Connecting the dots: Sensory cueing enhances functional connectivity between pre-motor and supplementary motor areas in Parkinson's disease.

People with Parkinson's disease often exhibit improvements in motor tasks when exposed to external sensory cues. While the effects of different types of sensory cues on motor functions in Parkinson's disease have been widely studied, the underlying neural mechanism of these effects and the potential of sensory cues to alter the motor cortical activity patterns and functional connectivity of cortical motor areas are still unclear. This study aims to compare changes in oxygenated haemoglobin, deoxygenated haemoglobin and correlations among different cortical regions of interest during wrist movement under different external stimulus conditions between people with Parkinson's disease and controls. Ten Parkinson's disease patients and 10 age- and sex-matched neurologically healthy individuals participated, performing repetitive wrist flexion and extension tasks under auditory and visual cues. Changes in oxygenated and deoxygenated haemoglobin in motor areas were measured using functional near-infrared spectroscopy, along with electromyograms from wrist muscles and wrist movement kinematics. The functional near-infrared spectroscopy data revealed significantly higher neural activity changes in the Parkinson's disease group's pre-motor area compared to controls (p = 0.006), and functional connectivity between the supplementary motor area and pre-motor area was also significantly higher in the Parkinson's disease group when external sensory cues were present (p = 0.016). These results indicate that external sensory cues' beneficial effects on motor tasks are linked to changes in the functional connectivity between motor areas responsible for planning and preparation of movements.

Spaced conditioned stimulus presentation facilitates the extinction of strong fear memory in mice.

Inducing fear memory extinction by re-presenting a conditioned stimulus (CS) is the foundation of exposure therapy for post-traumatic stress disorder (PTSD). Investigating differences in the ability of different CS presentation patterns to induce extinction learning is crucial for improving this type of therapy. Using a trace fear conditioning paradigm in mice, we demonstrate that spaced presentation of the CS facilitated the extinction of a strong fear memory to a greater extent than continuous CS presentation. These results lay the groundwork for developing more effective exposure therapy techniques for PTSD.

Older adults can rely on an auditory cue to generate anticipatory postural adjustments prior to an external perturbation.

To minimize the potential postural disturbance induced by predictable external perturbations, humans generate anticipatory postural adjustments (APAs) using visual information about a perturbation. However, it is unknown whether older adults can generate APAs relying on auditory information. Ten older adults received external perturbations (a) with visual information but no auditory information available, (b) without neither visual nor auditory information, (c) with both visual and auditory information available, and (d) participated in training with only auditory information available. In addition, they were tested again after 1 week of washout period. Electromyography activities of eight leg and trunk muscles and ground reaction forces were recorded and analyzed during the anticipatory and compensatory phases. Outcome measures included the latencies and integrals of muscle activities, and center-of-pressure displacements. After a short period of training, participants were able to rely on the auditory cue only to generate APAs close to that when the visual information was available. In addition, after 1 week of washout period, they were able to partially retain the skill to rely on auditory cues to generate APAs. The outcome provides a foundation for future studies focusing on utilizing auditory cues to optimize postural control in individuals who have balance or vision deficit.

Effects of rhythmic auditory stimulation on upper-limb movement speed in patients with schizophrenia spectrum disorders.

Movement slowness, linked to dysfunctional basal ganglia and cerebellum, is prevalent but lacks effective therapy in patients with schizophrenia spectrum disorders. This study was to examine immediate effects of rhythmic auditory stimulation (RAS) on upper-limb movement speed in patients. Thirty patients and 30 psychiatrically healthy people executed the right-hand task and the both-hand task of the Purdue Pegboard Test when listening to RAS with two tempi: normal (equal to the fastest movement tempo for each participant without RAS) and fast (120% of the normal tempo). The testing order of the RAS tempi for each participant was randomized. Patients had lower scores of right-hand and both-hand tasks than did psychiatrically healthy people. Scores of right-hand and both-hand tasks were higher in the fast-RAS condition than the normal-RAS condition in participants. This is the first study to explore the possibility of applying RAS to movement therapy for patients with schizophrenia spectrum disorders. The results demonstrated that faster RAS was effective in inducing faster upper-limb movements in patients and psychiatrically healthy people, suggesting that manipulating RAS may be a feasible therapeutic strategy utilized to regulate movement speed. The RAS may involve alternative neural pathways to modulate movement speed and thus to compensate for impaired function of basal ganglia and cerebellum in patients.

Determining social power: Do Chinese preschoolers integrate verbal and nonverbal cues?

Children aged about 5 years can use verbal and nonverbal cues to determine social power. However, it is not clear what kinds of nonverbal cues preschoolers can use and whether they can integrate each nonverbal cue with verbal content when determining social power. Therefore, this research examined the ability of Chinese preschoolers to use and integrate visual, auditory, and verbal cues when determining social power as well as how this ability develops with age. In Study 1 (N = 478), 4- to 6-year-old Chinese children were recruited to judge the social power in visual, auditory, content, visual-auditory, visual-content, auditory-content, visual-auditory-content, and neutral cues. The results showed that 4-year-olds could not use any cues, 5-year-olds could not use content cues, and 6-year-olds could use all the cues. Children's performance improved with increasing age and the number of channels. Study 2 (N = 240) investigated whether speech tempo influences children's performance when using content cues. The results showed that 5- and 6-year-olds, but not 4-year-olds, could use content cues to determine social power under normal and slow tempos. Study 3 (N = 80) was conducted to clarify which visual cues 5- and 6-year-olds can use and the age difference. The results revealed that 5- and 6-year-olds could use head, eye, posture, and mixed cues to determine social power, whereas their performance was steady for both 5- and 6-year-olds. In sum, the results show that Chinese children aged about 5 years are already able to use and integrate visual, auditory, and content cues when determining social power. Moreover, the ability to determine social power is gradually improved with age during the preschool period.

The role of an auditory cue in generating anticipatory postural adjustments in response to an external perturbation.

Anticipatory postural adjustments (APAs) are usually generated to minimize the potential postural disturbance induced by predictable external perturbations. Visual information about a perturbation is important for the generation of APAs, but whether people can rely on auditory information to generate APAs is unknown. The aim of this study was to investigate the role of an auditory cue in generating APAs when visual information is not available. Fifteen young adults participated in the study. They received external perturbations a) with visual information but no auditory information available, b) without neither visual nor auditory information, c) with both visual and auditory information available, and d) with only auditory information available. Electromyography (EMG) activities of eight leg and trunk muscles and displacements of the center-of-pressure (COP) were recorded and analyzed during the anticipatory and compensatory (CPAs) phases. Outcome measures included the latencies and integrals of muscle activities, COP displacements, and indices of co-contraction and reciprocal activation of muscles. The results showed that after a short training, participants were able to rely only on the auditory cue to generate APAs comparable to that when the visual information was available. In addition, a training effect was found such that the participants demonstrated stronger APAs and less demands for CPAs through the training trials. The outcome provides a foundation for future studies focusing on the utilization of auditory cues for postural control in older adults and individuals who have vision deficit.

Deficits in tapping accuracy and variability in tremor patients.

BACKGROUND: The basal ganglia and cerebellum are brain structures involved in movement initiation, execution and termination. They are thought to be involved in the tremor generation and movement deficits in Parkinson's disease (PD) and essential tremor (ET). Especially in PD, maintaining cyclic movement, such as walking or tapping can be significantly disturbed. Providing external cues improves timing of these movements in PD but its effect on ET has not yet been studied in depth. The aim of this study is to evaluate the usefulness of a bimanual tapping task as a tool during clinical decision making. METHOD: Hand movements and tremor was recorded using accelerometers and EMG (m. extensor carpi ulnaris) from PD and ET patients and healthy controls during a bimanual tapping task as a way to distinguish PD from ET. All subjects performed the tapping task at two different frequencies, 2 Hz and 4 Hz, with and without the presence of auditory cues. RESULTS: No significant intra-group differences were found in the patient groups. Acceleration data revealed significantly less accurate tapping and more variable tapping in PD than in ET and healthy controls. ET subjects tapped less accurate and with a greater variability than healthy controls during the 4 Hz tapping task. Most interestingly the tapping accuracy improved in PD patients when kinetic tremor was recorded with EMG during the task. CONCLUSION: Providing ET and PD patients with an external cue results in different tapping performances between patient groups and healthy controls. Furthermore, the findings suggest that kinetic tremor in PD enables patients to perform the task with a greater accuracy. So far this has not been shown in other studies.

Domestication reduces caterpillar response to auditory predator cues.

Domestication can lead to significant changes in the growth and behavior of organisms. While the threat of predation is a strong selective force in the wild, the relaxation or removal of this threat in captive-rearing environments selects for reduced sensitivity to biotic stressors. Previous work has documented such changes in other taxa, but no work has been done on domestication-related losses of predation risk sensitivity in insects. We exposed both wild and domesticated (>50 generations in captivity) Lymantria dispar dispar (Lepidoptera: Erebidae) larvae to recordings of predators (wasp buzzing), nonpredators (mosquito buzzing), or no sound to compare the effects of predation risk on the two stocks. Wasp buzzing, but not mosquito buzzing, decreased survival of wild caterpillars relative to the control; domesticated caterpillars showed no such response. Domesticated L. dispar larvae appear to have reduced sensitivity to predation risk cues, suggesting that captive-reared insects may not always be analogs to their wild counterparts for risk-related behavioral studies.

Auditory predator cues decrease herbivore survival and plant damage.

The high fitness cost of predation selects prey capable of detecting risk cues and responding in ways that reduce their vulnerability. While the impacts of auditory predator cues have been extensively researched in vertebrate prey, much less is known about invertebrate species' responses and their potential to affect the wider food web. We exposed larvae of Spodoptera exigua, a slow-moving and vulnerable herbivore hunted by aerial predators, to recordings of wasp buzzing (risk cue), mosquito buzzing (no-risk cue), or a no-sound control in both laboratory and field settings. In the laboratory, wasp buzzing (but not mosquito buzzing) reduced survival relative to the control; there was, however, no effect on time to or weight at pupation in survivors. In the field, wasp buzzing reduced caterpillar herbivory and increased plant biomass relative to the control treatment. In contrast, mosquito buzzing reduced herbivory less than wasp buzzing and had no effect on plant biomass. The fact that wasp cues evoked strong responses in both experiments, while mosquito buzzing generally did not, indicates that caterpillars were responding to predation risk rather than sound per se. Such auditory cues may have an important but largely unappreciated impacts on terrestrial invertebrate herbivores and their host plants.

Facilitating or disturbing? An investigation about the effects of auditory frequencies on prefrontal cortex activation and postural sway.

Auditory stimulation activates brain areas associated with higher cognitive processes, like the prefrontal cortex (PFC), and plays a role in postural control regulation. However, the effects of specific frequency stimuli on upright posture maintenance and PFC activation patterns remain unknown. Therefore, the study aims at filling this gap. Twenty healthy adults performed static double- and single-leg stance tasks of 60s each under four auditory conditions: 500, 1000, 1500, and 2000 Hz, binaurally delivered through headphones, and in quiet condition. Functional near-infrared spectroscopy was used to measure PFC activation through changes in oxygenated hemoglobin concentration, while an inertial sensor (sealed at the L5 vertebra level) quantified postural sway parameters. Perceived discomfort and pleasantness were rated through a 0-100 visual analogue scale (VAS). Results showed that in both motor tasks, different PFC activation patterns were displayed at the different auditory frequencies and the postural performance worsened with auditory stimuli, compared to quiet conditions. VAS results showed that higher frequencies were considered more discomfortable than lower ones. Present data prove that specific sound frequencies play a significant role in cognitive resources recruitment and in the regulation of postural control. Furthermore, it supports the importance of exploring the relationship among tones, cortical activity, and posture, also considering possible applications with neurological populations and people with hearing dysfunctions.

The effects of treadmill training with visual feedback and rhythmic auditory cue on gait and balance in chronic stroke patients: A randomized controlled trial.

BACKGROUND: Many stroke patients show reduced walking abilities, characterized by asymmetric walking patterns. For such patients, restoration of walking symmetry is important. OBJECTIVE: This study investigates the effect of treadmill training with visual feedback and rhythmic auditory cue (VF+RAC) for walking symmetry on spatiotemporal gait parameters and balance abilities. METHODS: Thirty-two patients with chronic stroke participated in this study. Participants were randomized to either the VF+RAC (n = 16) or the Control (n = 16) group. The VF+RAC group received treadmill training with VF and RAC, and the Control group underwent treadmill training without any visual and auditory stimulation. VF+RAC and Control groups were trained three times per week for eight weeks. After eight weeks of training, the spatiotemporal gait parameters, Timed up and go test, and Berg balance scale were measured. RESULTS: The VF+RAC group significantly improved balance and spatiotemporal parameters except for non-paretic single limb support compared to the Control group. CONCLUSIONS: This study demonstrated that treadmill training with VF+RAC significantly improved spatiotemporal gait symmetry, including other gait parameters, and enhanced balance abilities in stroke patients. Therefore, treadmill training with VF+RAC could be a beneficial intervention in clinical settings for stroke patients who need improvement in their gait and balance abilities.

Frequency Nesting Interactions in the Subthalamic Nucleus Correlate With the Step Phases for Parkinson's Disease.

Gait disturbance in Parkinson's disease (PD) can be ameliorated by sound stimulation. Given that excessive ß synchronization in basal ganglia is linked to motor impairment in PD, whether the frequency nesting interactions are associated with the gait problem is far from clear. To this end, the masking phase-amplitude coupling (PAC) method was proposed to overcome the trade-off between intrinsic nonlinearity/non-stationarity and demand for predetermined frequencies, normally extracted by the filter. In this study, we analyzed LFPs recorded from 13 patients (one female) with PD during stepping with bilateral deep brain electrodes implanted in the subthalamic nucleus (STN). We found that not only high-frequency oscillation (100-300 Hz) was modulated by ß (13-30 Hz) but also ß and γ amplitude were modulated by their low-frequency components in δ/θ/α and δ/θ/α/ß bands. These PAC values were suppressed by sound stimulation, along with an improvement in gait. We also showed that gait-related high-ß (Hß) modulation in the STN was sensitive to auditory cues, and Hß gait-phase modulation increased with a metronome. Meanwhile, phase-locking values (PLVs) across all frequencies were significantly suppressed around contralateral heel strikes, manifesting the contralateral step as a critical gait phase in gait initiation for PD. Only the PLVs around contralateral steps were sensitive to auditory cues. Our results support masking PAC as an effective method in exploring frequency nesting interactions in LFPs and reveal the linkages between sound stimulation and couplings related to gait phases in the STN. These findings raise the possibility that nesting interactions in the STN work as feasible biomarkers in alleviating gait disorders.

An inferior-superior colliculus circuit controls auditory cue-directed visual spatial attention.

Selective attention modulates neuronal activity in multiple brain regions, but the origins of attention signals remain unclear. We show that, during a visual task requiring spatial attention directed by an auditory cue, an inferior-superior colliculus circuit provides the key attention signal. In mice performing a task based on a visual stimulus in the cued hemifield while ignoring a conflicting stimulus on the uncued side, the visual cortex (V1) and superior colliculus (SC) showed strong attentional modulation, with a shorter latency in the SC. The nucleus of the brachium of the inferior colliculus (nBIC), which provides auditory inputs to the SC, was activated not only at auditory cue onset but also during the delay period before the visual stimulus. The delay activity, but not cue onset activity, was crucial for task performance and attentional modulation in the SC and V1. These results establish a new behavioral paradigm for studying visual attention in mice and identify a midbrain signal controlling auditory cue-directed spatial attention.

Tone deaf: Association of an auditory stimulus with predation risk by zebrafish Danio rerio does not generalize to another auditory stimulus.

Predator recognition by small fishes can be acquired when chemical alarm cues released from damaged skin (by a predator attack) are paired with a novel stimulus, such as the appearance or odor of a predator. Once learned, fish can extend recognition of risk by generalizing to associate risk with additional stimuli that are similar to the conditioned novel stimulus. Here, we trained zebrafish to associate a novel auditory stimulus with predation risk, and then tested to see if they generalize risk to all sound stimuli or whether the conditioned response is limited to the sound frequency of the conditioning stimulus. We found that zebrafish Danio rerio readily associated risk of predation with Tone 1 (285 Hz), as evidenced by reduction in activity, increased time spent near the substratum and increased shelter use, but fish conditioned to fear Tone 1 completely ignored presentation of a second tone of 762 Hz. These data suggest that generalization does not occur as easily for auditory cues as they do for olfactory and visual cues, perhaps due to differences in the properties of sensory biology or the cognitive mechanisms that process information in different sensory modalities.

Spatial reorientation with non-visual cues: Failure to spontaneously use auditory information.

Among the environmental stimuli that can guide navigation in space, most attention has been dedicated to visual information. The process of determining where you are and which direction you are facing (called reorientation) has been extensively examined by providing the navigator with two sources of information-typically the shape of the environment and its features-with an interest in the extent to which they are used. Similar questions with non-visual cues are lacking. Here, blindfolded sighted participants had to learn the location of a target in a real-world, circular search space. In Experiment 1, two ecologically relevant non-visual cues were provided: the slope of the floor and an array of two identical auditory landmarks. Slope successfully guided behaviour, suggesting that proprioceptive/kinesthetic access is sufficient to navigate on a slanted environment. However, despite the fact that participants could localise the auditory sources, this information was not encoded. In Experiment 2, the auditory cue was made more useful for the task because it had greater predictive value and there were no competing spatial cues. Nonetheless, again, the auditory landmark was not encoded. Finally, in Experiment 3, after being prompted, participants were able to reorient by using the auditory landmark. Overall, participants failed to spontaneously rely on the auditory cue, regardless of how informative it was.

Motor Adaptation in Parkinson's Disease During Prolonged Walking in Response to Corrective Acoustic Messages.

Wearable sensing technology is a new way to deliver corrective feedback. It is highly applicable to gait rehabilitation for persons with Parkinson's disease (PD) because feedback potentially engages spared neural function. Our study characterizes participants' motor adaptation to feedback signaling a deviation from their normal cadence during prolonged walking, providing insight into possible novel therapeutic devices for gait re-training. Twenty-eight persons with PD (15 with freezing, 13 without) and 13 age-matched healthy elderly (HE) walked for two 30-minute sessions. When their cadence varied, they heard either intelligent cueing (IntCue: bouts of ten beats indicating normal cadence) or intelligent feedback (IntFB: verbal instruction to increase or decrease cadence). We created a model that compares the effectiveness of the two conditions by quantifying the number of steps needed to return to the target cadence for every deviation. The model fits the short-term motor responses to the external step inputs (collected with wearable sensors). We found some significant difference in motor adaptation among groups and subgroups for the IntCue condition only. Both conditions were instead able to identify different types of responders among persons with PD, although showing opposite trends in their speed of adaptation. Increasing rather than decreasing the pace appeared to be more difficult for both groups. In fact, under IntFB the PD group required about seven steps to increase their cadence, whereas they only needed about three steps to decrease their cadence. However, it is important to note that this difference was not significant; perhaps future work could include more participants and/or more sessions, increasing the total number of deviations for analysis. Notably, a significant negative correlation, r = -0.57 (p-value = 0.008), was found between speed of adaptation and number of deviations during IntCue, but not during IntFB, suggesting that, for people who struggle with gait, such as those with PD, verbal instructions rather than metronome beats might be more effective at restoring normal cadence. Clinicians and biofeedback developers designing novel therapeutic devices could apply our findings to determine the optimal timing for corrective feedback, optimizing gait rehabilitation while minimizing the risk of cue-dependency.

Influence of visual feedback and rhythmic auditory cue on walking of chronic stroke patient induced by treadmill walking in real-time basis.

BACKGROUND: Many patients who have suffered from a stroke show decreased walking ability, characterized by asymmetric gait. For such patients, the recovery of symmetry in walking is important. OBJECTIVES: The purpose of this study is to investigate the effect of visual feedback with rhythmic auditory cue on treadmill gait in persons with chronic stroke. METHODS: Seventeen chronic subjects who have been diagnosed at least six months or before were recruited in G Hospital, located in Incheon. The subjects who were enrolled in this study were instructed to walk randomly on a treadmill: treadmill with visual feedback and rhythmic auditory cue (VF+RAC), treadmill with visual feedback (VF), general treadmill (Control; non-feedback). Three factors were observed and estimated: paretic step length, non-paretic step length, spatial asymmetry ratio at comfortable speed. Also, in VF and VF+RAC, calibration of feedback accuracy was measured. RESULTS: Results showed that paretic step length and spatial asymmetry ratio were significantly improved in VF+RAC compared to that of the VF and control (p < 0.05). Accuracies of paretic and non-paretic leg were significantly increased in VF+RAC than that of the VF (p < 0.05). CONCLUSIONS: According to this result, it seems that application of VF+RAC in treadmill gait significantly improved gait of these patients. Also, we can conclude that VF+RAC in treadmill gait is thought to be useful in clinical settings where there are many chronic patients who are in need of improvement in their gait ability.

Prolonged Walking with a Wearable System Providing Intelligent Auditory Input in People with Parkinson's Disease.

Rhythmic auditory cueing is a well-accepted tool for gait rehabilitation in Parkinson's disease (PD), which can now be applied in a performance-adapted fashion due to technological advance. This study investigated the immediate differences on gait during a prolonged, 30 min, walk with performance-adapted (intelligent) auditory cueing and verbal feedback provided by a wearable sensor-based system as alternatives for traditional cueing. Additionally, potential effects on self-perceived fatigue were assessed. Twenty-eight people with PD and 13 age-matched healthy elderly (HE) performed four 30 min walks with a wearable cue and feedback system. In randomized order, participants received: (1) continuous auditory cueing; (2) intelligent cueing (10 metronome beats triggered by a deviating walking rhythm); (3) intelligent feedback (verbal instructions triggered by a deviating walking rhythm); and (4) no external input. Fatigue was self-scored at rest and after walking during each session. The results showed that while HE were able to maintain cadence for 30 min during all conditions, cadence in PD significantly declined without input. With continuous cueing and intelligent feedback people with PD were able to maintain cadence (p = 0.04), although they were more physically fatigued than HE. Furthermore, cadence deviated significantly more in people with PD than in HE without input and particularly with intelligent feedback (both: p = 0.04). In PD, continuous and intelligent cueing induced significantly less deviations of cadence (p = 0.006). Altogether, this suggests that intelligent cueing is a suitable alternative for the continuous mode during prolonged walking in PD, as it induced similar effects on gait without generating levels of fatigue beyond that of HE.

Hearing visuo-tactile synchrony - Sound-induced proprioceptive drift in the invisible hand illusion.

The rubber hand illusion (RHI) and its variant the invisible hand illusion (IHI) are useful for investigating multisensory aspects of bodily self-consciousness. Here, we explored whether auditory conditioning during an RHI could enhance the trisensory visuo-tactile-proprioceptive interaction underlying the IHI. Our paradigm comprised of an IHI session that was followed by an RHI session and another IHI session. The IHI sessions had two parts presented in counterbalanced order. One part was conducted in silence, whereas the other part was conducted on the backdrop of metronome beats that occurred in synchrony with the brush movements used for the induction of the illusion. In a first experiment, the RHI session also involved metronome beats and was aimed at creating an associative memory between the brush stroking of a rubber hand and the sounds. An analysis of IHI sessions showed that the participants' perceived hand position drifted more towards the body-midline in the metronome relative to the silent condition without any sound-related session differences. Thus, the sounds, but not the auditory RHI conditioning, influenced the IHI. In a second experiment, the RHI session was conducted without metronome beats. This confirmed the conditioning-independent presence of sound-induced proprioceptive drift in the IHI. Together, these findings show that the influence of visuo-tactile integration on proprioceptive updating is modifiable by irrelevant auditory cues merely through the temporal correspondence between the visuo-tactile and auditory events.

The Influence of Auditory Information on Visual Size Adaptation.

Size perception can be influenced by several visual cues, such as spatial (e.g., depth or vergence) and temporal contextual cues (e.g., adaptation to steady visual stimulation). Nevertheless, perception is generally multisensory and other sensory modalities, such as auditory, can contribute to the functional estimation of the size of objects. In this study, we investigate whether auditory stimuli at different sound pitches can influence visual size perception after visual adaptation. To this aim, we used an adaptation paradigm (Pooresmaeili et al., 2013) in three experimental conditions: visual-only, visual-sound at 100 Hz and visual-sound at 9,000 Hz. We asked participants to judge the size of a test stimulus in a size discrimination task. First, we obtained a baseline for all conditions. In the visual-sound conditions, the auditory stimulus was concurrent to the test stimulus. Secondly, we repeated the task by presenting an adapter (twice as big as the reference stimulus) before the test stimulus. We replicated the size aftereffect in the visual-only condition: the test stimulus was perceived smaller than its physical size. The new finding is that we found the auditory stimuli have an effect on the perceived size of the test stimulus after visual adaptation: low frequency sound decreased the effect of visual adaptation, making the stimulus perceived bigger compared to the visual-only condition, and contrarily, the high frequency sound had the opposite effect, making the test size perceived even smaller.