Evidence that anterograde learning interference depends on the stage of learning of the interferer: blocked versus interleaved training.

Training on one task (task A) can disrupt learning on a subsequently trained task (task B), illustrating anterograde learning interference. We asked whether the induction of anterograde learning interference depends on the learning stage that task A has reached when the training on task B begins. To do so, we drew on previous observations in perceptual learning in which completing all training on one task before beginning training on another task (blocked training) yielded markedly different learning outcomes than alternating training between the same two tasks for the same total number of trials (interleaved training). Those blocked versus interleaved contrasts suggest that there is a transition between two differentially vulnerable learning stages that is related to the number of consecutive training trials on each task, with interleaved training presumably tapping acquisition, and blocked training tapping consolidation. Here, we used the blocked versus interleaved paradigm in auditory perceptual learning in a case in which blocked training generated anterograde-but not its converse, retrograde-learning interference (A→B, not B←A). We report that anterograde learning interference of training on task A (interaural time difference discrimination) on learning on task B (interaural level difference discrimination) occurred with blocked training and diminished with interleaved training, with faster rates of interleaving leading to less interference. This pattern held for across-day, within-session, and offline learning. Thus, anterograde learning interference only occurred when the number of consecutive training trials on task A surpassed some critical value, consistent with other recent evidence that anterograde learning interference only arises when learning on task A has entered the consolidation stage.

Discrimination thresholds for interaural-time differences and interaural-level differences in naïve listeners: Sex differences and learning.

The two primary cues to sound-source location on the horizontal plane are interaural time differences (ITDs) and interaural level differences (ILDs). Here we asked whether the ability to discriminate small changes in each of these interaural cues differs between the sexes. We tested one group of males (n = 43) and females (n = 94) on ITD discrimination at 0.5 kHz and a separate group of males (n = 80) and females (n = 166) on ILD discrimination at 4 kHz. None of the participants had any prior experience with psychoacoustic tasks. Testing of each participant was completed in a single testing session of 4-5 blocks of 60 trials. For ILD discrimination, the overall mean threshold, as well as the mean threshold for each block, was statistically significantly lower for males than for females. Despite that, males and females learned at an equal rate over the course of testing. For ITD discrimination, in contrast, thresholds did not differ significantly between the sexes for the overall mean or for any block. There also was no statistically significant learning across blocks for either sex. For both tasks and both sexes, the individual thresholds spanned a wide range. The presence of a statistically significant sex difference and learning for ILD but not for ITD discrimination, along with a larger effect size for ILD than for ITD discrimination, suggests that the factors responsible for these outcomes acted upon an ILD-specific neural pathway, and not upon an ITD-specific pathway, nor any pathway common to the two cues. Because the ILD and ITD specific pathways are most separable initially, the factors associated with sex and learning may have acted upon the ILD-specific pathway at an early stage.

Non-sensory Influences on Auditory Learning and Plasticity.

Distinguishing between regular and irregular heartbeats, conversing with speakers of different accents, and tuning a guitar-all rely on some form of auditory learning. What drives these experience-dependent changes? A growing body of evidence suggests an important role for non-sensory influences, including reward, task engagement, and social or linguistic context. This review is a collection of contributions that highlight how these non-sensory factors shape auditory plasticity and learning at the molecular, physiological, and behavioral level. We begin by presenting evidence that reward signals from the dopaminergic midbrain act on cortico-subcortical networks to shape sound-evoked responses of auditory cortical neurons, facilitate auditory category learning, and modulate the long-term storage of new words and their meanings. We then discuss the role of task engagement in auditory perceptual learning and suggest that plasticity in top-down cortical networks mediates learning-related improvements in auditory cortical and perceptual sensitivity. Finally, we present data that illustrates how social experience impacts sound-evoked activity in the auditory midbrain and forebrain and how the linguistic environment rapidly shapes speech perception. These findings, which are derived from both human and animal models, suggest that non-sensory influences are important regulators of auditory learning and plasticity and are often implemented by shared neural substrates. Application of these principles could improve clinical training strategies and inform the development of treatments that enhance auditory learning in individuals with communication disorders.

Humans attend to signal duration but not temporal structure for sound detection: Steady-state versus pulse-train signals.

Most sounds fluctuate in amplitude, but do listeners attend to the temporal structure of those fluctuations when trying to detect the mere presence of those sounds? This question was addressed by leading listeners to expect a faint sound with a fixed temporal structure (pulse train or steady-state tone) and total duration (300 ms) and measuring their ability to detect equally faint sounds of unexpected temporal structure (pulse train when expecting steady state) and/or total duration (<300 ms). Detection was poorer for sounds with unexpected than with expected total durations, replicating previous outcomes, but was uninfluenced by the temporal structure of the expected sound. The results disagree with computational predictions of the multiple-look model, which posits that listeners attend to both the total duration and temporal structure of the signal, but agree with predictions of the matched-window energy-detector model, which posits that listeners attend to the total duration but not the temporal structure of the signal. Moreover, the matched-window energy-detector model could also account for previous results, including some that were originally interpreted as supporting the multiple-look model. Taken together, at least when detecting faint sounds, listeners appear to attend to the total duration of expected sounds but to ignore their detailed temporal structure.

Extended high frequency hearing and speech perception implications in adults and children.

Extended high frequencies (EHF), above 8 kHz, represent a region of the human hearing spectrum that is generally ignored by clinicians and researchers alike. This article is a compilation of contributions that, together, make the case for an essential role of EHF in both normal hearing and auditory dysfunction. We start with the fundamentals of biological and acoustic determinism - humans have EHF hearing for a purpose, for example, the detection of prey, predators, and mates. EHF hearing may also provide a boost to speech perception in challenging conditions and its loss, conversely, might help explain difficulty with the same task. However, it could be that EHF are a marker for damage in the conventional frequency region that is more related to speech perception difficulties. Measurement of EHF hearing in concert with otoacoustic emissions could provide an early warning of age-related hearing loss. In early life, when EHF hearing sensitivity is optimal, we can use it for enhanced phonetic identification during language learning, but we are also susceptible to diseases that can prematurely damage it. EHF audiometry techniques and standardization are reviewed, providing evidence that they are reliable to measure and provide important information for early detection, monitoring and possible prevention of hearing loss in populations at-risk. To better understand the full contribution of EHF to human hearing, clinicians and researchers can contribute by including its measurement, along with measures of speech in noise and self-report of hearing difficulties and tinnitus in clinical evaluations and studies.

Semi-supervised learning of a nonnative phonetic contrast: How much feedback is enough?

Semi-supervised learning refers to learning that occurs when feedback about performance is provided on only a subset of training trials. Algorithms for semi-supervised learning are popular in machine learning because of their minimal reliance on labeled data. There have been, however, only a few reports of semi-supervised learning in humans. Here we document human semi-supervised learning on a nonnative phonetic classification task. Classification performance remained unchanged when 60 feedback trials were provided on each of the two days of training. In contrast, performance improved when 60 feedback trials were combined with 240 no-feedback trials each day. In variants of this successful semi-supervised regimen, increasing the daily number of feedback trials from 60 to 240 did not increase the amount of learning, while decreasing that number to 30 abolished learning. Finally, replacing the no-feedback trials with stimulus exposure alone had little effect on the outcome. These results were an unexpected consequence of combining training periods with feedback and testing periods without feedback, illustrating that no-feedback testing can influence learning outcomes. More broadly, these data suggest that task performance with feedback can function as an all-or-none trigger for recruiting the contribution of trials without feedback, or mere stimulus exposures, to human learning.

Perceptual-learning evidence for inter-onset-interval- and frequency-specific processing of fast rhythms.

Rhythm is fundamental to music and speech, yet little is known about how even simple rhythmic patterns are processed. Here we investigated the processing of isochronous rhythms in the short inter-onset-interval (IOI) range (IOIs < 250-400 ms) using a perceptual-learning paradigm. Trained listeners (n=8) practiced anisochrony detection with a 100-ms IOI marked by 1-kHz tones, 720 trials per day for 7 days. Between pre- and post-training tests, trained listeners improved significantly more than controls (no training; n=8) on the anisochrony-detection condition that the trained listeners practiced. However, the learning on anisochrony detection did not generalize to temporal-interval discrimination with the trained IOI (100 ms) and marker frequency (1 kHz) or to anisochrony detection with an untrained marker frequency (4 kHz or variable frequency vs. 1 kHz), and generalized negatively to anisochrony detection with an untrained IOI (200 ms vs. 100 ms). Further, pre-training thresholds were correlated among nearly all of the conditions with the same IOI (100-ms IOIs), but not between conditions with different IOIs (100-ms vs. 200-ms IOIs). Thus, it appears that some task-, IOI-, and frequency-specific processes are involved in fast-rhythm processing. These outcomes are most consistent with a holistic rhythm-processing model in which a holistic "image" of the stimulus is compared to a stimulus-specific template.

Inducing musical-interval learning by combining task practice with periods of stimulus exposure alone.

A key component of musical proficiency is the ability to discriminate between and identify musical intervals, or fixed ratios between pitches. Acquiring these skills requires training, but little is known about how to best arrange the trials within a training session. To address this issue, learning on a musical-interval comparison task was evaluated for two four-day training regimens that employed equal numbers of stimulus presentations per day. A regimen of continuous practice yielded no learning, but a regimen that combined practice and stimulus exposure alone generated clear improvement. Learning in the practice-plus-exposure regimen was due to the combination of the two experiences, because two control groups who received only either the practice or the exposure from that regimen did not learn. Posttest performance suggested that this improvement in comparison learning generalized to an untrained stimulus type and an untrained musical-interval identification task. Naïve comparison performance, but not learning, was better for larger pitch-ratio differences and for individuals with more musical experience. The reported benefits of the practice-plus-exposure regimen mirror the outcomes for fine-grained discrimination and speech tasks, suggesting that a general learning principle is involved. In practical terms, it appears that combining practice and stimulus exposure alone is a particularly effective configuration for improving musical-interval perception.

The Value of Homework: Exposure to Odors in the Home Cage Enhances Odor-Discrimination Learning in Mice.

Perceptual learning is an enhancement in discriminability of similar stimuli following experience with those stimuli. Here, we examined the efficacy of adding additional active training following a standard training session, compared with additional stimulus exposure in the absence of associated task performance. Mice were trained daily in an odor-discrimination task, and then, several hours later each day, received 1 of 3 different manipulations: 1) a second active-training session, 2) non-task-related odor exposure in the home cage, or 3) no second session. For home-cage exposure, odorants were presented in small tubes that mice could sniff and investigate for a similar period of time as in the active discrimination task each day. The results demonstrate that daily home-cage exposure was equivalent to active odor training in supporting improved odor discrimination. Daily home-cage exposure to odorants that did not match those used in the active task did not improve learning, yielding outcomes similar to those obtained with no second session. Piriform cortical local field potential recordings revealed that both sampling in the active learning task and investigation in the home cage evoked similar beta band oscillatory activity. Together the results suggest that odor-discrimination learning can be significantly enhanced by addition of odor exposure outside of the active training task, potentially because of the robust activity evoked in the olfactory system by both exposure paradigms. They further suggest that odorant exposure alone could enhance or maintain odor-discrimination abilities in conditions associated with olfactory impairment, such as aging or dementia.

Transient sex differences during adolescence on auditory perceptual tasks.

Many perceptual abilities differ between the sexes. Because these sex differences have been documented almost exclusively in adults, they have been attributed to sex-specific neural circuitry that emerges during development and is maintained in the mature perceptual system. To investigate whether behavioral sex differences in perception can also have other origins, we compared performance between males and females ranging in age from 8 to 30 years on auditory temporal-interval discrimination and tone-in-noise detection tasks on which there are no sex differences in adults. If sex differences in perception arise only from the establishment and subsequent maintenance of sex-specific neural circuitry, there should be no sex differences during development on these tasks. In contrast, sex differences emerged in adolescence but resolved by adulthood on two of the six conditions, with signs of a similar pattern on a third condition. In each case, males reached mature performance earlier than females, resulting in a sex difference in the interim. These results suggest that sex differences in perception may arise from differences in the maturational timing of common circuitry used by both sexes. They also imply that sex differences in perceptual abilities may be more prevalent than previously thought based on adult data alone.

Disruption of Perceptual Learning by a Brief Practice Break.

Some forms of associative learning require only a single experience to create a lasting memory [1, 2]. In contrast, perceptual learning often requires extensive practice within a day for performance to improve across days [3, 4]. This suggests that the requisite practice for durable perceptual learning is integrated throughout each day. If the total amount of daily practice is the only important variable, then a practice break within a day should not disrupt across-day improvement. To test this idea, we trained human listeners on an auditory frequency-discrimination task over multiple days and compared the performance of those who engaged in a single continuous practice session each day [4] with those who were given a 30-min break halfway through each practice session. Continuous practice yielded significant perceptual learning [4]. In contrast, practice with a rest break led to no improvement, indicating that the integration process had decayed within 30 min. In a separate experiment, a 30-min practice break also disrupted durable learning on a non-native phonetic classification task. These results suggest that practice trials are integrated up to a learning threshold within a transient memory store before they are sent en masse into a memory that lasts across days. Thus, the oft cited benefits of distributed over massed training [5, 6] may arise from different mechanisms depending on whether the breaks occur before or after a learning threshold has been reached. Trial integration could serve as an early gatekeeper to plasticity, helping to ensure that longer-lasting changes are only made when deemed worthwhile.

Detection of tones of unexpected frequency in amplitude-modulated noise.

Detection of a tonal signal in amplitude-modulated noise can improve with increases in noise bandwidth if the pattern of amplitude fluctuations is uniform across frequency, a phenomenon termed comodulation masking release (CMR). Most explanations for CMR rely on an assumption that listeners monitor frequency channels both at and remote from the signal frequency in conditions that yield the effect. To test this assumption, detectability was assessed for signals presented at expected and unexpected frequencies in wideband amplitude-modulated noise. Detection performance was high even for signals of unexpected frequency, suggesting that listeners were monitoring multiple frequency channels, as has been assumed.

Enhancing speech learning by combining task practice with periods of stimulus exposure without practice.

Language acquisition typically involves periods when the learner speaks and listens to the new language, and others when the learner is exposed to the language without consciously speaking or listening to it. Adaptation to variants of a native language occurs under similar conditions. Here, speech learning by adults was assessed following a training regimen that mimicked this common situation of language immersion without continuous active language processing. Experiment 1 focused on the acquisition of a novel phonetic category along the voice-onset-time continuum, while Experiment 2 focused on adaptation to foreign-accented speech. The critical training regimens of each experiment involved alternation between periods of practice with the task of phonetic classification (Experiment 1) or sentence recognition (Experiment 2) and periods of stimulus exposure without practice. These practice and exposure periods yielded little to no improvement separately, but alternation between them generated as much or more improvement as did practicing during every period. Practice appears to serve as a catalyst that enables stimulus exposures encountered both during and outside of the practice periods to contribute to quite distinct cases of speech learning. It follows that practice-plus-exposure combinations may tap a general learning mechanism that facilitates language acquisition and speech processing.

A late-emerging auditory deficit in autism.

OBJECTIVE: Individuals with autism spectrum disorders (ASD) show enhanced perceptual and memory abilities in the domain of pitch, but also perceptual deficits in other auditory domains. The present study investigated their skills with respect to "echoic memory," a form of short-term sensory memory intimately tied to auditory perception, using a developmental perspective. METHOD: We tested 23 high-functioning participants with ASD and 26 typically developing (TD) participants, distributed in two age groups (children vs. young adults; mean ages: ∼11 and ∼21 years). By means of an adaptive psychophysical procedure, we measured the longest period for which periodic (i.e., repeated) noise could be reliably discriminated from nonperiodic (i.e., plain random) noise. On each experimental trial, a single noise sample was presented to the participant, who had to classify this sound as periodic or nonperiodic. RESULTS: The TD adults performed, on average, much better than the other three groups, who performed similarly overall. As a function of practice, the measured thresholds improved for the TD participants, but did not change for the ASD participants. Thresholds were not correlated to performance in a test assessing verbal memory. The variance of the participants' response biases was larger among the ASD participants than among the TD participants. CONCLUSION: The results mainly suggest that echoic memory takes a long time to fully develop in TD humans, and that this development stops prematurely in persons with ASD.

Learning one task by interleaving practice with another task.

Perceptual learning is a sustainable improvement in performance on a perceptual task following training. A hallmark of perceptual learning is task specificity - after participants have trained on and learned a particular task, learning rarely transfers to another task, even with identical stimuli. Accordingly, it is assumed that performing a task throughout training is a requirement for learning to occur on that specific task. Thus, interleaving training trials of a target task, with those of another task, should not improve performance on the target task. However, recent findings in audition show that interleaving two tasks during training can facilitate perceptual learning, even when the training on neither task yields learning on its own. Here we examined the role of cross-task training in the visual domain by training 4 groups of human observers for 3 consecutive days on an orientation comparison task (target task) and/or spatial-frequency comparison task (interleaving task). Interleaving small amounts of training on each task, which were ineffective alone, not only enabled learning on the target orientation task, as in audition, but also surpassed the learning attained by training on that task alone for the same total number of trials. This study illustrates that cross-task training in visual perceptual learning can be more effective than single-task training. The results reveal a comparable learning principle across modalities and demonstrate how to optimize training regimens to maximize perceptual learning.

Learning, worsening, and generalization in response to auditory perceptual training during adolescence.

While it is commonly held that the capacity to learn is greatest in the young, there have been few direct comparisons of the response to training across age groups. Here, adolescents (11-17 years, n = 20) and adults (≥18 years, n = 11) practiced detecting a backward-masked tone for ∼1 h/day for 10 days. Nearly every adult, but only half of the adolescents improved across sessions, and the adolescents who learned did so more slowly than adults. Nevertheless, the adolescent and adult learners showed the same generalization pattern, improving on untrained backward- but not forward- or simultaneous-masking conditions. Another subset of adolescents (n = 6) actually got worse on the trained condition. This worsening, unlike learning, generalized to an untrained forward-masking, but not backward-masking condition. Within sessions, both age groups got worse, but the worsening was greater for adolescents. These maturational changes in the response to training largely followed those previously reported for temporal-interval discrimination. Overall, the results suggest that late-maturing processes affect the response to perceptual training and that some of these processes may be shared between tasks. Further, the different developmental rates for learning and generalization, and different generalization patterns for learning and worsening imply that learning, generalization, and worsening may have different origins.

Accent-independent adaptation to foreign accented speech.

Foreign-accented speech can be difficult to understand but listeners can adapt to novel talkers and accents with appropriate experience. Previous studies have demonstrated talker-independent but accent-dependent learning after training on multiple talkers from a single language background. Here, listeners instead were exposed to talkers from five language backgrounds during training. After training, listeners generalized their learning to novel talkers from language backgrounds both included and not included in the training set. These findings suggest that generalization of foreign-accent adaptation is the result of exposure to systematic variability in accented speech that is similar across talkers from multiple language backgrounds.

Different patterns of perceptual learning on spectral modulation detection between older hearing-impaired and younger normal-hearing adults.

Young adults with normal hearing (YNH) can improve their sensitivity to basic acoustic features with practice. However, it is not known to what extent the influence of the same training regimen differs between YNH listeners and older listeners with hearing impairment (OHI)--the largest population seeking treatment in audiology clinics. To examine this issue, we trained OHI listeners on a basic auditory task (spectral modulation detection) using a training regimen previously administered to YNH listeners (≈ 1 h/session for seven sessions on a single condition). For the trained conditions on which pretraining performance was not already at asymptote, the YNH listeners who received training learned more than matched controls who received none, but that learning did not generalize to any untrained spectral modulation frequency. In contrast, the OHI-trained listeners and controls learned similar amounts on the trained condition, implying no effect of the training itself. However, surprisingly the OHI-trained listeners improved over the training phase and on an untrained spectral modulation frequency. These population differences suggest that learning consolidated more slowly, and that training modified an aspect of processing that had broader tuning to spectral modulation frequency, in OHI than YNH listeners. More generally, these results demonstrate that conclusions about perceptual learning that come from examination of one population do not necessarily apply to another.

Perceptual learning evidence for tuning to spectrotemporal modulation in the human auditory system.

Natural sounds are characterized by complex patterns of sound intensity distributed across both frequency (spectral modulation) and time (temporal modulation). Perception of these patterns has been proposed to depend on a bank of modulation filters, each tuned to a unique combination of a spectral and a temporal modulation frequency. There is considerable physiological evidence for such combined spectrotemporal tuning. However, direct behavioral evidence is lacking. Here we examined the processing of spectrotemporal modulation behaviorally using a perceptual-learning paradigm. We trained human listeners for ∼1 h/d for 7 d to discriminate the depth of spectral (0.5 cyc/oct; 0 Hz), temporal (0 cyc/oct; 32 Hz), or upward spectrotemporal (0.5 cyc/oct; 32 Hz) modulation. Each trained group learned more on their respective trained condition than did controls who received no training. Critically, this depth-discrimination learning did not generalize to the trained stimuli of the other groups or to downward spectrotemporal (0.5 cyc/oct; -32 Hz) modulation. Learning on discrimination also led to worsening on modulation detection, but only when the same spectrotemporal modulation was used for both tasks. Thus, these influences of training were specific to the trained combination of spectral and temporal modulation frequencies, even when the trained and untrained stimuli had one modulation frequency in common. This specificity indicates that training modified circuitry that had combined spectrotemporal tuning, and therefore that circuits with such tuning can influence perception. These results are consistent with the possibility that the auditory system analyzes sounds through filters tuned to combined spectrotemporal modulation.

Perceptual learning of auditory spectral modulation detection.

Normal sensory perception requires the ability to detect and identify patterns of activity distributed across the receptor surface. In the visual system, the ability to perceive these patterns across the retina improves with training. This learning differs in magnitude for different trained stimuli and does not generalize to untrained spatial frequencies or retinal locations. Here, we asked whether training to detect patterns of activity across the cochlea yields learning with similar characteristics. Differences in learning between the visual and auditory systems would be inconsistent with the suggestion that the ability to detect these patterns is limited by similar constraints in these two systems. We trained three groups of normal-hearing listeners to detect spectral envelopes with a sinusoidal shape (spectral modulation) at 0.5, 1, or 2 cycles/octave and compared the performance of each group to that of a separate group that received no training. On average, as the trained spectral modulation frequency increased, the magnitude of training-induced improvement and the time to reach asymptotic performance decreased, while the tendency for performance to worsen within a training session increased. The training-induced improvements did not generalize to untrained spectral modulation frequencies or untrained carrier spectra. Thus, for both visual-spatial and auditory spectral modulation detection, learning depended upon and was specific to analogous features of the trained stimulus. Such similarities in learning could arise if, as has been suggested, similar constraints limit the ability to detect patterns across the receptor surface between the auditory and visual systems.