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.

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.

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.

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.

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.

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.

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.

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.

Generalization lags behind learning on an auditory perceptual task.

The generalization of learning from trained to untrained conditions is of great potential value because it markedly increases the efficacy of practice. In principle, generalization and the learning itself could arise from either the same or distinct neural changes. Here, we assessed these two possibilities in the realm of human perceptual learning by comparing the time course of improvement on a trained condition (learning) to that on an untrained condition (generalization) for an auditory temporal-interval discrimination task. While significant improvement on the trained condition occurred within 2 d, generalization to the untrained condition lagged behind, only emerging after 4 d. The different time courses for learning and generalization suggest that these two types of perceptual improvement can arise from at least partially distinct neural changes. The notably longer time course for generalization than learning demonstrates that increasing the duration of training can be an effective means to increase the number of conditions to which learning generalizes on perceptual tasks.

Enhancing perceptual learning by combining practice with periods of additional sensory stimulation.

Perceptual skills can be improved even in adulthood, but this learning seldom occurs by stimulus exposure alone. Instead, it requires considerable practice performing a perceptual task with relevant stimuli. It is thought that task performance permits the stimuli to drive learning. A corresponding assumption is that the same stimuli do not contribute to improvement when encountered separately from relevant task performance because of the absence of this permissive signal. However, these ideas are based on only two types of studies, in which the task was either always performed or not performed at all. Here we demonstrate enhanced perceptual learning on an auditory frequency-discrimination task in human listeners when practice on that target task was combined with additional stimulation. Learning was enhanced regardless of whether the periods of additional stimulation were interleaved with or provided exclusively before or after target-task performance, and even though that stimulation occurred during the performance of an irrelevant (auditory or written) task. The additional exposures were only beneficial when they shared the same frequency with, though they did not need to be identical to, those used during target-task performance. Their effectiveness also was diminished when they were presented 15 min after practice on the target task and was eliminated when that separation was increased to 4 h. These data show that exposure to an acoustic stimulus can facilitate learning when encountered outside of the time of practice on a perceptual task. By properly using additional stimulation one may markedly improve the efficiency of perceptual training regimens.

Late maturation of auditory perceptual learning.

Adults can improve their performance on many perceptual tasks with training, but when does the response to training become mature? To investigate this question, we trained 11-year-olds, 14-year-olds and adults on a basic auditory task (temporal-interval discrimination) using a multiple-session training regimen known to be effective for adults. The adolescents all began with performance in the adult range. However, while all of the adults improved across sessions, none of the 11-year-olds and only half of the 14-year-olds did. The adolescents who failed to learn did so even though the 10-session training regimen provided twice the number of sessions required by adults to reach asymptotic performance. Further, over the course of each session, the performance of the adults was stable but that of the adolescents, including those who learned, deteriorated. These results demonstrate that the processes that underlie perceptual learning can continue to develop well into adolescence.

Auditory filter shapes and high-frequency hearing in adults who have impaired speech in noise performance despite clinically normal audiograms.

Some individuals complain of hearing difficulties in the presence of background noise even in the absence of clinically significant hearing loss (obscure auditory dysfunction). Previous evidence suggests that these listeners have impaired frequency resolution, but there has been no thorough characterization of auditory filter shapes in this population. Here, the filter shapes of adults (n = 14) who self-reported speech recognition problems in noise and performed poorly on a sentence-in-noise perception test despite having clinically normal audiograms were compared to those of controls (n = 10). The filter shapes were evaluated using a 2-kHz probe with a fixed level of 30, 40, or 50 dB sound pressure level (SPL) and notched-noise simultaneous maskers that were varied in level to determine the masker level necessary to just mask the probe. The filters of the impaired group were significantly wider than those of controls at all probe levels owing to an unusual broadening of the upper slope of the filter. In addition, absolute thresholds were statistically indistinguishable between the groups at the standard audiometric frequencies, but were elevated in the impaired listeners at higher frequencies. These results strengthen the idea that this population has a variety of hearing deficits that go undetected by standard audiometry.

Perceptual learning and generalization resulting from training on an auditory amplitude-modulation detection task.

Fluctuations in sound amplitude provide important cues to the identity of many sounds including speech. Of interest here was whether the ability to detect these fluctuations can be improved with practice, and if so whether this learning generalizes to untrained cases. To address these issues, normal-hearing adults (n = 9) were trained to detect sinusoidal amplitude modulation (SAM; 80-Hz rate, 3-4 kHz bandpass carrier) 720 trials/day for 6-7 days and were tested before and after training on related SAM-detection and SAM-rate-discrimination conditions. Controls (n = 9) only participated in the pre- and post-tests. The trained listeners improved more than the controls on the trained condition between the pre- and post-tests, but different subgroups of trained listeners required different amounts of practice to reach asymptotic performance, ranging from 1 (n = 6) to 4-6 (n = 3) sessions. This training-induced learning did not generalize to detection with two untrained carrier spectra (5 kHz low-pass and 0.5-1.5 kHz bandpass) or to rate discrimination with the trained rate and carrier spectrum, but there was some indication that it generalized to detection with two untrained rates (30 and 150 Hz). Thus, practice improved the ability to detect amplitude modulation, but the generalization of this learning to untrained cases was somewhat limited.

Differential rates of consolidation of conceptual and stimulus learning following training on an auditory skill.

Training-induced improvements on perceptual skills can be attributed to at least two learning types: learning of general aspects of the trained condition (conceptual learning) and learning of specific feature values of the stimulus used in training (stimulus learning). Here we asked whether conceptual and stimulus learning on interaural time difference (ITD) discrimination emerge along different time courses. Conceptual learning was clearly evident 10 h after training, when performance on a target ITD condition was equivalent following training on that condition or on a non-target condition differing only in the stimulus, and was better in both cases than immediately after training. In contrast, stimulus learning emerged 24 h after training. At that time, performance on the target ITD condition was better following target- than non-target training, due to a worsening in performance between 10 and 24 h after non-target training rather than from additional improvements over this time period after target training. Training amount influenced performance immediately, but not 10 or 24 h, after training. Thus, conceptual learning emerged before stimulus learning, and each manifested through different improvement trajectories many hours after training. These results suggest that on ITD discrimination, conceptual learning is consolidated earlier, and with different behavioral consequences, than stimulus learning.

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.

Contributions of procedure and stimulus learning to early, rapid perceptual improvements.

Improvements in performance on many perceptual skills can occur with only a single training session. Of interest here is what aspects of the training experience are being learned during this brief exposure. Although there is considerable evidence that learning associated with specific feature values of the stimulus used in training (stimulus learning) contributes to these rapid improvements, there has been little direct investigation of the possibility that other types of learning do so as well. Here the authors show that not only stimulus learning but also learning of more general aspects of the training experience (procedure learning) contributed to rapid improvements in performance on interaural time difference discrimination. However, practice on the type of judgment to be made did not appear to aid performance (no task learning). These results are consistent with physiological reports that different neural mechanisms are engaged at different time points during even a brief training session, and imply that the circuits that are engaged and likely modified vary in the degree of their selectivity to the target condition. Such changes presumably enable further learning.