Pre-stimulus beta power varies as a function of auditory-motor synchronization and temporal predictability.

Introduction: Auditory-motor interactions can support the preparation for expected sensory input. We investigated the periodic modulation of beta activity in the electroencephalogram to assess the role of active auditory-motor synchronization. Pre-stimulus beta activity (13-30 Hz) has been interpreted as a neural signature of the preparation for expected sensory input. Methods: In the current study, participants silently counted frequency deviants in sequences of pure tones either during a physically inactive control condition or while pedaling on a cycling ergometer. Tones were presented either rhythmically (at 1 Hz) or arrhythmically with variable intervals. In addition to the pedaling conditions with rhythmic (auditory-motor synchronization, AMS) or arrhythmic stimulation, a self-generated stimulus condition was used in which tones were presented in sync with the participants' spontaneous pedaling. This condition served to explore whether sensory predictions are driven primarily by the auditory or by the motor system. Results: Pre-stimulus beta power increased for rhythmic compared to arrhythmic stimulus presentation in both sitting and pedaling conditions but was strongest in the AMS condition. Furthermore, beta power in the AMS condition correlated with motor performance, i.e., the better participants synchronized with the rhythmic stimulus sequence, the higher was pre-stimulus beta power. Additionally, beta power was increased for the self-generated stimulus condition compared with arrhythmic pedaling, but there was no difference between the self-generated and the AMS condition. Discussion: The current data pattern indicates that pre-stimulus beta power is not limited to neuronal entrainment (i.e., periodic stimulus presentation) but represents a more general correlate of temporal anticipation. Its association with the precision of AMS supports the role of active behavior for auditory predictions.

Cognitive effects of rhythmic auditory stimulation in Parkinson's disease: A P300 study.

Rhythmic auditory stimulation (RAS) may compensate dysfunctions of the basal ganglia (BG), involved with intrinsic evaluation of temporal intervals and action initiation or continuation. In the cognitive domain, RAS containing periodically presented tones facilitates young healthy participants' attention allocation to anticipated time points, indicated by better performance and larger P300 amplitudes to periodic compared to random stimuli. Additionally, active auditory-motor synchronization (AMS) leads to a more precise temporal encoding of stimuli via embodied timing encoding than stimulus presentation adapted to the participants' actual movements. Here we investigated the effect of RAS and AMS in Parkinson's disease (PD). 23 PD patients and 23 healthy age-matched controls underwent an auditory oddball task. We manipulated the timing (periodic/random/adaptive) and setting (pedaling/sitting still) of stimulation. While patients elicited a general timing effect, i.e., larger P300 amplitudes for periodic versus random tones for both, sitting and pedaling conditions, controls showed a timing effect only for the sitting but not for the pedaling condition. However, a correlation between P300 amplitudes and motor variability in the periodic pedaling condition was obtained in control participants only. We conclude that RAS facilitates attentional processing of temporally predictable external events in PD patients as well as healthy controls, but embodied timing encoding via body movement does not affect stimulus processing due to BG impairment in patients. Moreover, even with intact embodied timing encoding, such as healthy elderly, the effect of AMS depends on the degree of movement synchronization performance, which is very low in the current study.

Pre-encoding gamma-band activity during auditory working memory.

Previous magnetoencephalography (MEG) studies have revealed gamma-band activity at sensors over parietal and fronto-temporal cortex during the delay phase of auditory spatial and non-spatial match-to-sample tasks, respectively. While this activity was interpreted as reflecting the memory maintenance of sound features, we noted that task-related activation differences might have been present already prior to the onset of the sample stimulus. The present study focused on the interval between a visual cue indicating which sound feature was to be memorized (lateralization or pitch) and sample sound presentation to test for task-related activation differences preceding stimulus encoding. MEG spectral activity was analyzed with cluster randomization tests (N = 15). Whereas there were no differences in frequencies below 40 Hz, gamma-band spectral amplitude (about 50-65 and 90-100 Hz) was higher for the lateralization than the pitch task. This activity was localized at right posterior and central sensors and present for several hundred ms after task cue offset. Activity at 50-65 Hz was also increased throughout the delay phase for the lateralization compared with the pitch task. Apparently cortical networks related to auditory spatial processing were activated after participants had been informed about the task.

Actively but not passively synchronized motor activity amplifies predictive timing.

Previous studies have shown that the effect of temporal predictability of presented stimuli on attention allocation is enhanced by auditory-motor synchronization (AMS). The present P300 event-related potential study (N=20) investigated whether this enhancement depends on the process of actively synchronizing one's motor output with the acoustic input or whether a passive state of auditory-motor synchrony elicits the same effect. Participants silently counted frequency deviants in sequences of pure tones either during a physically inactive control condition or while pedaling on a cycling ergometer. Tones were presented either at fixed or variable intervals. In addition to the pedaling conditions with fixed or variable stimulation, there was a third condition in which stimuli were adaptively presented in sync with the participants' spontaneous pedaling. We replicated the P300 enhancement for fixed versus variable stimulation and the amplification of this effect by AMS. Synchronization performance correlated positively with P300 amplitude in the fixed stimulation condition. Most interestingly, P300 amplitude was significantly reduced for the passive synchronization condition by adaptive stimulus presentation as compared to the fixed stimulation condition. For the first time we thus provide evidence that it is not the passive state of (even perfect) auditory-motor synchrony that facilitates attention allocation during AMS but rather the active process of synchronizing one's movements with external stimuli.

Treadmill walking during vocabulary encoding improves verbal long-term memory.

Moderate physical activity improves various cognitive functions, particularly when it is applied simultaneously to the cognitive task. In two psychoneuroendocrinological within-subject experiments, we investigated whether very low-intensity motor activity, i.e. walking, during foreign-language vocabulary encoding improves subsequent recall compared to encoding during physical rest. Furthermore, we examined the kinetics of brain-derived neurotrophic factor (BDNF) in serum and salivary cortisol. Previous research has associated both substances with memory performance.In both experiments, subjects performed better when they were motorically active during encoding compared to being sedentary. BDNF in serum was unrelated to memory performance. In contrast we found a positive correlation between salivary cortisol concentration and the number of correctly recalled items. In summary, even very light physical activity during encoding is beneficial for subsequent recall.

Attentional modulation of the inner ear: a combined otoacoustic emission and EEG study.

Attending to a single stimulus in a complex multisensory environment requires the ability to select relevant information while ignoring distracting input. The underlying mechanism and involved neuronal levels of this attentional gain control are still a matter of debate. Here, we investigated the influence of intermodal attention on different levels of auditory processing in humans. It is known that the activity of the cochlear amplifier can be modulated by efferent neurons of the medial olivocochlear complex. We used distortion product otoacoustic emission (DPOAE) measurements to monitor cochlear activity during an intermodal cueing paradigm. Simultaneously, central auditory processing was assessed by electroencephalography (EEG) with a steady-state paradigm targeting early cortical responses and analysis of alpha oscillations reflecting higher cognitive control of attentional modulation. We found effects of selective attention at all measured levels of the auditory processing: DPOAE levels differed significantly between periods of visual and auditory attention, showing a reduction during visual attention, but no change during auditory attention. Primary auditory cortex activity, as measured by the auditory steady-state response (ASSR), differed between conditions, with higher ASSRs during auditory than visual attention. Furthermore, the analysis of cortical oscillatory activity revealed increased alpha power over occipitoparietal and frontal regions during auditory compared with visual attention, putatively reflecting suppression of visual processing. In conclusion, this study showed both enhanced processing of attended acoustic stimuli in early sensory cortex and reduced processing of distracting input, both at higher cortical levels and at the most peripheral level of the hearing system, the cochlea.

Auditory-motor synchronization facilitates attention allocation.

Temporal predictability of auditory events induces larger P300 amplitudes and shorter P300 latencies compared to stimulus presentation with variable onset asynchronies. This suggests that periodic stimuli lead to neuronal entrainment resulting in a more efficient allocation of attentional resources. Simultaneous synchronized motor activity should facilitate the precise temporal encoding of acoustic sequences. Therefore the current event-related potential study investigated whether embodied stimulus encoding enhances the reported effects of stimulus periodicity. We found that simultaneous pedaling on an ergometer compared to a physically passive situation amplified the predictability effect on the P300 component. Furthermore, the temporal variability of cycling behavior correlated positively with both P300 latency and P300 amplitude. These findings indicate that auditory-motor synchronization enhances the attentional processing of periodical auditory stimuli.

P3b reflects periodicity in linguistic sequences.

Temporal predictability is thought to affect stimulus processing by facilitating the allocation of attentional resources. Recent studies have shown that periodicity of a tonal sequence results in a decreased peak latency and a larger amplitude of the P3b compared with temporally random, i.e., aperiodic sequences. We investigated whether this applies also to sequences of linguistic stimuli (syllables), although speech is usually aperiodic. We compared aperiodic syllable sequences with two temporally regular conditions. In one condition, the interval between syllable onset was fixed, whereas in a second condition the interval between the syllables' perceptual center (p-center) was kept constant. Event-related potentials were assessed in 30 adults who were instructed to detect irregularities in the stimulus sequences. We found larger P3b amplitudes for both temporally predictable conditions as compared to the aperiodic condition and a shorter P3b latency in the p-center condition than in both other conditions. These findings demonstrate that even in acoustically more complex sequences such as syllable streams, temporal predictability facilitates the processing of deviant stimuli. Furthermore, we provide first electrophysiological evidence for the relevance of the p-center concept in linguistic stimulus processing.

The auditory cortex of the bat Molossus molossus: disproportionate search call frequency representation.

The extent of the auditory cortex in the bat Molossus molossus was electrophysiologically investigated. Best frequencies and minimum thresholds of neural tuning curves were analyzed to define the topography of the auditory cortex. The auditory cortex encompasses an average cortical surface area of 5mm(2). Characteristic frequencies are tonotopically organized with low frequencies being represented caudally and high frequencies rostrally. However, a large interindividual variability in the tonotopic organization was found. In most animals, the caudal 50% was tonotopically organized. More anterior, a variable area was found. A distinct field with reversed topography was not consistently found. Within the demarcated auditory cortex, frequencies of 30-40 kHz, which correspond to the frequency range of search calls emitted during hunting, are overrepresented, occupying 49% of the auditory cortex surface. High minimum thresholds >50 dB SPL were found in a narrow dorsal narrow area. Neurons with multipeaked tuning curves (20%) preferentially were located in the dorsal part of the auditory cortex. In accordance with studies in other bat species, the auditory cortex of M. molossus is highly sensitive to the dominant frequencies of biosonar search calls.

Contralateral acoustic stimulation modulates low-frequency biasing of DPOAE: efferent influence on cochlear amplifier operating state?

The mammalian efferent medial olivocochlear system modulates active amplification of low-level sounds in the cochlea. Changes of the cochlear amplifier can be monitored by distortion product otoacoustic emissions (DPOAEs). The quadratic distortion product f2-f1 is known to be sensitive to changes in the operating point of the amplifier transfer function. We investigated the effect of contralateral acoustic stimulation (CAS), known to elicit efferent activity, on DPOAEs in the gerbil. During CAS, a significant increase of the f2-f1 level occurred already at low contralateral noise levels (20 dB SPL), whereas 2f1-f2 was much less affected. The effect strength depended on the CAS level and as shown in experiments with pure tones on the frequency of the contralateral stimulus. In a second approach, we biased the position of the cochlear partition and thus the cochlear amplifier operating point periodically by a ipsilateral low-frequency tone, which resulted in a phase-related amplitude modulation of f2-f1. This modulation pattern was changed considerably during contralateral noise stimulation, in dependence on the noise level. The experimental results were in good agreement with a simple model of distortion product generation and suggest that the olivocochlear efferents might change the operating state of cochlear amplification.

Sensitive response to low-frequency cochlear distortion products in the auditory midbrain.

During auditory stimulation with several frequency components, distortion products (DPs) are generated as byproduct of nonlinear cochlear amplification. After generated, DP energy is reemitted into the ear channel where it can be measured as DP otoacoustic emission (DPOAE), and it also induces an excitatory response at cochlear places related to the DP frequencies. We measured responses of 91 inferior colliculus (IC) neurons in the gerbil during two-tone stimulation with frequencies well above the unit's receptive field but adequate to generate a distinct distortion product (f2-f1 or 2f1-f2) at the unit's characteristic frequency (CF). Neuronal responses to DPs could be accounted for by the simultaneously measured DPOAEs for DP frequencies >1.3 kHz. For DP frequencies <1.3 kHz (n = 25), there was a discrepancy between intracochlear DP magnitude and DPOAE level, and most neurons responded as if the intracochlear DP level was significantly higher than the DPOAE level in the ear channel. In 12% of those low-frequency neurons, responses to the DPs could be elicited even if the stimulus tone levels were below the threshold level of the neuron at CF. High intracochlear f2-f1 and 2f1-f2 DP-levels were verified by cancellation of the neuronal DP response with a third phase-adjusted tone stimulus at the DP frequency. A frequency-specific reduction of middle ear gain at low frequencies is possibly involved in the reduction of DPOAE level. The results indicate that pitch-related properties of complex stimuli may be produced partially by high intracochlear f2-f1 distortion levels.