neurons tested for duration responded exclusively to both the sweep direction and duration of the biosonar pulse. 4. Downward FM-selective neurons, with one exception, were generally insensitive to the rate of frequency change of the FM sweep, as well as the frequency range and bandwidth of the sweep. They responded similarly to both the full 60- to 30-kHz sweep and to 5-kHz bandwidth portions of the full sweep.(ABSTRACT TRUNCATED AT 400 WORDS)


1975 ◽  
Vol 38 (5) ◽  
pp. 1196-1207 ◽  
Author(s):  
L. M. Aitkin ◽  
W. R. Webster ◽  
J. L. Veale ◽  
D. C. Crosby

The responses of 150 units in the central (ICC), pericentral (ICP), and external nuclei (ICX) of the inferior colliculus of the anesthetized cat were studied in relation to their tuning characteristics and binaural responses to tonal stimuli. Units in ICC were characterized by sharp tuning and binaural responses, while those in ICP and ICX were frequently very broadly tuned with a poorly defined best frequency. Nonetheless, in the latter nuclei a tendency existed for tonotopic organization to occur with high frequencies located externally and low frequencies at the margins of the central nucleus. Tuning measurements were hampered by the common occurrence of habituation in the discharges of single units in ICP and, to a lesser extend, ICX. The majority of units in ICP could be differentiated from those in ICX by their monaural input. Speculations were advanced linking anatomical cell types to physiological responses in the three nuclei and into the possible functional significance of the different behavior of units to tonal stimuli.


2006 ◽  
Vol 96 (6) ◽  
pp. 3064-3073 ◽  
Author(s):  
Yuri B. Saalmann ◽  
Ian G. Morgan ◽  
Mike B. Calford

Fast inhibitory neurotransmission in the brain is largely mediated by the γ-aminobutyric acid-type A (GABAA) receptor. The 3α,5α-reduced neurosteroids (e.g., allopregnanolone) are the most potent endogenous modulators of the GABAA receptor. Although it is known that 3α,5α-reduced neurosteroid levels change during stress or depression and over the estrus cycle, a basic physiological role consistent with their pharmacological action remains elusive. We used the unique architecture of the auditory midbrain to reveal a role for 3α,5α-reduced neurosteroids in regulating inhibitory efficacy. After blocking the massive GABAergic projection from the dorsal nucleus of the lateral lemniscus (DNLL) to the contralateral central nucleus of the inferior colliculus (ICC) in anesthetized rats, a reactive increase in the efficacy of other inhibitory circuits in the ICC (separable because of the dominant ear that drives each circuit) was demonstrated with physiological measures—single-neuron activity and a neural-population-evoked response. This effect was prevented by blocking 3α,5α-reduced neurosteroid synthesis with a 5α-reductase inhibitor: finasteride. Immunohistochemistry confirmed that the DNLL blockade induced an increase in 3α,5α-reduced neurosteroids in the contralateral ICC. This study shows that when GABAergic inhibition is reduced, the brain compensates within minutes by locally increasing synthesis of neurosteroids, thereby balancing excitatory and inhibitory inputs in complex neural circuits.


1999 ◽  
Vol 81 (2) ◽  
pp. 722-734 ◽  
Author(s):  
Alan R. Palmer ◽  
Dan Jiang ◽  
David McAlpine

Desynchronizing responses to correlated noise: a mechanism for binaural masking level differences at the inferior colliculus. We examined the adequacy of decorrelation of the responses to dichotic noise as an explanation for the binaural masking level difference (BMLD). The responses of 48 low-frequency neurons in the inferior colliculus of anesthetized guinea pigs were recorded to binaurally presented noise with various degrees of interaural correlation and to interaurally correlated noise in the presence of 500-Hz tones in either zero or π interaural phase. In response to fully correlated noise, neurons’ responses were modulated with interaural delay, showing quasiperiodic noise delay functions (NDFs) with a central peak and side peaks, separated by intervals roughly equivalent to the period of the neuron’s best frequency. For noise with zero interaural correlation (independent noises presented to each ear), neurons were insensitive to the interaural delay. Their NDFs were unmodulated, with the majority showing a level of activity approximately equal to the mean of the peaks and troughs of the NDF obtained with fully correlated noise. Partial decorrelation of the noise resulted in NDFs that were, in general, intermediate between the fully correlated and fully decorrelated noise. Presenting 500-Hz tones simultaneously with fully correlated noise also had the effect of demodulating the NDFs. In the case of tones with zero interaural phase, this demodulation appeared to be a saturation process, raising the discharge at all noise delays to that at the largest peak in the NDF. In the majority of neurons, presenting the tones in π phase had a similar effect on the NDFs to decorrelating the noise; the response was demodulated toward the mean of the peaks and troughs of the NDF. Thus the effect of added tones on the responses of delay-sensitive inferior colliculus neurons to noise could be accounted for by a desynchronizing effect. This result is entirely consistent with cross-correlation models of the BMLD. However, in some neurons, the effects of an added tone on the NDF appeared more extreme than the effect of decorrelating the noise, suggesting the possibility of additional inhibitory influences.


1999 ◽  
Vol 82 (1) ◽  
pp. 164-175 ◽  
Author(s):  
Kevin A. Davis ◽  
Ramnarayan Ramachandran ◽  
Bradford J. May

Single units in the central nucleus of the inferior colliculus (ICC) of unanesthetized decerebrate cats can be grouped into three distinct types (V, I, and O) according to the patterns of excitation and inhibition revealed in contralateral frequency response maps. This study extends the description of these response types by assessing their ipsilateral and binaural response map properties. Here the nature of ipsilateral inputs is evaluated directly using frequency response maps and compared with results obtained from methods that rely on sensitivity to interaural level differences (ILDs). In general, there is a one-to-one correspondence between observed ipsilateral input characteristics and those inferred from ILD manipulations. Type V units receive ipsilateral excitation and show binaural facilitation (EE properties); type I and type O units receive ipsilateral inhibition and show binaural excitatory/inhibitory (EI) interactions. Analyses of binaural frequency response maps show that these ILD effects extend over the entire receptive field of ICC units. Thus the range of frequencies that elicits excitation from type V units is expanded with increasing levels of ipsilateral stimulation, whereas the excitatory bandwidth of type I and O units decreases under the same binaural conditions. For the majority of ICC units, application of bicuculline, an antagonist for GABAA-mediated inhibition, does not alter the basic effects of binaural stimulation; rather, it primarily increases spontaneous and maximum discharge rates. These results support our previous interpretations of the putative dominant inputs to ICC response types and have important implications for midbrain processing of competing free-field sounds that reach the listener with different directional signatures.


2006 ◽  
Vol 95 (3) ◽  
pp. 1926-1935 ◽  
Author(s):  
Liang-Fa Liu ◽  
Alan R. Palmer ◽  
Mark N. Wallace

In the auditory system, some ascending pathways preserve the precise timing information present in a temporal code of frequency. This can be measured by studying responses that are phase-locked to the stimulus waveform. At each stage along a pathway, there is a reduction in the upper frequency limit of the phase-locking and an increase in the steady-state latency. In the guinea pig, phase-locked responses to pure tones have been described at various levels from auditory nerve to neocortex but not in the inferior colliculus (IC). Therefore we made recordings from 161 single units in guinea pig IC. Of these single units, 68% (110/161) showed phase-locked responses. Cells that phase-locked were mainly located in the central nucleus but also occurred in the dorsal cortex and external nucleus. The upper limiting frequency of phase-locking varied greatly between units (80−1,034 Hz) and between anatomical divisions. The upper limits in the three divisions were central nucleus, >1,000 Hz; dorsal cortex, 700 Hz; external nucleus, 320 Hz. The mean latencies also varied and were central nucleus, 8.2 ± 2.8 (SD) ms; dorsal cortex, 17.2 ms; external nucleus, 13.3 ms. We conclude that many cells in the central nucleus receive direct inputs from the brain stem, whereas cells in the external and dorsal divisions receive input from other structures that may include the forebrain.


1994 ◽  
Vol 266 (6) ◽  
pp. G978-G986 ◽  
Author(s):  
W. Hasibeder ◽  
R. Germann ◽  
H. Sparr ◽  
M. Haisjackl ◽  
B. Friesenecker ◽  
...  

The mucosa of the small intestine has some unique microcirculatory features that may result in significant tissue oxygenation changes even under physiological conditions. To prove this hypothesis we investigated mucosal and serosal oxygenation in an autoperfused, innervated jejunal segment in pigs. Eight animals (30-40 kg) were anesthetized, paralyzed, and normoventilated. A small segment of the jejunal mucosa and serosa was exposed by a midline laparotomy and an antimesenteric incision. Mucosal and serosal oxygen tensions were measured using Clark-type surface oxygen electrodes. Mucosal hemoglobin saturation and concentration were determined by tissue reflectance spectrophotometry. Systemic hemodynamics, mesenteric-venous acid base, and blood gas variables, as well as systemic acid-base and blood gas variables and jejunal electromyogenic potentials, were recorded. Measurements were performed after a rest period at 0, 30, 60, and 90 min. All animals remained hemodynamically stable. At time 0 the jejunal oxygen extraction ratio was 0.33 +/- 0.05, the mean serosal PO2 was 60.25 +/- 7.69, the mean mucosal PO2 was 25.47 +/- 4.41 mmHg, and the mean mucosal hemoglobin saturation was 46.36 +/- 6.22%. Mean values did not change with time. In contrast to serosal PO2, mucosal PO2, mucosal hemoglobin oxygen saturation, and hemoglobin concentration showed rhythmic oscillations with a frequency of 3.4-5 cycles/min that were unrelated to systemic hemodynamic parameters, respiratory frequency, and intestinal peristalsis. From this we concluded that the jejunal mucosa demonstrates significant, regular changes in oxygenation parameters that are locally mediated. We speculate that the physiological basis for this phenomenon is the countercurrent arrangement of microvessels in conjunction with vasomotion.(ABSTRACT TRUNCATED AT 250 WORDS)


1997 ◽  
Vol 78 (1) ◽  
pp. 417-428 ◽  
Author(s):  
Mary Kate Worden ◽  
Maria Bykhovskaia ◽  
John T. Hackett

Worden, Mary Kate, Maria Bykhovskaia, and John T. Hackett. Facilitation at the lobster neuromuscular junction: a stimulus-dependent mobilization model. J. Neurophysiol. 78: 417–428, 1997. Frequency facilitation is a process whereby neurosecretion increases as a function of stimulation frequency during repetitive synaptic activity. To examine the physiological basis underlying facilitation, we have estimated the frequency dependence of the synaptic parameters n (number of units capable of responding to a nerve impulse) and P (average probability of responding) at the lobster neuromuscular junction. Both n and P increase as a function of frequency, suggesting that the efficiency of quantal docking and quantal fusion is regulated by repetitive synaptic activity. In experiments in which facilitation is strong and quantal content does not saturate over the frequency range tested, the value of P saturates at low frequencies of stimulation, and increases in quantal content at higher frequencies of stimulation are due to an increase in n. Therefore the value of P does not limit facilitation. We propose that transmitter release is limited by the rates of quantal mobilization and demobilization, and that each excitatory stimulus causes additional mobilization of quanta to dock at the presynaptic release sites. In such a model the binomial parameter n will correspond to the number of quanta docked at the release sites and available for release. We have developed and solved kinetic equations that describe how the number of docked quanta changes as a function of time and of stimulation frequency. The stimulus-dependent mobilization model of facilitation predicts that the reciprocal value of the quantal content depends linearly on the reciprocal product of the stimulation frequency and the probability of release. Fits of the experimental data confirm the accuracy of this prediction, showing that the model proposed here quantitatively describes frequency facilitation. The model predicts that high rates of quantal demobilization will produce strong frequency facilitation.


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