Intracellular Responses of Neurons in the Mouse Inferior Colliculus to Sinusoidal Amplitude-Modulated Tones

2009 ◽  
Vol 101 (4) ◽  
pp. 2002-2016 ◽  
Author(s):  
H.-R. Geis ◽  
J. G. G. Borst

Changes in the temporal envelope are important defining features of natural acoustic signals. Many cells in the inferior colliculus (IC) respond preferentially to certain modulation frequencies, but how they accomplish this is not yet clear. We therefore made whole cell patch-clamp recordings in the IC of anesthetized mice while presenting sinusoidal amplitude-modulated (SAM) tones. The relation between the number of evoked spikes and modulation frequency was used to construct rate modulation transfer functions (rMTFs). We observed different types of rate tuning, including band-pass (16%), band-reject (13%), high-pass (6%), and low-pass (6%) tuning. In the high-pass rMTF neurons and some of the low-pass rMTF neurons, the tuning characteristics appeared to be already present in the inputs. In both band-pass and band-reject rMTF neurons, the nonlinear relation between membrane potential and spike probability ensured preferential spiking during only a small part of the modulation period. Band-pass rMTF neurons had rapidly rising excitatory postsynaptic potentials, allowing good phase-locking to brief tones and intermediate modulation frequencies. At low modulation frequencies, adaptation of their spike threshold contributed to the onset response. In contrast, band-reject rMTF neurons responded with small excitatory or inhibitory postsynaptic potentials to brief tones. In these cells, a power law could describe the supralinear relation between average membrane potential and spike rate. Differences in timing of synaptic input and presence or absence of spike adaptation therefore define band-pass and band-reject rate tuning to SAM tones in the mouse IC.

2017 ◽  
Vol 26 (07) ◽  
pp. 1750121 ◽  
Author(s):  
Thanat Nonthaputha ◽  
Montree Kumngern

This paper presents new programmable universal biquadratic filters using current conveyor transconductance amplifiers (CCTAs) by which both voltage- and current-mode filters can be obtained. The proposed filters use second-generation current conveyor (CCII) which is the first stage of CCTA to operate as current conveyor analog switch (CCAS) and this CCAS will be used to program the filtering functions such as low-pass, high-pass, band-pass, band-stop and all-pass filters. Unlike previous universal filters, the filtering functions of the proposed filters can be programmed using the bias currents of CCTAs without changing any input and output connections. The natural frequency and quality factor of all filtering functions can be controlled electronically and orthogonally using the bias currents of transconductance amplifiers. Also gain response of all transfer functions can be adjusted. The active and passive sensitivities of the filters are low. The proposed programmable filters have been simulated using 0.18[Formula: see text][Formula: see text]m CMOS process from TSMC. PSPICE simulation results are included to confirm workability of the proposed circuits.


1998 ◽  
Vol 79 (5) ◽  
pp. 2303-2315 ◽  
Author(s):  
Yong Lu ◽  
Philip H.-S. Jen ◽  
Min Wu

Lu, Yong, Philip H.-S. Jen, and Min Wu. GABAergic disinhibition affects responses of bat inferior collicular neurons to temporally patterned sound pulses. J. Neurophysiol. 79: 2303–2315, 1998. Using the big brown bat, Eptesicus fuscus, as a model mammalian auditory system, we studied the effect of GABAergic disinhibition by bicuculline on the responses of inferior collicular (IC) neurons to temporally patterned trains of sound pulses delivered at different pulse repetition rates (PRRs) under free-field stimulation conditions. All 66 neurons isolated from eight bats either discharged one to two impulses (phasic on responders, n = 41, 62%), three to eight impulses (phasic bursters, n = 19, 29%), or many impulses throughout the entire duration of the stimulus (tonicresponders, n = 6, 9%). Whereas 50 neurons responded vigorously to frequency-modulated (FM) pulses, 16 responded poorly ornot at all to FM pulses. Bicuculline application increased the number of impulses of all 66 neurons in response to 4 ms pulses by 15–1,425%. The application also changed most phasic on responders into phasic bursters or tonic responders, resulting in 12 (18%) phasic on responders, 34 (52%) phasic bursters, and 20 (30%) tonic responders. Response latencies of these neurons were either shortened ( n = 25, 38%) by 0.5–6.0 ms, lengthened ( n = 9, 14%) by 0.5–2.5 ms or not changed ( n = 32, 48%) on bicuculline application. Each neuron had a highest response repetition rate beyond which the neuron failed to respond. Bicuculline application increased the highest response repetition rates of 62 (94%) neurons studied. The application also increased the highest 100% pulse-locking repetition rates of 21 (32%) neurons and facilitated 27 (41%) neurons in response to more pulses at the same PRR than predrug conditions. According to average rate-based modulation transfer functions (average rate MTFs), all 66 neurons had low-pass filtering characteristics both before and after bicuculline application. According to total discharge rate-based modulation transfer functions (total rate MTFs), filtering characteristics of these neurons can be described as band-pass ( n = 52, 79%), low-pass ( n = 12, 18%), or high-pass ( n = 2, 3%) before bicuculline application. Bicuculline application changed the filtering characteristics of 14 (21%) neurons. According to synchronization coefficient-based modulation transfer functions, filtering characteristics of these neurons can be described as low-pass ( n = 41, 62%), all-pass ( n = 11, 17%), band-suppression ( n = 7, 10.5%), and band-suppression–band-pass filters ( n = 7, 10.5%). Bicuculline application changed filtering characteristics of 19 (29%) neurons.


2006 ◽  
Vol 96 (5) ◽  
pp. 2388-2398 ◽  
Author(s):  
Ranjan Batra

Fluctuations in the amplitude of a sound play an important role in our perception of pitch and acoustic space, but their neural analysis has not been fully elucidated. The ventral nucleus of the lateral lemniscus (VNLL) has been implicated in the processing of such temporal features of a sound. This study examines responses of neurons in the VNLL of unanesthetized rabbits to sinusoidally amplitude modulated tones, a type of stimulus that has often been used to investigate encoding of temporal information. Modulation transfer functions of responses were calculated in two ways: based on discharge rates (rMTFs) and on synchronization to the envelope (tMTFs). Among the variety of rMTFs, two types were readily identifiable: flat and band-pass. The responses of neurons exhibiting these types of rMTF differed in several ways. Neurons with flat rMTFs typically had moderate rates of spontaneous activity, sustained responses to short tone bursts, and low-pass or band-pass tMTFs. Neurons with band-pass rMTFs typically had low spontaneous activity, onset responses to short tone bursts, and flat tMTFs. The vast majority synchronized strongly to the modulation envelope. The best modulation frequencies of neurons with band-pass rMTFs extended from 14 to 283 Hz. The presence of neurons with band-pass rMTFs in the VNLL suggests that this nucleus plays a role in converting the temporal code for modulation frequency used in lower structures into a rate-based code for use higher in the auditory pathway. The substantial number of neurons with more complex modulation transfer functions indicates that the VNLL has other functions.


2003 ◽  
Vol 90 (1) ◽  
pp. 477-490 ◽  
Author(s):  
Huiming Zhang ◽  
Jack B. Kelly

Recordings were made from single neurons in the rat inferior colliculus in response to sinusoidally amplitude-modulated sounds (10-s duration) presented to the contralateral ear. Neural responses were determined for different rates of modulation (0.5 Hz to 1 kHz) at a depth of 100%, and modulation transfer functions were generated based on firing rate (MTFFR) and vector strength (MTFVS). The effects of AMPA, NMDA, and GABAA receptor antagonists were examined by releasing drugs iontophoretically through a multibarrel pipette attached to a single-barrel recording pipette. Both the AMPA receptor antagonist, 1,2,3,4-tetrahydro-6-nitro-2,3-dioxo-benzo[f]quinoxaline-7-sulfonamide disodium (NBQX), and the NMDA receptor antagonist, (±)-3-(2-carboxypiperazin-4-yl)-propyl-1-phosphonic acid (CPP) resulted in a decrease in firing rate, and the GABAA receptor antagonist, bicuculline, produced an increase in the firing rate in most of the cells examined. In some cases, the shape of the MTFFR was modified slightly by receptor antagonists, but in most cases, the peak firing rate that determined a neuron's best modulation frequency remained the same. Also there were no changes during delivery of either excitatory or inhibitory antagonists in the maximum response synchrony at the peak of the MTFVS although some changes were noticed at off-peak modulation rates particularly with the AMPA receptor antagonist, NBQX.


2009 ◽  
Vol 101 (6) ◽  
pp. 3031-3041 ◽  
Author(s):  
Martin Pienkowski ◽  
Greg Shaw ◽  
Jos J. Eggermont

An extension of the Wiener-Volterra theory to a Poisson-distributed impulse train input was used to characterize the temporal response properties of neurons in primary auditory cortex (AI) of the ketamine-anesthetized cat. Both first- and second-order “Poisson-Wiener” (PW) models were tested on their predictions of temporal modulation transfer functions (tMTFs), which were derived from extracellular spike responses to periodic click trains with click repetition rates of 2–64 Hz. Second-order (i.e., nonlinear) PW fits to the measured tMTFs could be described as very good in a majority of cases (e.g., predictability ≥80%) and were almost always superior to first-order (i.e., linear) fits. In all sampled neurons, second-order PW kernels showed strong compressive nonlinearities (i.e., a depression of the impulse response) but never expansive nonlinearities (i.e., a facilitation of the impulse response). In neurons with low-pass tMTFs, the depression decayed exponentially with the interstimulus lag, whereas in neurons with band-pass tMTFs, the depression was typically double-peaked, and the second peak occurred at a lag that correlated with the neuron's best modulation frequency. It appears that modulation-tuning in AI arises in part from an interplay of two nonlinear processes with distinct time courses.


2006 ◽  
Vol 96 (6) ◽  
pp. 2905-2914 ◽  
Author(s):  
Huiming Zhang ◽  
Jack B. Kelly

Recordings were made from single neurons in the rat's ventral nucleus of the lateral lemniscus (VNLL) to determine responses to amplitude-modulated (AM) tones. The neurons were first characterized on the basis of their response to tone bursts presented to the contralateral ear and a distinction was made between those with transient onset responses and those with sustained responses. Sinusoidal AM tones were then presented to the contralateral ear with a carrier that matched the neuron's characteristic frequency (CF). Modulation transfer functions were generated on the basis of firing rate (MTFFR) and vector strength (MTFVS). Ninety-two percent of onset neurons that responded continuously to AM tones had band-pass MTFFRs with best modulation frequencies from 10 to 300 Hz. Fifty-four percent of sustained neurons had band-pass MTFFRs with best modulation frequencies from 10 to 500 Hz; other neurons had band-suppressed, all-pass, low-pass, or high-pass functions. Most neurons showed either band-pass or low-pass MTFVS. Responses were well synchronized to the modulation cycle with maximum vector strengths ranging from 0.37 to 0.98 for sustained neurons and 0.78 to 0.99 for onset neurons. The upper frequency limit for response synchrony was higher than that reported for inferior colliculus, but lower than that seen in more peripheral structures. Results suggest that VNLL neurons, especially those with onset responses to tone bursts, are sensitive to temporal features of sounds and narrowly tuned to different modulation rates. However, there was no evidence of a topographic relation between dorsoventral position along the length of VNLL and best modulation frequency as determined by either firing rate or vector strength.


2007 ◽  
Vol 16 (04) ◽  
pp. 507-516 ◽  
Author(s):  
SHAHRAM MINAEI ◽  
ERKAN YUCE

In this paper, a universal current-mode second-order active-C filter for simultaneously realizing low-pass, band-pass and high-pass responses is proposed. The presented filter employs only three plus-type second-generation current-controlled conveyors (CCCII+s). This filter needs no critical active and passive component matching conditions and no additional active and passive elements for realizing high output impedance low-pass, band-pass and high-pass characteristics. The angular resonance frequency (ω0) and quality factor (Q) of the proposed resistorless filter can be tuned electronically. To verify the theoretical analysis and to exhibit the performance of the proposed filter, it is simulated with SPICE program.


2005 ◽  
Vol 14 (01) ◽  
pp. 159-164 ◽  
Author(s):  
SUDHANSHU MAHESHWARI ◽  
IQBAL A. KHAN

A novel voltage-mode universal filter employing only two current differencing buffered amplifiers (CDBAs) is proposed. The filter uses four inputs and single output to realize six responses, viz. low-pass, high-pass, inverting band-pass, noninverting band-pass, band-elimination, and all-pass through input selection with independent pole-Q control. Computer simulation results using SPICE are also given to verify the theory.


1991 ◽  
Vol 65 (3) ◽  
pp. 424-445 ◽  
Author(s):  
A. S. Feng ◽  
J. C. Hall ◽  
S. Siddique

1. Physiological recordings were made from single auditory fibers in the frog eighth nerve to determine quantitatively how the different behaviorally relevant temporal parameters (the signal rise-fall time, duration, and rate of amplitude modulation) of complex sounds are encoded in the auditory periphery. Individual temporal parameters were varied. Response functions (RFs) were constructed with respect to each of these parameters using each unit's best excitatory frequency (BF) as the carrier. 2. In response to a change in signal rise-fall time, auditory nerve fibers showed little change in the mean spike count or firing rate, i.e., all fibers displayed ALL-PASS RFrfts. But the transient components, particularly the early phasic component, of responses varied with rise-fall times; these components were more pronounced in the responses to stimuli with shorter rise-fall times. 3. In response to an increase in signal duration, auditory nerve fibers showed a corresponding increase in firing duration and thus in the mean spike count, giving rise to HIGH-PASS RFdurs. The shape of response curves differed among fibers; the difference appeared to be related to the fiber's temporal adaptation characteristic. When the firing rate was measured, all fibers displayed higher mean firing rates in response to shorter duration stimuli than they did to longer duration stimuli, thus giving rise to LOW-PASS response functions. 4. To determine the response transfer functions to modulation rate, pulsed (PAM) and sinusoidally (SAM) amplitude-modulated signals were used. These signals differed substantially in terms of their envelopes and how they varied with AM rate. Data were analyzed by 1) plotting spike counts against the AM rate to derive modulation transfer functions (MTFspks) and 2) plotting synchronization coefficients (SCs) against the AM rate to generate MTFscs. 5. In response to PAM stimuli, all fibers showed an increase in mean spike count with modulation frequency over the range examined, giving rise to HIGH-PASS MTFspks. 6. For SAM stimuli, the average energy and duty cycle are independent of AM rate. Most (79%) auditory fibers showed little selectivity for AM rate over a range of 5-400 Hz, giving rise to ALL-PASS MTFspks. The remaining auditory fibers displayed LOW-PASS MTFspks, i.e., there was a distinct decline in the mean spike count with increasing AM rate. 7. In response to PAM stimuli, most fibers showed good response synchrony at low AM rates but the SC declined with an increase in the AM rate (i.e., LOW-PASS MTFscs). The cut-off frequency was typically very high, averaging 90 pulses/s.(ABSTRACT TRUNCATED AT 400 WORDS)


1988 ◽  
Vol 60 (6) ◽  
pp. 1799-1822 ◽  
Author(s):  
G. Langner ◽  
C. E. Schreiner

1. Temporal properties of single- and multiple-unit responses were investigated in the inferior colliculus (IC) of the barbiturate-anesthetized cat. Approximately 95% of recording sites were located in the central nucleus of the inferior colliculus (ICC). Responses to contralateral stimulation with tone bursts and amplitude-modulated tones (100% sinusoidal modulation) were recorded. Five response parameters were determined for neurons at each location: 1) characteristic frequency (CF); 2) onset latency of responses to CF-tones 60 dB above threshold; 3) Q10 dB (CF divided by bandwidth of tuning curve 10 dB above threshold); 4) best modulation frequency for firing rate (rBMF or BMF; amplitude modulation frequency that elicited the highest firing rate); and 5) best modulation frequency for synchronization (sBMF; amplitude modulation frequency that elicited the highest degree of phase-locking to the modulation frequency). 2. Response characteristics for single units and multiple units corresponded closely. A BMF was obtained at almost all recording sites. For units with a similar CF, a range of BMFs was observed. The upper limit of BMF increased approximately proportional to CF/4 up to BMFs as high as 1 kHz. The lower limit of encountered BMFs for a given CF also increased slightly with CF. BMF ranges for single-unit and multiple-unit responses were similar. Twenty-three percent of the responses revealed rBMFs between 10 and 30 Hz, 51% between 30 and 100 Hz, 18% between 100 and 300 Hz, and 8% between 300 and 1000 Hz. 3. For single units with modulation transfer functions of bandpass characteristics, BMFs determined for firing rate and synchronization were similar (r2 = 0.95). 4. Onset latencies for responses to CF tones 60 dB above threshold varied between 4 and 120 ms. Ninety percent of the onset latencies were between 5 and 18 ms. A range of onset latencies was recorded for different neurons with any given CF. The onset response latency of a given unit or unit cluster was significantly correlated with the period of the BMF and the period of the CF (P less than 0.05). 5."Intrinsic oscillations" of short duration, i.e., regularly timed discharges of units in response to stimuli without a corresponding temporal structure, were frequently observed in the ICC. Oscillation intervals were commonly found to be integer multiples of 0.4 ms. Changes of stimulus frequency or intensity had only minor influences on these intrinsic oscillations.(ABSTRACT TRUNCATED AT 400 WORDS)


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