Effect of age on lateralized auditory processing.

The lateralization of processing in the auditory cortex for different acoustic parameters differs depending on stimuli and tasks. Thus, processing complex auditory stimuli requires an efficient hemispheric interaction. Anatomical connectivity decreases with aging and consequently affects the functional interaction between the left and right auditory cortex and lateralization of auditory processing. Here we studied with magnetic resonance imaging the effect of aging on the lateralization of processing and hemispheric interaction during two tasks utilizing the contralateral noise procedure. Categorization of tones according to their direction of frequency modulations (FM) is known to be processed mainly in the right auditory cortex. Sequential comparison of the same tones according to their FM direction strongly involves additionally the left auditory cortex and therefore a stronger hemispheric interaction than the categorization task. The results showed that older adults more strongly recruit the auditory cortex especially during the comparison task that requires stronger hemispheric interaction. This was the case although the task difficulty was adapted to achieve similar performance as the younger adults. Additionally, functional connectivity from auditory cortex to other brain areas was stronger in older than younger adults especially during the comparison task. Diffusion tensor imaging data showed a reduction in fractional anisotropy and an increase in mean diffusivity in the corpus callosum of older adults compared to younger adults. These changes indicate a reduction of anatomical interhemispheric connections in older adults that makes larger processing capacity necessary when tasks require functional hemispheric interaction.

Dorsal posterior cingulate cortex responds to negative feedback information supporting learning and relearning of response policies.

Many challenges in life come without explicit instructions. Instead, humans need to test, select, and adapt their behavioral responses based on feedback from the environment. While reward-centric accounts of feedback processing primarily stress the reinforcing aspect of positive feedback, feedback's central function from an information-processing perspective is to offer an opportunity to correct errors, thus putting a greater emphasis on the informational content of negative feedback. Independent of its potential rewarding value, the informational value of performance feedback has recently been suggested to be neurophysiologically encoded in the dorsal portion of the posterior cingulate cortex (dPCC). To further test this association, we investigated multidimensional categorization and reversal learning by comparing negative and positive feedback in an event-related functional magnetic resonance imaging experiment. Negative feedback, compared with positive feedback, increased activation in the dPCC as well as in brain regions typically involved in error processing. Only in the dPCC, subarea d23, this effect was significantly enhanced in relearning, where negative feedback signaled the need to shift away from a previously established response policy. Together with previous findings, this result contributes to a more fine-grained functional parcellation of PCC subregions and supports the dPCC's involvement in the adaptation to behaviorally relevant information from the environment.

Neural Mechanisms Underlying the Effects of Novel Sounds on Task Performance in Children With and Without ADHD.

Distractibility is one of the key features of attention deficit hyperactivity disorder (ADHD) and has been associated with alterations in the neural orienting and alerting networks. Task-irrelevant stimuli are thus expected to have detrimental effects on the performance of patients with ADHD. However, task-irrelevant presentation of novel sounds seems to have the opposite effect and improve subsequent attentional performance particularly in patients with ADHD. Here, we aimed to understand the neural modulations of the attention networks underlying these improvements. Fifty boys (25 with ADHD) participated in a functional magnetic resonance imaging (fMRI) study in which unique (novel) or repeatedly presented (familiar) sounds were placed before a visual flanker task in 2/3 of the trials. We found that presenting any sound improved task performance in all participants, but the underlying neural mechanisms differed for the type of sound. Familiar sounds led to a stronger increase in activity in the left posterior insula in patients with ADHD compared to typically developing peers. Novel sounds led to activations of the fronto-temporoparietal ventral attention network, likewise in ADHD and TD. These changes in signaling by novelty in the right inferior frontal gyrus were directly related to improved response speed showing that neural orienting network activity following novel sounds facilitated subsequent attentional performance. This mechanism of behavioral enhancement by short distractions could potentially be useful for cognitive trainings or homework situations.

Why do humans have unique auditory event-related fields? Evidence from computational modeling and MEG experiments.

Auditory event-related fields (ERFs) measured with magnetoencephalography (MEG) are useful for studying the neuronal underpinnings of auditory cognition in human cortex. They have a highly subject-specific morphology, albeit certain characteristic deflections (e.g., P1m, N1m, and P2m) can be identified in most subjects. Here, we explore the reason for this subject-specificity through a combination of MEG measurements and computational modeling of auditory cortex. We test whether ERF subject-specificity can predominantly be explained in terms of each subject having an individual cortical gross anatomy, which modulates the MEG signal, or whether individual cortical dynamics is also at play. To our knowledge, this is the first time that tools to address this question are being presented. The effects of anatomical and dynamical variation on the MEG signal is simulated in a model describing the core-belt-parabelt structure of the auditory cortex, and with the dynamics based on the leaky-integrator neuron model. The experimental and simulated ERFs are characterized in terms of the N1m amplitude, latency, and width. Also, we examine the waveform grand-averaged across subjects, and the standard deviation of this grand average. The results show that the intersubject variability of the ERF arises out of both the anatomy and the dynamics of auditory cortex being specific to each subject. Moreover, our results suggest that the latency variation of the N1m is largely related to subject-specific dynamics. The findings are discussed in terms of how learning, plasticity, and sound detection are reflected in the auditory ERFs. The notion of the grand-averaged ERF is critically evaluated.

Dorsal posterior cingulate cortex encodes the informational value of feedback in human-computer interaction.

In communication between humans as well as in human-computer interaction, feedback is ubiquitous. It is essential for keeping up the dialogue between interaction partners, evaluating the adequacy of an action, or improving task performance. While the neuroscientific view on feedback has largely focused on its function as reward, more general definitions also emphasise its function as information about aspects of one's task performance. Using fMRI in a computer-controlled auditory categorisation task, we studied the neural correlates of the informational value of computer-given feedback independent of reward. Feedback about the correctness of a decision, compared with feedback only indicating the registration of a decision, increases activation of the dorsal posterior cingulate cortex, supporting this region's role in adapting to behaviourally relevant information. Both conditions elicit equally strong activation of the dorsal striatum which does not support an interpretation of feedback information as a type of reward. Instead, we suggest that it reflects a more fundamental aspect of human interaction behaviour, namely the establishment of a state that enables us to continue with the next step of the interaction.

Machine learning identifies the dynamics and influencing factors in an auditory category learning experiment.

Human learning is one of the main topics in psychology and cognitive neuroscience. The analysis of experimental data, e.g. from category learning experiments, is a major challenge due to confounding factors related to perceptual processing, feedback value, response selection, as well as inter-individual differences in learning progress due to differing strategies or skills. We use machine learning to investigate (Q1) how participants of an auditory category-learning experiment evolve towards learning, (Q2) how participant performance saturates and (Q3) how early we can differentiate whether a participant has learned the categories or not. We found that a Gaussian Mixture Model describes well the evolution of participant performance and serves as basis for identifying influencing factors of task configuration (Q1). We found early saturation trends (Q2) and that CatBoost, an advanced classification algorithm, can separate between participants who learned the categories and those who did not, well before the end of the learning session, without much degradation of separation quality (Q3). Our results show that machine learning can model participant dynamics, identify influencing factors of task design and performance trends. This will help to improve computational models of auditory category learning and define suitable time points for interventions into learning, e.g. by tutorial systems.

Predictive cues for auditory stream formation in humans and monkeys.

Auditory perception is improved when stimuli are predictable, and this effect is evident in a modulation of the activity of neurons in the auditory cortex as shown previously. Human listeners can better predict the presence of duration deviants embedded in stimulus streams with fixed interonset interval (isochrony) and repeated duration pattern (regularity), and neurons in the auditory cortex of macaque monkeys have stronger sustained responses in the 60-140 ms post-stimulus time window under these conditions. Subsequently, the question has arisen whether isochrony or regularity in the sensory input contributed to the enhancement of the neuronal and behavioural responses. Therefore, we varied the two factors isochrony and regularity independently and measured the ability of human subjects to detect deviants embedded in these sequences as well as measuring the responses of neurons the primary auditory cortex of macaque monkeys during presentations of the sequences. The performance of humans in detecting deviants was significantly increased by regularity. Isochrony enhanced detection only in the presence of the regularity cue. In monkeys, regularity increased the sustained component of neuronal tone responses in auditory cortex while isochrony had no consistent effect. Although both regularity and isochrony can be considered as parameters that would make a sequence of sounds more predictable, our results from the human and monkey experiments converge in that regularity has a greater influence on behavioural performance and neuronal responses.

The impact of task difficulty on the lateralization of processing in the human auditory cortex.

Perception of complex auditory stimuli like speech requires the simultaneous processing of different fundamental acoustic parameters. The contribution of left and right auditory cortex (AC) in the processing of these parameters differs. In addition, activity within the AC can vary positively or negatively with task performance depending on the type of task. This might affect the allocation of processing to the left and right AC. Here we studied with functional magnetic resonance imaging the impact of task difficulty on the degree of involvement of the left and right AC in two tasks that have previously been shown to differ in hemispheric involvement: categorization and sequential comparison of the direction of frequency modulations (FM). Task difficulty was manipulated by changing the speed of modulation and by that the frequency range covered by the FM. To study the impact of task-difficulty despite covarying the stimulus parameters, we utilized the contralateral noise procedure that allows comparing AC activation unconfounded by bottom-up driven activity. The easiest conditions confirmed the known right AC involvement during the categorization task and the left AC involvement during the comparison task. The involvement of the right AC increased with increasing task difficulty for both tasks presumably due to the common task component of categorizing FM direction. The involvement of left AC varied with task difficulty depending on the task. Thus, task difficulty has a strong impact on lateralized processing in AC. This connection must be taken into account when interpreting future results on lateralized processing in the AC.

A European Perspective on Auditory Processing Disorder-Current Knowledge and Future Research Focus.

Current notions of "hearing impairment," as reflected in clinical audiological practice, do not acknowledge the needs of individuals who have normal hearing pure tone sensitivity but who experience auditory processing difficulties in everyday life that are indexed by reduced performance in other more sophisticated audiometric tests such as speech audiometry in noise or complex non-speech sound perception. This disorder, defined as "Auditory Processing Disorder" (APD) or "Central Auditory Processing Disorder" is classified in the current tenth version of the International Classification of diseases as H93.25 and in the forthcoming beta eleventh version. APDs may have detrimental effects on the affected individual, with low esteem, anxiety, and depression, and symptoms may remain into adulthood. These disorders may interfere with learning per se and with communication, social, emotional, and academic-work aspects of life. The objective of the present paper is to define a baseline European APD consensus formulated by experienced clinicians and researchers in this specific field of human auditory science. A secondary aim is to identify issues that future research needs to address in order to further clarify the nature of APD and thus assist in optimum diagnosis and evidence-based management. This European consensus presents the main symptoms, conditions, and specific medical history elements that should lead to auditory processing evaluation. Consensus on definition of the disorder, optimum diagnostic pathway, and appropriate management are highlighted alongside a perspective on future research focus.

A Cognitive Modeling Approach to Strategy Formation in Dynamic Decision Making.

Decision-making is a high-level cognitive process based on cognitive processes like perception, attention, and memory. Real-life situations require series of decisions to be made, with each decision depending on previous feedback from a potentially changing environment. To gain a better understanding of the underlying processes of dynamic decision-making, we applied the method of cognitive modeling on a complex rule-based category learning task. Here, participants first needed to identify the conjunction of two rules that defined a target category and later adapt to a reversal of feedback contingencies. We developed an ACT-R model for the core aspects of this dynamic decision-making task. An important aim of our model was that it provides a general account of how such tasks are solved and, with minor changes, is applicable to other stimulus materials. The model was implemented as a mixture of an exemplar-based and a rule-based approach which incorporates perceptual-motor and metacognitive aspects as well. The model solves the categorization task by first trying out one-feature strategies and then, as a result of repeated negative feedback, switching to two-feature strategies. Overall, this model solves the task in a similar way as participants do, including generally successful initial learning as well as reversal learning after the change of feedback contingencies. Moreover, the fact that not all participants were successful in the two learning phases is also reflected in the modeling data. However, we found a larger variance and a lower overall performance of the modeling data as compared to the human data which may relate to perceptual preferences or additional knowledge and rules applied by the participants. In a next step, these aspects could be implemented in the model for a better overall fit. In view of the large interindividual differences in decision performance between participants, additional information about the underlying cognitive processes from behavioral, psychobiological and neurophysiological data may help to optimize future applications of this model such that it can be transferred to other domains of comparable dynamic decision tasks.

Effect of sequential comparison on active processing of sound duration.

Previous studies on active duration processing on sounds showed opposing results regarding the predominant involvement of the left or right hemisphere. Duration of an acoustic event is normally judged relative to other sounds. This requires sequential comparison as auditory events unfold over time. We hypothesized that increasing the demand on sequential comparison in a task increases the involvement of the left auditory cortex. With the current fMRI study, we investigated the effect of sequential comparison in active duration discrimination by comparing a categorical with a comparative task. During the categorical task, the participant had to categorize the tones according to their duration (short vs long). During the comparative task, they had to decide for each tone whether its length matched the tone presented before. We used the contralateral noise procedure to reveal the degree of participation of the left and right auditory cortex during these tasks. We found that both tasks more strongly involve the left than the right auditory cortex. Furthermore, the left auditory cortex was more strongly involved during comparison than during categorization. Together with previous studies, this suggests that additional demand for sequential comparison during processing of different basic acoustic parameters leads to an increased recruitment of the left auditory cortex. In addition, the comparison task more strongly involved several brain areas outside the auditory cortex, which may also be related to the demand for additional cognitive resources as compared to the more efficient categorization of sounds. Hum Brain Mapp 38:4459-4469, 2017. © 2017 Wiley Periodicals, Inc.

Reversal Learning in Humans and Gerbils: Dynamic Control Network Facilitates Learning.

Biologically plausible modeling of behavioral reinforcement learning tasks has seen great improvements over the past decades. Less work has been dedicated to tasks involving contingency reversals, i.e., tasks in which the original behavioral goal is reversed one or multiple times. The ability to adjust to such reversals is a key element of behavioral flexibility. Here, we investigate the neural mechanisms underlying contingency-reversal tasks. We first conduct experiments with humans and gerbils to demonstrate memory effects, including multiple reversals in which subjects (humans and animals) show a faster learning rate when a previously learned contingency re-appears. Motivated by recurrent mechanisms of learning and memory for object categories, we propose a network architecture which involves reinforcement learning to steer an orienting system that monitors the success in reward acquisition. We suggest that a model sensory system provides feature representations which are further processed by category-related subnetworks which constitute a neural analog of expert networks. Categories are selected dynamically in a competitive field and predict the expected reward. Learning occurs in sequentialized phases to selectively focus the weight adaptation to synapses in the hierarchical network and modulate their weight changes by a global modulator signal. The orienting subsystem itself learns to bias the competition in the presence of continuous monotonic reward accumulation. In case of sudden changes in the discrepancy of predicted and acquired reward the activated motor category can be switched. We suggest that this subsystem is composed of a hierarchically organized network of dis-inhibitory mechanisms, dubbed a dynamic control network (DCN), which resembles components of the basal ganglia. The DCN selectively activates an expert network, corresponding to the current behavioral strategy. The trace of the accumulated reward is monitored such that large sudden deviations from the monotonicity of its evolution trigger a reset after which another expert subnetwork can be activated-if it has already been established before-or new categories can be recruited and associated with novel behavioral patterns.

Probing neural mechanisms underlying auditory stream segregation in humans by transcranial direct current stimulation (tDCS).

One hypothesis concerning the neural underpinnings of auditory streaming states that frequency tuning of tonotopically organized neurons in primary auditory fields in combination with physiological forward suppression is necessary for the separation of representations of high-frequency A and low-frequency B tones. The extent of spatial overlap between the tonotopic activations of A and B tones is thought to underlie the perceptual organization of streaming sequences into one coherent or two separate streams. The present study attempts to interfere with these mechanisms by transcranial direct current stimulation (tDCS) and to probe behavioral outcomes reflecting the perception of ABAB streaming sequences. We hypothesized that tDCS by modulating cortical excitability causes a change in the separateness of the representations of A and B tones, which leads to a change in the proportions of one-stream and two-stream percepts. To test this, 22 subjects were presented with ambiguous ABAB sequences of three different frequency separations (∆F) and had to decide on their current percept after receiving sham, anodal, or cathodal tDCS over the left auditory cortex. We could confirm our hypothesis at the most ambiguous ∆F condition of 6 semitones. For anodal compared with sham and cathodal stimulation, we found a significant decrease in the proportion of two-stream perception and an increase in the proportion of one-stream perception. The results demonstrate the feasibility of using tDCS to probe mechanisms underlying auditory streaming through the use of various behavioral measures. Moreover, this approach allows one to probe the functions of auditory regions and their interactions with other processing stages.

Auditory intensity processing: Effect of MRI background noise.

Studies on active auditory intensity discrimination in humans showed equivocal results regarding the lateralization of processing. Whereas experiments with a moderate background found evidence for right lateralized processing of intensity, functional magnetic resonance imaging (fMRI) studies with background scanner noise suggest more left lateralized processing. With the present fMRI study, we compared the task dependent lateralization of intensity processing between a conventional continuous echo planar imaging (EPI) sequence with a loud background scanner noise and a fast low-angle shot (FLASH) sequence with a soft background scanner noise. To determine the lateralization of the processing, we employed the contralateral noise procedure. Linearly frequency modulated (FM) tones were presented monaurally with and without contralateral noise. During both the EPI and the FLASH measurement, the left auditory cortex was more strongly involved than the right auditory cortex while participants categorized the intensity of FM tones. This was shown by a strong effect of the additional contralateral noise on the activity in the left auditory cortex. This means a massive reduction in background scanner noise still leads to a significant left lateralized effect. This suggests that the reversed lateralization in fMRI studies with loud background noise in contrast to studies with softer background cannot be fully explained by the MRI background noise.

Delays in Human-Computer Interaction and Their Effects on Brain Activity.

The temporal contingency of feedback is an essential requirement of successful human-computer interactions. The timing of feedback not only affects the behavior of a user but is also accompanied by changes in psychophysiology and neural activity. In three fMRI experiments we systematically studied the impact of delayed feedback on brain activity while subjects performed an auditory categorization task. In the first fMRI experiment, we analyzed the effects of rare and thus unexpected delays of different delay duration on brain activity. In the second experiment, we investigated if users can adapt to frequent delays. Therefore, delays were presented as often as immediate feedback. In a third experiment, the influence of interaction outage was analyzed by measuring the effect of infrequent omissions of feedback on brain activity. The results show that unexpected delays in feedback presentation compared to immediate feedback stronger activate inter alia bilateral the anterior insular cortex, the posterior medial frontal cortex, the left inferior parietal lobule and the right inferior frontal junction. The strength of this activation increases with the duration of the delay. Thus, delays interrupt the course of an interaction and trigger an orienting response that in turn activates brain regions of action control. If delays occur frequently, users can adapt, delays become expectable, and the brain activity in the observed network diminishes over the course of the interaction. However, introducing rare omissions of expected feedback reduces the system's trustworthiness which leads to an increase in brain activity not only in response to such omissions but also following frequently occurring and thus expected delays.

Neuronal correlates of voluntary facial movements.

Whereas the somatotopy of finger movements has been extensively studied with neuroimaging, the neural foundations of facial movements remain elusive. Therefore, we systematically studied the neuronal correlates of voluntary facial movements using the Facial Action Coding System (FACS, Ekman et al., 2002). The facial movements performed in the MRI scanner were defined as Action Units (AUs) and were controlled by a certified FACS coder. The main goal of the study was to investigate the detailed somatotopy of the facial primary motor area (facial M1). Eighteen participants were asked to produce the following four facial movements in the fMRI scanner: AU1+2 (brow raiser), AU4 (brow lowerer), AU12 (lip corner puller) and AU24 (lip presser), each in alternation with a resting phase. Our facial movement task induced generally high activation in brain motor areas (e.g., M1, premotor cortex, supplementary motor area, putamen), as well as in the thalamus, insula, and visual cortex. BOLD activations revealed overlapping representations for the four facial movements. However, within the activated facial M1 areas, we could find distinct peak activities in the left and right hemisphere supporting a rough somatotopic upper to lower face organization within the right facial M1 area, and a somatotopic organization within the right M1 upper face part. In both hemispheres, the order was an inverse somatotopy within the lower face representations. In contrast to the right hemisphere, in the left hemisphere the representation of AU4 was more lateral and anterior compared to the rest of the facial movements. Our findings support the notion of a partial somatotopic order within the M1 face area confirming the "like attracts like" principle (Donoghue et al., 1992). AUs which are often used together or are similar are located close to each other in the motor cortex.

Superior memorizers employ different neural networks for encoding and recall.

Superior memorizers often employ the method of loci (MoL) to memorize large amounts of information. The MoL, known since ancient times, relies on a complex process where information to be memorized is bound to landmarks along mental routes in a previously memorized environment. However, functional magnetic resonance imaging data on groups of trained superior memorizer are rare. Based on the memorizing strategy reported by superior memorizers, we developed a scheme of the processes successively employed during memorizing and recalling digits and relate these to brain activation that is specific for the encoding and recall period. In the examined superior memorizers several regions, suggested to be involved in mental navigation and digit-to-word processing, were specifically activated during encoding: bilateral early visual cortex, retrosplenial cortex, left parahippocampus, left visual cortex, and left superior parietal cortex. Although the scheme suggests that some steps during encoding and recall seem to be analog, none of the encoding areas were specifically activated during the recall. Instead, we found strong activation in left anterior superior temporal gyrus, which we relate to recalling the sequential order of the digits, and right motor cortex that may be related to reciting the digits.

Decision making and ambiguity in auditory stream segregation.

Researchers of auditory stream segregation have largely taken a bottom-up view on the link between physical stimulus parameters and the perceptual organization of sequences of ABAB sounds. However, in the majority of studies, researchers have relied on the reported decisions of the subjects regarding which of the predefined percepts (e.g., one stream or two streams) predominated when subjects listened to more or less ambiguous streaming sequences. When searching for neural mechanisms of stream segregation, it should be kept in mind that such decision processes may contribute to brain activation, as also suggested by recent human imaging data. The present study proposes that the uncertainty of a subject in making a decision about the perceptual organization of ambiguous streaming sequences may be reflected in the time required to make an initial decision. To this end, subjects had to decide on their current percept while listening to ABAB auditory streaming sequences. Each sequence had a duration of 30 s and was composed of A and B harmonic tone complexes differing in fundamental frequency (ΔF). Sequences with seven different ΔF were tested. We found that the initial decision time varied non-monotonically with ΔF and that it was significantly correlated with the degree of perceptual ambiguity defined from the proportions of time the subjects reported a one-stream or a two-stream percept subsequent to the first decision. This strong relation of the proposed measures of decision uncertainty and perceptual ambiguity should be taken into account when searching for neural correlates of auditory stream segregation.

Auditory intensity processing: Categorization versus comparison.

Intensity is an important parameter for the perception of complex auditory stimuli like speech. The results of previous studies on the processing of intensity are diverse since left-lateralized, right-lateralized and non-lateralized processing was suggested. A clear dependence of the lateralization on the kind of stimuli and/or task is not apparent. With the present functional magnetic resonance imaging (fMRI) study, we directly investigated the differences between a categorical and comparative task. To determine hemispheric involvement we used a method with contralateral noise presentation. Harmonic complexes were presented monaurally without and with contralateral noise. Both categorization and comparison of harmonic complexes according to their intensity more strongly involved the left than the right auditory cortex shown by a stronger effect of the additional noise on the activity in the left auditory cortex. Together with previous results, this suggests that left-lateralized processing of intensity in the auditory cortex can be observed independent of task and stimuli. The comparison task more strongly engaged the left auditory cortex than the categorization task probably due the additional need for sequential comparison and the right auditory cortex probably due to capacity reasons. Comparison also more strongly engaged areas associated with attentional processes and areas responsible for motor response selection. We suggest this to be caused by a more difficult response selection and by the need for continuous update of information in reference memory during the comparison task.