Bilateral Ablation of Auditory Cortex in Mongolian Gerbil Affects Discrimination of Frequency Modulated Tones but not of Pure Tones

This study examines the role of auditory cortex in the Mongolian gerbil in differential conditioning to pure tones and to linearly frequency-modulated (FM) tones by analyzing the effects of bilateral auditory cortex ablation. Learning behavior and performance were studied in a GO/NO-GO task aiming at avoidance of a mild foot shock by crossing a hurdle in a two-way shuttle box. Hurdle crossing as the conditioned response to the reinforced stimulus (CR+), as false alarm in response to the unreinforced stimulus (CR−), intertrial activity, and reaction times were monitored. The analysis revealed no effects of lesion on pure tone discrimination but impairment of FM tone discrimination. In the latter case lesion effects were dependent on timing of lesion relative to FM tone discrimination training. Lesions before training in naive animals led to a reduced CR+ rate and had no effect on CR− rate. Lesions in pretrained animals led to an increased CR− rate without effects on the CR+ rate. The results suggest that auditory cortex plays a more critical role in discrimination of FM tones than in discrimination of pure tones. The different lesion effects on FM tone discrimination before and after training are compatible with both the hypothesis of a purely sensory deficit in FM tone processing and the hypothesis of a differential involvement of auditory cortex in acquisition and retention, respectively.     

Perceptual grouping in audition

The present experiments evaluated the effect of relative frequency as a determinant of the figure-ground organization of sequences of auditory tones. Observers counted sequences of 20 ms tones that were presented at the same frequency or that alternated between two different frequencies. The alternating tones differed in frequency by one whole tone, seven tones, or nineteen tones. Counting accuracy increased with increases in the silent interval between the tones. When the alternating tones differed by seven or nineteen tones, counting was disrupted at rates of presentation of eight tones per second or slower. In contrast to this decrement in the counting of tones that alternated by over an octave, very little decrement was observed when the tones alternated by just one whole tone. The best subjects counted these alternating tones more accurately than the tones presented at the same frequency. The poorest subjects showed a small decrement even when the tones alternated by just one whole tone. The results were discussed in terms of determinants of figure-ground organization in auditory information processing.     

Maturation of rapid auditory temporal processing and subsequent nonword repetition performance in children

According to the rapid auditory processing theory, the ability to parse incoming auditory information underpins learning of oral and written language. There is wide variation in this low-level perceptual ability, which appears to follow a protracted developmental course. We studied the development of rapid auditory processing using event-related potentials (ERPs) elicited by tone pairs presented at varying inter-stimulus intervals (25, 50, 100, 200, and 400 ms) in a sample of children (N = 103) aged 7-9 years initially and again at 9-11 years. We also assessed their ability to repeat nonsense words at both time-points. The amount of difference between the ERP to single tones and paired tones (as assessed by the intra-class correlation coefficient, ICC) provided a measure of the brain's capacity to discriminate auditory information delivered at different presentation rates. Results showed that older children showed greater neural discrimination to tone pairs than younger children at rapid presentation rates, although these differences were reduced at slower presentation rates. The ICC at time 1 significantly predicted nonword repetition scores two years later, providing support for the view that rapid auditory temporal processing ability affects oral language development in typically developing children.     

Electrophysiologic Manifestations of Impaired Temporal Lobe Auditory Processing in Verbal Auditory Agnosia

The present study examined the extent to which verbal auditory agnosia (VAA) is primarily a phonemic decoding disorder, as contrasted to a more global defect in acoustic processing. Subjects were six young adults who presented with VAA in childhood and who, at the time of testing, showed varying degrees of residual auditory discrimination impairment. They were compared to a group of young adults with normal language development matched for age and gender. Cortical event-related potentials (ERPs) were recorded to tones and to consonant-vowel stimuli presented in an "oddball" discrimination paradigm. In addition to cortical ERPs, auditory brainstem responses (ABRs) and middle latency responses (MLRs) were recorded. Cognitive and language assessments were obtained for the VAA subjects. ABRs and MLRs were normal. In comparison with the control group, the cortical ERPs of the VAA subjects showed a delay in the N1 component recorded over lateral temporal cortex both to tones and to speech sounds, despite an N1 of normal latency overlying the frontocentral region of the scalp. These electrophysiologic findings indicate a slowing of processing of both speech and nonspeech auditory stimuli and suggest that the locus of this abnormality is within the secondary auditory cortex in the lateral surface of the temporal lobes.     

Avoidance responding as a function of stimulus duration and relation to free shock

Response-independent pairings of a tone and a brief shock were superimposed on uncued avoidance responding in four groups of rhesus monkeys. For one group, tone presentations were immediately followed by an unavoidable electric shock; for the remaining groups, gaps of 5, 20, and 80 sec intervened between tone termination and shock delivery. These temporal values subsume paradigms usually treated as discrete procedures; the conditioned emotional response procedure (0-sec gap between tone and shock), trace procedure (5-sec gap) and safety-signal training (80-sec gap). Within each group, tone durations of 10, 20, 40, and 80 sec were examined. A response pattern marked by maximum response rate in the initial 5 sec of the tone followed by deceleration before shock was observed when shock immediately followed the tone, but not when gaps were interposed between the tone and shock. Response rates in the first 5 sec of the tone were a function of both tone duration and duration of the gap. When the gap was 0 to 5 sec, initial response rates were highest in longer duration tones; this relationship between tone duration and initial tone response rate was not observed for longer gaps.     

Echoic memory in the rat: effects of inspection time, retention interval, and the spectral composition of masking noise.

Memory for tones (1100 vs. 2330 Hz) was studied in 4 rats (Rattus norvegicus), as affected by the durations of both target tones (30 to 620 ms) and noise-filled retention intervals (0 to 480 ms). With a 0-ms delay, performance was near asymptotic with the 30-ms tone, but the memory of this brief tone suffered a massive decrement at retention intervals as brief as 60 ms; in contrast, memory for the 340-ms tone was stable for at least 240 ms. If the retention interval was filled by band-stop noise (with targets presented in the spectral gap), then the rat's memory for brief tones was superior to that obtained with the standard broad-band noise filler, and band-stop noise was better than a band-pass noise that had the tones embedded in the region of its spectral energy. These findings are consistent with the hypotheses that auditory memory in the rat consists of a transient sensorylike echoic store and a short-term store more resistant to the effects of retroactive interference.     

The effect of exogenous spatial attention on auditory information processing

This study investigated the effect of exogenous spatial attention on auditory information processing. In Experiments 1, 2 and 3, temporal order judgment tasks were performed to examine the effect. In Experiment 1 and 2, a cue tone was presented to either the left or right ear, followed by sequential presentation of two target tones. The subjects judged the order of presentation of the target tones. The results showed that subjects heard both tones simultaneously when the target tone, which was presented on the same side as the cue tone, was presented after the target tone on the opposite side. This indicates that spatial exogenous attention was aroused by the cue tone, and facilitated subsequent auditory information processing. Experiment 3 examined whether both cue position and frequency influence the resulting information processing. The same effect of spatial attention was observed, but the effect of attention to a certain frequency was only partially observed. In Experiment 4, a tone fusion judgment task was performed to examine whether the effect of spatial attention occurred in the initial stages of hearing. The result suggests that the effect occurred in the later stages of hearing.     

Immature cortical responses to auditory stimuli in specific language impairment: evidence from ERPs to rapid tone sequences

Event-related potentials (ERPs) to tone pairs and single tones were measured for 16 participants with specific language impairment (SLI) and 16 age-matched controls aged from 10 to 19 years The tone pairs were separated by an inter-stimulus interval (ISI) of 20, 50 or 150 ms. The intraclass correlation (ICC) was computed for each participant between the ERP to a single tone and the ERP to the tone pair. A high ICC indicates that the brain response to a tone pair is similar to that for a single tone. ICCs were significantly higher at short than at long ISIs. At 50-ms ISI, ICCs were higher for younger than older participants. Age and ISI interacted with SLI status: ERPs of older participants with SLI differed from age-matched controls, and resembled ERPs of younger controls, consistent with a theory of immature auditory processing in SLI.     

Learning strategy determines auditory cortical plasticity

Learning modifies the primary auditory cortex (A1) to emphasize the processing and representation of behaviorally relevant sounds. However, the factors that determine cortical plasticity are poorly understood. While the type and amount of learning are assumed to be important, the actual strategies used to solve learning problems might be critical. To investigate this possibility, we trained two groups of adult male Sprague-Dawley rats to bar-press (BP) for water contingent on the presence of a 5.0 kHz tone using two different strategies: BP during tone presence or BP from tone-onset until receiving an error signal after tone cessation. Both groups achieved the same high levels of correct performance and both groups revealed equivalent learning of absolute frequency during training. Post-training terminal "mapping" of A1 showed no change in representational area of the tone signal frequency but revealed other substantial cue-specific plasticity that developed only in the tone-onset-to-error strategy group. Threshold was decreased approximately 10 dB and tuning bandwidth was narrowed by approximately 0.7 octaves. As sound onsets have greater perceptual weighting and cortical discharge efficacy than continual sound presence, the induction of specific learning-induced cortical plasticity may depend on the use of learning strategies that best exploit cortical proclivities. The present results also suggest a general principle for the induction and storage of plasticity in learning, viz., that the representation of specific acquired information may be selected by neurons according to a match between behaviorally selected stimulus features and circuit/network response properties.     

Generalization of free-operant avoidance behavior in pigeons

Three groups of four pigeons, trained to press a treadle on a free-operant avoidance schedule, were given auditory discrimination training. Alternating 2-min components of avoidance and no shock were paired with either a tone or white noise. The pigeons were subsequently given two types of generalization tests, with and without avoidable shocks scheduled. Two of the groups, trained interdimensionally, produced excitatory and inhibitory generalization gradients along the tone frequency dimension. A predicted post-discrimination gradient was computed from the algebraic summation of these gradients of excitation and inhibition. The predicted gradient was compared with the actual post-discrimination gradient obtained from the third group of pigeons that had been given intradimensional discrimination training on the tone frequency dimension. The predicted postdiscrimination gradient agreed in shape with the empirical postdiscrimination gradient. The results in general support Spence's (1937) gradient interaction theory.     

Auditory ERP in extremely premature 5-year-old children

The aim of this research was to test the hypothesis of a functional impairment in the automatic detection of deviant tones in 141 children born after 25 to 28 weeks of gestational age, as compared to 45 age-matched full-term control children. All of them were assessed at age 5 years 9 months and instructed to listen passively to two different pure tones (1000 vs 1200 Hz; 20 vs 80%) counterbalanced between ears. Rarity was thus defined by specific ear by tone combinations. The temporal N100 showed a clear contralateral functional organization of the central auditory pathway, especially for the left ear, but without group difference. By contrast, in full-term controls but not in premature children, the central N200 was specifically increased over frontal leads to rare stimuli as compared to frequent. Premature children demonstrated a lack of brain response when more complex processing integrating different informations was required.     

Temporal preparation decreases perceptual latency: Evidence from a clock paradigm

A clock paradigm was employed to assess whether temporal preparation decreases the time to detect the onset of a stimulus—that is, perceptual latency. In four experiments participants watched a revolving clock hand while listening to soft or loud target tones under high or low temporal preparation. At the end of each trial, participants reported the clock hand position at the onset of the target tone. The deviation of the reported clock hand position from the actual position indexed perceptual latency. As expected, perceptual latency decreased with target tone intensity. Most importantly, however, greater temporal preparation decreased perceptual latency in all four experiments, especially for soft tones, which supports rather directly the idea that temporal preparation diminishes the duration of perceptual processing.     

Age‐related changes in transient and oscillatory brain responses to auditory stimulation during early adolescence

Maturational changes in the capacity to process quickly the temporal envelope of sound have been linked to language abilities in typically developing individuals. As part of a longitudinal study of brain maturation and cognitive development during adolescence, we employed dense‐array EEG and spatiotemporal source analysis to characterize maturational changes in the timing of brain responses to temporal variations in sound. We found significant changes in the brain responses compared longitudinally at two time points in early adolescence, namely 10 years (65 subjects) and 11.5 years (60 of the 65 subjects), as well as large differences between adults, studied with the same protocol ( Poulsen, Picton & Paus, 2007 ), and the children at 10 and 11.5 years of age. The transient auditory evoked potential to tone onset showed decreases in the latency of vertex and T‐complex components, and a highly significant increase in the amplitude of the N1 wave with increasing age. The auditory steady state response to a 40‐Hz frequency‐modulated tone increased in amplitude with increasing age. The peak frequency of the envelope‐following response to sweeps of amplitude‐modulated white noise also increased significantly with increasing age. These results indicate persistent maturation of the cortical mechanisms for auditory processing from childhood into middle adulthood.     

Auditory habituation in the fetus and neonate: an fMEG study

Habituation – the most basic form of learning – is used to evaluate central nervous system (CNS) maturation and to detect abnormalities in fetal brain development. In the current study, habituation, stimulus specificity and dishabituation of auditory evoked responses were measured in fetuses and newborns using fetal magnetoencephalography (fMEG). An auditory habituation paradigm consisting of 100 trains of five 500 Hz tones, one 750 Hz tone (dishabituator) and two more 500 Hz tones, respectively, were presented to 41 fetuses (gestational age 30–39 weeks) and 22 newborns or babies (age 6–89 days). A response decrement between the first and fifth tones (habituation), an increment between the fifth tone and the dishabituator (stimulus specificity) and an increment between the fifth (last tone before the dishabituator) and seventh tones (first tone after the dishabituator) (dishabituation) were expected. Fetuses showed weak responses to the first tone. However, a significant response decrement between the second and fifth tones (habituation) and a significant increment between the fifth tone and the dishabituator (stimulus specificity) were found. No significant difference was found for dishabituation nor was a developmental trend found at the group level. From the neonatal data, significant values for stimulus specificity were found. Sensory fatigue or adaptation was ruled out as a reason for the response decrement due to the strong reactions to the dishabituator. Taken together, the current study used fMEG to directly show fetal habituation and provides evidence of fetal learning in the last trimester of pregnancy.     

On insirumental response interaction as explaining the influences of Pavlovian CSs upon avoidance behavior

The finding that Pavlovian signals for food or shock influence avoidance responding might be explained either by interaction of conditioned central mediational states or interaction of learned instrumental responses. Using three groups of dogs, the two hypotheses were pited one against the other in a three-stage transfer-of-control experiment. In the initial conditioning phase, tones were established as signals for food, shock, or neither; additionally the tones also cued a common instrumental response. Following avoidance training, the tone was tested for its influence upon avoidance. If the lone had signaled food, avoidance was suppressed; if shock, avoidance was facilitated; if neither, avoidance was unaffected. This was interpreted as supporting the hypothesis that interaction of central states mediates the transfer-of-control.     

非言语声音影响汉语听者言语声音的知觉

采用启动范式, 以汉语听者为被试, 考察了非言语声音是否影响言语声音的知觉。实验1考察了纯音对辅音范畴连续体知觉的影响, 结果发现纯音影响到辅音范畴连续体的知觉, 表现出频谱对比效应。实验2考察了纯音和复合音对元音知觉的影响, 结果发现与元音共振峰频率一致的纯音或复合音加快了元音的识别, 表现出启动效应。两个实验一致发现非言语声音能够影响言语声音的知觉, 表明言语声音知觉也需要一个前言语的频谱特征分析阶段, 这与言语知觉听觉理论的观点一致。     

Long-term consolidation and retention of learning-induced tuning plasticity in the auditory cortex of the guinea pig.

The major goal of this study was to determine whether classical conditioning produces long-term neural consolidation of frequency tuning plasticity in the auditory cortex. Local field potentials (LFPs) were obtained from chronically implanted adult male Hartley guinea pigs that were divided into conditioning (n = 4) and sensitization control (n = 3) groups. Tuning functions were determined in awake subjects for average LFPs (approximately 0.4 to 36.0 kHz, -20 to 80 dB) immediately before training as well as 1 h and 1, 3, 7, and 10 days after training; sensitization subjects did not have a 10-day retention test. Conditioning consisted of a single session of 30 to 45 trials of a 6-s tone (CS, 70 dB) that was not the best frequency (BF, peak of a tuning curve), followed by a brief leg shock (US) at CS offset. Sensitization control animals received the same density of CS and US presentations unpaired. Heart rate recordings showed that the conditioning group developed conditioned bradycardia, whereas the sensitization control group did not. Local field potentials in the conditioning group, but not in the sensitization group, developed tuning plasticity. The ratio of responses to the CS frequency versus the BF were increased 1 h after training, and this increase was retained for the 10-day period of the study. Both tuning plasticity and retention were observed across stimulus levels (10-80 dB). Most noteworthy, tuning plasticity exhibited consolidation (i.e., developed greater CS-specific effects across retention periods), attaining asymptote at 3 days. The findings indicate that LFPs in the auditory cortex have three cardinal features of behavioral memory: associative tuning plasticity, long-term retention, and long-term consolidation. Potential cellular and subcellular mechanisms of LFP tuning plasticity and long-term consolidation are discussed.     

Differential pavlovian fear conditioning as a function of the qualitative nature of the UCS: Constant versus pulsating shock

Dogs were trained in a shuttlebox to perform an instrumental avoidance response to a visual signal. Then, while pharmacologically immobilized in a hammock, they received differential Pavlovian fear conditioning with two tones: one paired with a constant shock and one with a pulsating shock. During Pavlovian conditioning EKG’s were recorded. Later, in the shuttlebox, Ss received tests for the transfer of control of avoidance responding. Although no significant heart rate differences were found,Ss hurdled the barrier significantly more quickly in response to the tone previously paired with the pulsating shock than to the tone paired with the constant shock, indicating that pulsating shock results in better fear conditioning.     

Signal clustering modulates auditory cortical activity in humans

Auditory streaming and its relevance to attentional processing was examined using event-related brain potentials (ERPs) in situations facilitating perception of one or two streams of sounds. Subjects listened to sequences of brief tones of three different frequencies presented in random order. In evenly spaced (ES) conditions, the three frequencies were equidistant on the musical scale. In clustered, easy (CE) conditions, the attended frequency was distinct, while the middle and extreme distractor tones were clustered together. In clustered, hard (CH) conditions, the attended frequency was clustered with one of the distractors. The subjects pressed a button in response to occasional target tones of longer duration at a prespecified frequency. The subjects were faster and more accurate in CE conditions than they were in ES conditions, and ERP attention effects were enhanced in amplitude in CE conditions. Conversely, the subjects were slower and less accurate in CH conditions and ERP attention effects were delayed in latency and decreased in amplitude. Clustering effects suggest that the processing of stimuli belonging to the attended stream was promoted and the processing of those falling outside the stream was inhibited. The timing and scalp distribution of clustering-related changes in ERPs suggest that clustering modulates early sensory processing in auditory cortex.     

Feature processing during high-rate auditory selective attention

Auditory event-related brain potentials (ERPs) and reaction times were, analyzed in a selective attention task in which subjects attended to tone pips presented at high rates-Xinterstimulua intervals [ISIs] of 40-200 msec). Subjects responded to infrequent target tones of a specified frequency (250 or 4000 Hz) and location (left or right ear) that were louder than otherwise identical tones presented randomly to the left and right ears. Negative difference (Nd) waves were isolated by subtracting ERPs to tones with no target features from ERPs to the same tones when they shared target location, frequency, or both frequency and location cues. Nd waves began 60-70 msec after tone onset and lasted until 252–350 msec after tone onset, even for tones with single attended cues. The duration of Nd waves exceeded the ISIs between successive tones, implying that several stimuli underwent concurrent analysis. Nd waves associated with frequency processing had scalp distributions different from those associated with location processing, implying that the features were analyzed in distinct cortical areas. Nd waves specific to auditory feature conjunction were isolated. These began at latencies of 110–120 msec, some 30-40 msec after the Nds to single features. The relative timing of the different Nd waves suggests that auditory feaure conjunction begins after a brief parallel analysis of individual features but before feature analysis is complete.