Activity related to perceptual judgment and action in primary auditory cortex (A1)

In this paper we report the results of extensive research on phase modulation (PM) and amplitude modulation (AM) noise in linear bipolar junction transistor (BJT) amplifiers. BJT amplifiers exhibit 1/f PM and AM noise about a carrier signal that is much larger than the amplifier's thermal noise at those frequencies in the absence of the carrier signal. Our work shows that the 1/f PM noise of a BJT based amplifier is accompanied by 1/f AM noise which can be higher, lower, or nearly equal depending on the circuit implementation. The 1/f AM and PM noise in BJTs is primarily the result of 1/f fluctuations in transistor current, transistor capacitance, circuit supply voltages, circuit impedances, and circuit configuration. We discuss the theory and present experimental data in reference to common emitter amplifiers, but the analysis can be applied to other configurations as well. This study provides the functional dependence of 1/f AM and PM noise on transistor parameters, circuit parameters, and signal frequency, thereby laying the groundwork for a comprehensive theory of 1/f AM and PM noise in BJT amplifiers. We show that in many cases the 1/f PM and AM noise can be reduced below the thermal noise of the amplifier.

A practical implementation of a portable secondary standard for phase modulation (PM) and amplitude modulation (AM) noise at 5, 10, and 100 MHz is described. The accuracy of the standard for both PM and AM noise is +or-0.14 dB, and the temperature coefficient is less than 0.02 dB/K. The noise floor of the standard for PM noise measurements is less than -190 dBC rel to 1 rad/sup 2//Hz at 5, 10, and 100 MHz. The noise floor for AM measurements depends on the configuration. A calibrated level of PM and AM noise of approximately-130+or-0.2 dB rel to 1 rad/sup 2//Hz is used to evaluate the accuracy versus Fourier frequency. This standard can also be used as an alternative to the normal method of the PM/AM detector for PM/AM measurements. >

A portable amplitude-modulation (AM) and phase-modulation (PM) noise standard has been developed for carrier frequencies of 10.6, 21.2 and 42.4 GHz. The noise level from the carrier 1/2 S/sub /spl phi//(f) (phase noise) or 1/2 S/sub a/(f) (amplitude noise) is -122.6 dBc/Hz at 10.6 GHz, -119.6 dBc/Hz at 21.2 GHz and -116.3 dBc/Hz at 42.4 GHz. For Fourier frequencies of 1 kHz to 10 MHz the added noise is flat with an uncertainty of /spl plusmn/0.25 dB. The uncertainty of the noise level for Fourier frequencies of 1 kHz to 100 MHz is /spl plusmn/0.5 dB. The residual PM noise floor between the signal and reference outputs at 100 MHz is less than -182 dBc/Hz.

Tuning of cortical neurons to behaviorally salient acoustic signals

Developmental dyslexia is associated with deficits in the processing of basic auditory stimuli. Yet it is unclear how these sensory impairments might contribute to poor reading skills. This study better characterizes the relationship between phonological decoding skills, the lack of which is generally accepted to comprise the core deficit in reading disabilities, and auditory sensitivity to amplitude modulation (AM) and frequency modulation (FM). Thirty-eight adult subjects, 17 of whom had a history of developmental dyslexia, completed a battery of psychophysical measures of sensitivity to FM and AM at different modulation rates, along with a measure of pseudoword reading accuracy and standardized assessments of literacy and cognitive skills. The subjects with a history of dyslexia were significantly less sensitive than controls to 2-Hz FM and 20-Hz AM only. The absence of a significant group difference for 2-Hz AM shows that the dyslexics do not have a general deficit in detecting all slow modulations. Thresholds for detecting 2-Hz and 240-Hz FM and 20-Hz AM correlated significantly with pseudoword reading accuracy. After accounting for various cognitive skills, however, multiple regression analyses showed that detection thresholds for both 2-Hz FM and 20-Hz AM were significant and independent predictors of pseudoword reading ability in the entire sample. Thresholds for 2-Hz AM and 240-Hz FM did not explain significant additional variance in pseudoword reading skill. It is therefore possible that certain components of auditory processing of modulations are related to phonological decoding skills, whereas others are not.

The perception of spectrotemporal changes is crucial for distinguishing between acoustic signals, including vocalizations. Temporal modulation transfer functions (TMTFs) have been measured in many species and reveal that the discrimination of amplitude modulation suffers at rapid modulation frequencies. TMTFs were measured in six CBA/CaJ mice in an operant conditioning procedure, where mice were trained to discriminate an 800 ms amplitude modulated white noise target from a continuous noise background. TMTFs of mice show a bandpass characteristic, with an upper limit cutoff frequency of around 567 Hz. Within the measured modulation frequencies ranging from 5 Hz to 1280 Hz, the mice show a best sensitivity for amplitude modulation at around 160 Hz. To look for a possible parallel evolution between sound perception and production in living organisms, we also analyzed the components of amplitude modulations embedded in natural ultrasonic vocalizations (USVs) emitted by this strain. We found that the cutoff frequency of amplitude modulation in most of the individual USVs is around their most sensitive range obtained from the psychoacoustic experiments. Further analyses of the duration and modulation frequency ranges of USVs indicated that the broader the frequency ranges of amplitude modulation in natural USVs, the shorter the durations of the USVs.

When an RF carrier is amplified by a GaAs MESFET, amplitude modulation (AM) and phase modulation (PM) noises are imposed on the carrier. This is generally believed to be caused by transposition to the carrier frequency of the low-frequency flicker noise generated by the FET. The cross correlation between the AM and PM noises and the low-frequency noise observed on the drain of the FET is measured. While the AM noise and the low-frequency noise on the drain of the FET exhibit a high degree of correlation, the PM noise and the low-frequency drain noise are not highly correlated. The latter result may explain the limited success of oscillator phase noise reduction methods which rely on the existence of a large cross correlation between the PM and low-frequency noises. >

Human auditory perception and speech intelligibility have been successfully described based on the two concepts of spectral masking and amplitude modulation (AM) masking. The power-spectrum model (PSM) [Patterson and Moore (1986). Frequency Selectivity in Hearing, pp. 123-177] accounts for effects of spectral masking and critical bandwidth, while the envelope power-spectrum model (EPSM) [Ewert and Dau (2000). J. Acoust. Soc. Am. 108, 1181-1196] has been successfully applied to AM masking and discrimination. Both models extract the long-term (envelope) power to calculate signal-to-noise ratios (SNR). Recently, the EPSM has been applied to speech intelligibility (SI) considering the short-term envelope SNR on various time scales (multi-resolution speech-based envelope power-spectrum model; mr-sEPSM) to account for SI in fluctuating noise [Jørgensen, Ewert, and Dau (2013). J. Acoust. Soc. Am. 134, 436-446]. Here, a generalized auditory model is suggested combining the classical PSM and the mr-sEPSM to jointly account for psychoacoustics and speech intelligibility. The model was extended to consider the local AM depth in conditions with slowly varying signal levels, and the relative role of long-term and short-term SNR was assessed. The suggested generalized power-spectrum model is shown to account for a large variety of psychoacoustic data and to predict speech intelligibility in various types of background noise.

This study investigates the degree of similarity of three different auditory cortical areas with respect to the coding of periodic stimuli. Simultaneous single- and multiunit recordings in response to periodic stimuli were made from primary auditory cortex (AI), anterior auditory field (AAF), and secondary auditory cortex (AII) in the cat to addresses the following questions: is there, within each cortical area, a difference in the temporal coding of periodic click trains, amplitude-modulated (AM) noise bursts, and AM tone bursts? Is there a difference in this coding between the three cortical fields? Is the coding based on the temporal modulation transfer function (tMTF) and on the all-order interspike-interval (ISI) histogram the same? Is the perceptual distinction between rhythm and roughness for AM stimuli related to a temporal versus spatial representation of AM frequency in auditory cortex? Are interarea differences in temporal response properties related to differences in frequency tuning? The results showed that: 1) AM stimuli produce much higher best modulation frequencies (BMFs) and limiting rates than periodic click trains. 2) For periodic click trains and AM noise, the BMFs and limiting rates were not significantly different for the three areas. However, for AM tones the BMF and limiting rates were about a factor 2 lower in AAF compared with the other areas. 3) The representation of stimulus periodicity in ISIs resulted in significantly lower mean BMFs and limiting rates compared with those estimated from the tMTFs. The difference was relatively small for periodic click trains but quite large for both AM stimuli, especially in AI and AII. 4) Modulation frequencies <20 Hz were represented in the ISIs, suggesting that rhythm is coded in auditory cortex in temporal fashion. 5) In general only a modest interdependence of spectral- and temporal-response properties in AI and AII was found. The BMFs were correlated positively with characteristic frequency in AAF. The limiting rate was positively correlated with the frequency-tuning curve bandwidth in AI and AII but not in AAF. Only in AAF was a correlation between BMF and minimum latency was found. Thus whereas differences were found in the frequency-tuning curve bandwidth and minimum response latencies among the three areas, the coding of periodic stimuli in these areas was fairly similar with the exception of the very poor representation of AM tones in AII. This suggests a strong parallel processing organization in auditory cortex.

We show that a high-resolution software package can be adapted to calculate the amplitude modulation (AM) and phase modulation (PM) noise performance of Si Bipolar Junction Transistor (BJT) amplifiers. Our simulations predict that the PM noise induced by bias current noise can be greatly suppressed by operating at special bias conditions. Furthermore, a dc balance amplifier circuit is introduced to stabilize the base-to-collector voltage. The total sensitivity of PM noise to power supply noise using dc balance architecture can be reduced by 40 dB compared to basic amplifier architecture. This is experimentally verified.

The acquisition of new auditory skills can be facilitated by experiencing a conspecific performing a well-defined behavior (i.e., social learning). Although the neural bases for auditory social learning remain uncertain, one plausible hypothesis is that social experience induces long-term changes to auditory cortex, thereby facilitating the subsequent acquisition of an auditory skill. To explore this idea, we developed a social learning paradigm in which naïve observer gerbils are exposed to a demonstrator that performs an amplitude modulation (AM) rate discrimination task on the other side of an opaque divider. Thus, observers have no access to visual cues. When exposed to a demonstrator for five successive days, observers subsequently acquire the AM task more rapidly than controls. Two experiments suggest that auditory cortex is necessary and sufficient for social learning. First, inactivating the observer’s auditory cortex during each social exposure session significantly delayed task acquisition. Second, recordings from the observer’s auditory cortex revealed that individual neurons displayed a significant improvement in AM stimulus discrimination across the five exposure sessions, and the magnitude of neural improvement correlated with an animal’s subsequent rate of task acquisition. Together, these findings suggest that auditory cortex plasticity plays a pivotal role in social learning.

Temporal envelope processing is critical for speech comprehension, which is known to be affected by normal aging. Whereas the macaque is an excellent animal model for human cerebral cortical function, few studies have investigated neural processing in the auditory cortex of aged, nonhuman primates. Therefore, we investigated age-related changes in the spiking activity of neurons in primary auditory cortex (A1) of two aged macaque monkeys using amplitude-modulated (AM) noise and compared these responses with data from a similar study in young monkeys (Yin P, Johnson JS, O'Connor KN, Sutter ML. J Neurophysiol 105: 582-600, 2011). For each neuron, we calculated firing rate (rate code) and phase-locking using phase-projected vector strength (temporal code). We made several key findings where neurons in old monkeys differed from those in young monkeys. Old monkeys had higher spontaneous and driven firing rates, fewer neurons that synchronized with the AM stimulus, and fewer neurons that had differential responses to AM stimuli with both a rate and temporal code. Finally, whereas rate and temporal tuning functions were positively correlated in young monkeys, this relationship was lost in older monkeys at both the population and single neuron levels. These results are consistent with considerable evidence from rodents and primates of an age-related decrease in inhibition throughout the auditory pathway. Furthermore, this dual coding in A1 is thought to underlie the capacity to encode multiple features of an acoustic stimulus. The apparent loss of ability to encode AM with both rate and temporal codes may have consequences for stream segregation and effective speech comprehension in complex listening environments.

Characteristics of neurons in auditory cortex of monkeys performing a simple auditory task.

Whether attention exerts its impact already on primary sensory levels is still a matter of debate. Particularly in the auditory domain the amount of empirical evidence is scarce. Recently noninvasive and invasive studies have shown attentional modulations of the auditory Steady-State Response (aSSR). This evoked oscillatory brain response is of importance to the issue, because the main generators have been shown to be located in primary auditory cortex. So far, the issue whether the aSSR is sensitive to the predictive value of a cue preceding a target has not been investigated. Participants in the present study had to indicate on which ear the faster amplitude modulated (AM) sound of a compound sound (42 and 19 Hz AM frequencies) was presented. A preceding auditory cue was either informative (75%) or uninformative (50%) with regards to the location of the target. Behaviorally we could confirm that typical attentional modulations of performance were present in case of a preceding informative cue. With regards to the aSSR we found differences between the informative and uninformative condition only when the cue/target combination was presented to the right ear. Source analysis indicated this difference to be generated by a reduced 42 Hz aSSR in right primary auditory cortex. Our and previous data by others show a default tendency of "40 Hz" AM sounds to be processed by the right auditory cortex. We interpret our results as active suppression of this automatic response pattern, when attention needs to be allocated to right ear input.

This study investigates the auditory continuity illusion, combining psychophysics and magnetoencephalography. Stimuli consisted of amplitude-modulated (AM) noise interrupted by bursts of louder, unmodulated noise. Subjective judgments confirmed that the AM was perceived as continuous, a case of illusory continuity. Psychophysical measurements showed that the illusory modulation had little effect on the detection of a physical modulation, i.e., the illusory modulation produced no modulation masking. Duration discrimination thresholds for the AM noise segments, however, were elevated by the illusion. A whole-head magnetoencephalographic system was used to record brain activity when listeners attended passively to the stimuli. The AM noise produced a modulated magnetic activity, the auditory steady-state response. The illusory modulation did not produce such a response, instead, a possible neural correlate of the illusion was found in transient evoked responses. When the AM was interrupted by silence, oscillatory activity in the gamma-band range as well as slow evoked potentials were observed at each AM onset. In the case of the illusion, these neural responses were largely reduced. Both sets of results are inconsistent with a restoration of the modulation in the case of illusory continuity. Rather, they point to a role for onset-detection mechanisms in auditory scene analysis.