Bimodal Integration Research Articles

Aversive Bimodal Associations Differently Impact Visual and Olfactory Memory Performance in Drosophila.

Animals form sensory associations and store them as memories to guide behavioral decisions. Although unimodal learning has been studied extensively in insects, it is important to explore sensory cues in combination because most behaviors require multimodal inputs. In our study, we optimized the T-maze to employ both visual and olfactory cues in a classical aversive learning paradigm in Drosophila melanogaster. In contrast to unimodal training, bimodal training evoked a significant short-term visual memory after a single training trial. Interestingly, the same protocol did not enhance short-term olfactory memory and even had a negative impact. However, compromised long-lasting olfactory memory significantly improved after bimodal training. Our study demonstrates that the effect of bimodal integration on learning is not always beneficial and is conditional upon the formed memory strengths. We postulate that flies utilize information on a need-to basis: bimodal training augments weakly formed memories while stronger associations are impacted differently.

Experimental and theoretical probe on mechano- and chemosensory integration in the insect antennal lobe.

In nature, olfactory signals are delivered to detectors-for example, insect antennae-by means of turbulent air, which exerts concurrent chemical and mechanical stimulation on the detectors. The antennal lobe, which is traditionally viewed as a chemosensory module, sits downstream of antennal inputs. We review experimental evidence showing that, in addition to being a chemosensory structure, antennal lobe neurons also respond to mechanosensory input in the form of wind speed. Benchmarked with empirical data, we constructed a dynamical model to simulate bimodal integration in the antennal lobe, with model dynamics yielding insights such as a positive correlation between the strength of mechanical input and the capacity to follow high frequency odor pulses, an important task in tracking odor sources. Furthermore, we combine experimental and theoretical results to develop a conceptual framework for viewing the functional significance of sensory integration within the antennal lobe. We formulate the testable hypothesis that the antennal lobe alternates between two distinct dynamical regimes, one which benefits odor plume tracking and one which promotes odor discrimination. We postulate that the strength of mechanical input, which correlates with behavioral contexts such being mid-flight versus hovering near a flower, triggers the transition from one regime to the other.

Cross-Modal Interaction and Integration Through Stimulus-Specific Adaptation in the Thalamic Reticular Nucleus of Rats.

Stimulus-specific adaptation (SSA), defined as a decrease in responses to a common stimulus that only partially generalizes to other rare stimuli, is a widespread phenomenon in the brain that is believed to be related to novelty detection. Although cross-modal sensory processing is also a widespread phenomenon, the interaction between the two phenomena is not well understood. In this study, the thalamic reticular nucleus (TRN), which is regarded as a hub of the attentional system that contains multi-modal neurons, was investigated. The results showed that SSA existed in an interactive oddball stimulation, which mimics stimulation changes from one modality to another. In the bimodal integration, SSA to bimodal stimulation was stronger than to visual stimulation alone but similar to auditory stimulation alone, which indicated a limited integrative effect. Collectively, the present results provide evidence for independent cross-modal processing in bimodal TRN neurons.

Emotion-modulated Recall: Congruency Effects of Nonverbal Facial and Vocal Cues on Semantic Recall

The current study had two main goals: First, to replicate the ‘bimodal integration’ effect (i.e. the automatic integration of crossmodal stimuli, namely facial emotions and emotional prosody); and second, to investigate whether this phenomenon facilitates or impairs the intake and retention of unattended verbal content. The study borrowed from previous bimodal integration designs and included a two-alternative forced-choice (2AFC) task, where subjects were instructed to identify the emotion of a face (as either ‘angry’ or ‘happy’) while ignoring a concurrently presented sentence (spoken in an angry, happy, or neutral prosody), after which a surprise recall was administered to investigate effects on semantic content retention. While bimodal integration effects were replicated (i.e. faster and more accurate emotion identification under congruent conditions), congruency effects were not found for semantic recall. Overall, semantic recall was better for trials with emotional (vs. neutral) faces, and worse in trials with happy (vs. angry or neutral) prosody. Taken together, our findings suggest that when individuals focus their attention on evaluation of facial expressions, they implicitly integrate nonverbal emotional vocal cues (i.e. hedonic valence or emotional tone of accompanying sentences), and devote less attention to their semantic content. While the impairing effect of happy prosody on recall may indicate an emotional interference effect, more research is required to uncover potential prosody-specific effects. All supplemental online materials can be found on OSF (https://osf.io/am9p2/).

Bimodal Hearing with Pediatric Cochlear Implant Recipients: Effect of Acoustic Bandwidth.

The primary purpose of this study was to examine the effect of acoustic bandwidth on bimodal benefit for speech understanding in pediatric cochlear implant (CI) recipients. Ten children (6-13 years) with CIs utilizing a bimodal hearing configuration participated in this study. Speech understanding was assessed via recorded Pediatric AzBio sentences presented in a 10-talker babble. The CI stimulus was always unprocessed and the low-pass filtered acoustic stimuli were delivered to the non-CI ear with the following cutoff frequencies: 250, 500, 750, 1000, and 1500 Hz. Tertiary referral center. Sentence recognition in noise for the acoustic-alone, CI-alone, and bimodal listening conditions. The primary findings were: (1) children gained significant bimodal benefit with 250 Hz, and (2) children demonstrated no statistically significant additional bimodal benefit with increasing acoustic bandwidth. Acoustic bandwidth effects for pediatric CI recipients were significantly different than those documented in the literature for adult CI recipients. Specifically, this group of pediatric CI recipients demonstrated no increases in bimodal benefit with increases in acoustic bandwidth, primarily consistent with a segregation theory of bimodal integration.

Modulation of early auditory processing by visual information: Prediction or bimodal integration?

What happens if a visual cue misleads auditory expectations? Previous studies revealed an early visuo–auditory incongruency effect, so-called incongruency response (IR) of the auditory event-related brain potential (ERP), occurring 100 ms after onset of the sound being incongruent to the preceding visual cue. So far, this effect has been ascribed to reflect the mismatch between auditory sensory expectation activated by visual predictive information and the actual sensory input. Thus, an IR should be confined to an asynchronous presentation of visual cue and sound. Alternatively, one could argue that frequently presented congruent visual-cue–sound combinations are integrated into a bimodal representation whereby violation of the visual–auditory relationship results in a bimodal feature mismatch (the IR should be obtained with asynchronous and with synchronous presentation). In an asynchronous condition, an either high-pitched or low-pitched sound was preceded by a visual note symbol presented above or below a fixation cross (90% congruent; 10% incongruent), while in a synchronous condition, both were presented simultaneously. High-pitched and low-pitched sounds were presented with different probabilities (83% vs. 17%) to form a strong association between bimodal stimuli. In both conditions, tones with pitch incongruent with the location of the note symbols elicited incongruency effects in the N2 and P3 ERPs; however, the IR was only elicited in the asynchronous condition. This finding supports the sensorial prediction error hypothesis stating that the amplitude of the auditory ERP 100 ms after sound onset is enhanced in response to unexpected compared with expected but otherwise identical sounds.

Image and WLAN Bimodal Integration for Indoor User Localization

Recently, we experience the increasing prevalence of wearable cameras, some of which feature Wireless Local Area Network (WLAN) connectivity, and the abundance of mobile devices equipped with on-board camera and WLAN modules. Motivated by this fact, this work presents an indoor localization system that leverages both imagery and WLAN data for enabling and supporting a wide variety of envisaged location-aware applications ranging from ambient and assisted living to indoor mobile gaming and retail analytics. The proposed solution integrates two complementary localization approaches, i.e., one based on WLAN and another one based on image location-dependent data, using a fusion engine. Two fusion strategies are developed and investigated to meet different requirements in terms of accuracy, run time, and power consumption. The one is a light-weight threshold-based approach that combines the location outputs of two localization algorithms, namely a WLAN-based algorithm that processes signal strength readings from the surrounding wireless infrastructure using an extended Naive Bayes approach and an image-based algorithm that follows a novel approach based on hierarchical vocabulary tree of SURF (Speeded Up Robust Features) descriptors. The second fusion strategy employs a particle filter algorithm that operates directly on the WLAN and image readings and also includes prior position estimation information in the localization process. Extensive experimental results using real-life data from an indoor office environment indicate that the proposed fusion strategies perform well and are competitive against standalone WLAN and imaged-based algorithms, as well as alternative fusion localization solutions.

Visual and Auditory Information During Decision Making in Sport.

In 2 experiments, the authors investigated the effects of bimodal integration in a sport-specific task. Beach volleyball players were required to make a tactical decision, responding either verbally or via a motor response, after being presented with visual, auditory, or both kinds of stimuli in a beach volleyball scenario. In Experiment 1, players made the correct decision in a game situation more often when visual and auditory information were congruent than in trials in which they experienced only one of the modalities or incongruent information. Decision-making accuracy was greater when motor, rather than verbal, responses were given. Experiment 2 replicated this congruence effect using different stimulus material and showed a decreasing effect of visual stimulation on decision making as a function of shorter visual stimulus durations. In conclusion, this study shows that bimodal integration of congruent visual and auditory information results in more accurate decision making in sport than unimodal information.

Aversive Training of Honey Bees in an Automated Y-Maze.

Honeybees have remarkable learning abilities given their small brains, and have thus been established as a powerful model organism for the study of learning and memory. Most of our current knowledge is based on appetitive paradigms, in which a previously neutral stimulus (e.g., a visual, olfactory, or tactile stimulus) is paired with a reward. Here, we present a novel apparatus, the yAPIS, for aversive training of walking honey bees. This system consists in three arms of equal length and at 120° from each other. Within each arm, colored lights (λ = 375, 465 or 520 nm) or odors (here limonene or linalool) can be delivered to provide conditioned stimuli (CS). A metal grid placed on the floor and roof delivers the punishment in the form of mild electric shocks (unconditioned stimulus, US). Our training protocol followed a fully classical procedure, in which the bee was exposed sequentially to a CS paired with shocks (CS+) and to another CS not punished (CS-). Learning performance was measured during a second phase, which took advantage of the Y-shape of the apparatus and of real-time tracking to present the bee with a choice situation, e.g., between the CS+ and the CS-. Bees reliably chose the CS- over the CS+ after only a few training trials with either colors or odors, and retained this memory for at least a day, except for the shorter wavelength (λ = 375 nm) that produced mixed results. This behavior was largely the result of the bees avoiding the CS+, as no evidence was found for attraction to the CS-. Interestingly, trained bees initially placed in the CS+ spontaneously escaped to a CS- arm if given the opportunity, even though they could never do so during the training. Finally, honey bees trained with compound stimuli (color + odor) later avoided either components of the CS+. Thus, the yAPIS is a fast, versatile and high-throughput way to train honey bees in aversive paradigms. It also opens the door for controlled laboratory experiments investigating bimodal integration and learning, a field that remains in its infancy.

Multi-modal language input: A learned superadditive effect

AbstractReview of psychological and language acquisition research into seeing faces while listening, seeing gesture while listening, illustrated text, reading while listening, and same language subtitled video, confirms that bi-modal input has a consistently positive effect on language learning over a variety of input types. This effect is normally discussed using a simple additive model where bi-modal input increases the total amount of data and adds redundancy to duplicated input thus increasing comprehension and then learning. Parallel studies in neuroscience suggest that bi-modal integration is a general effect using common brain areas and following common neural paths. Neuroscience also shows that bi-modal effects are more complex than simple addition, showing early integration of inputs, a learning/developmental effect, and a superadditive effect for integrated bi-modal input. The different bodies of research produce a revised model of bi-modal input as a learned, active system. The implications for language learning are that bi- or multi-modal input can powerfully enhance language learning and that the learning benefits of such input will increase alongside the development of neurological integration of the inputs.

Bimodal benefits in Mandarin-speaking cochlear implant users with contralateral residual acoustic hearing

Objective: The present study aimed to measure bimodal benefits and probe their underlying mechanisms in Mandarin-speaking cochlear implant (CI) subjects who had contralateral residual acoustic hearing. Design: The subjects recognised words or phonemes from the Mandarin Lexical Neighborhood Test in noise at a 10-dB signal-to-noise ratio (SNR) with acoustic stimulation, electric stimulation or the combined bimodal stimulation. Study sample: Thirteen Mandarin-speaking subjects wore a CI in one ear and had residual acoustic hearing in the contralateral ear. Six of the subjects (5.2–13.0 years) had pre-lingual onset of severe hearing loss, and seven of them (8.6–45.8 years) had post-lingual onset of severe hearing loss. Results: Both groups of subjects produced a significant bimodal benefit in word recognition in noise. Consonants and tones accounted for the bimodal benefit. The bimodal integration efficiency was negatively correlated with the duration of deafness in the implanted ear for vowel recognition but positively correlated with CI or bimodal experience for consonant recognition. Conclusions: The present results support preservation of residual acoustic hearing, early cochlear implantation and continuous use of bimodal hearing for subjects who have significant residual hearing in the non-implanted ear.

Integrated multimodal network approach to PET and MRI based on multidimensional persistent homology.

Finding underlying relationships among multiple imaging modalities in a coherent fashion is one of the challenging problems in multimodal analysis. In this study, we propose a novel approach based on multidimensional persistence. In the extension of the previous threshold-free method of persistent homology, we visualize and discriminate the topological change of integrated brain networks by varying not only threshold but also mixing ratio between two different imaging modalities. The multidimensional persistence is implemented by a new bimodal integration method called 1D projection. When the mixing ratio is predefined, it constructs an integrated edge weight matrix by projecting two different connectivity information onto the one dimensional shared space. We applied the proposed methods to PET and MRI data from 23 attention deficit hyperactivity disorder (ADHD) children, 21 autism spectrum disorder (ASD), and 10 pediatric control subjects. From the results, we found that the brain networks of ASD, ADHD children and controls differ, with ASD and ADHD showing asymmetrical changes of connected structures between metabolic and morphological connectivities. The difference of connected structure between ASD and the controls was mainly observed in the metabolic connectivity. However, ADHD showed the maximum difference when two connectivity information were integrated with the ratio 0.6. These results provide a multidimensional homological understanding of disease-related PET and MRI networks that disclose the network association with ASD and ADHD. Hum Brain Mapp 38:1387-1402, 2017. © 2016 Wiley Periodicals, Inc.

Influence of attention on bimodal integration during emotional change decoding: ERP evidence

Recent findings on audiovisual emotional interactions suggest that selective attention affects cross-sensory interaction from an early processing stage. However, the influence of attention manipulation on facial-vocal integration during emotional change perception is still elusive at this point. To address this issue, we asked participants to detect emotional changes conveyed by prosodies (vocal task) or facial expressions (facial task) while facial, vocal, and facial-vocal expressions were presented. At the same time, behavioral responses and electroencephalogram (EEG) were recorded. Behavioral results showed that bimodal emotional changes were detected with shorter response latencies compared to each unimodal condition, suggesting that bimodal emotional cues facilitated emotional change detection. Moreover, while the P3 amplitudes were larger for the bimodal change condition than for the sum of the two unimodal conditions regardless of attention direction, the N1 amplitudes were larger for the bimodal emotional change condition than for the sum of the two unimodal conditions under the attend-voice condition, but not under the attend-face condition. These findings suggest that selective attention modulates facial-vocal integration during emotional change perception in early sensory processing, but not in late cognitive processing stages.

Identification of key regulators of pancreatic cancer progression through multidimensional systems-level analysis.

BackgroundPancreatic cancer is an aggressive cancer with dismal prognosis, urgently necessitating better biomarkers to improve therapeutic options and early diagnosis. Traditional approaches of biomarker detection that consider only one aspect of the biological continuum like gene expression alone are limited in their scope and lack robustness in identifying the key regulators of the disease. We have adopted a multidimensional approach involving the cross-talk between the omics spaces to identify key regulators of disease progression.MethodsMultidimensional domain-specific disease signatures were obtained using rank-based meta-analysis of individual omics profiles (mRNA, miRNA, DNA methylation) related to pancreatic ductal adenocarcinoma (PDAC). These domain-specific PDAC signatures were integrated to identify genes that were affected across multiple dimensions of omics space in PDAC (genes under multiple regulatory controls, GMCs). To further pin down the regulators of PDAC pathophysiology, a systems-level network was generated from knowledge-based interaction information applied to the above identified GMCs. Key regulators were identified from the GMC network based on network statistics and their functional importance was validated using gene set enrichment analysis and survival analysis.ResultsRank-based meta-analysis identified 5391 genes, 109 miRNAs and 2081 methylation-sites significantly differentially expressed in PDAC (false discovery rate ≤ 0.05). Bimodal integration of meta-analysis signatures revealed 1150 and 715 genes regulated by miRNAs and methylation, respectively. Further analysis identified 189 altered genes that are commonly regulated by miRNA and methylation, hence considered GMCs. Systems-level analysis of the scale-free GMCs network identified eight potential key regulator hubs, namely E2F3, HMGA2, RASA1, IRS1, NUAK1, ACTN1, SKI and DLL1, associated with important pathways driving cancer progression. Survival analysis on individual key regulators revealed that higher expression of IRS1 and DLL1 and lower expression of HMGA2, ACTN1 and SKI were associated with better survival probabilities.ConclusionsIt is evident from the results that our hierarchical systems-level multidimensional analysis approach has been successful in isolating the converging regulatory modules and associated key regulatory molecules that are potential biomarkers for pancreatic cancer progression.Electronic supplementary materialThe online version of this article (doi:10.1186/s13073-016-0282-3) contains supplementary material, which is available to authorized users.

Neural correlates of audiotactile phonetic processing in early-blind readers: an fMRI study.

Reading is a multisensory function that relies on arbitrary associations between auditory speech sounds and symbols from a second modality. Studies of bimodal phonetic perception have mostly investigated the integration of visual letters and speech sounds. Blind readers perform an analogous task by using tactile Braille letters instead of visual letters. The neural underpinnings of audiotactile phonetic processing have not been studied before. We used functional magnetic resonance imaging to reveal the neural correlates of audiotactile phonetic processing in 16 early-blind Braille readers. Braille letters and corresponding speech sounds were presented in unimodal, and congruent/incongruent bimodal configurations. We also used a behavioral task to measure the speed of blind readers in identifying letters presented via tactile and/or auditory modalities. Reaction times for tactile stimuli were faster. The reaction times for bimodal stimuli were equal to those for the slower auditory-only stimuli. fMRI analyses revealed the convergence of unimodal auditory and unimodal tactile responses in areas of the right precentral gyrus and bilateral crus I of the cerebellum. The left and right planum temporale fulfilled the 'max criterion' for bimodal integration, but activities of these areas were not sensitive to the phonetical congruency between sounds and Braille letters. Nevertheless, congruency effects were found in regions of frontal lobe and cerebellum. Our findings suggest that, unlike sighted readers who are assumed to have amodal phonetic representations, blind readers probably process letters and sounds separately. We discuss that this distinction might be due to mal-development of multisensory neural circuits in early blinds or it might be due to inherent differences between Braille and print reading mechanisms.

The effect of amplitude envelope on spatial ventriloquism

Spatial ventriloquism occurs when a visual event and an auditory event happen simultaneously; location judgments for the sound source become biased toward the location of the visual event. Evidence suggests the brain attributes greater weight to spatial information provided by the visual stimulus because the visual system offers better spatial acuity; when the visual stimulus is deteriorated, the visual bias is reduced. Thus, the brain performs optimal bimodal integration: greater weight is given to the modality which provides more information. The present study aims to determine whether the amplitude envelope of sounds provides spatial localization information to the perceptual system. We used a psychophysical staircase procedure to measure spatial ventriloquism experienced by participants for sounds with percussive, flat, and time-reversed percussive envelopes. We hypothesize that percussive and reverse-percussive sounds provide more information and thus better sound localization acuity than flat sounds, which would result in smaller degrees of spatial ventriloquism for the former than the latter. The results yield insight into the brain’s use of auditory cues in audio-visual integration.

Subjective evaluation of the audiovisual spatial congruence in the case of stereoscopic-3D video and Wave Field Synthesis

While 3D cinema is becoming increasingly established, little effort has been focused on the general problem of producing a 3D sound scene spatially coherent with the visual content of a stereoscopic-3D (s-3D) movie. The perceptual relevance of such a spatial audiovisual coherence is of significant interest. In this paper, a subjective experiment is carried out where an angular error between an s-3D video and a spatially accurate sound reproduced through Wave Field Synthesis (WFS) is simulated. The psychometric curve is measured with the method of constant stimuli, and the threshold for bimodal integration is estimated. The impact of the presence of background noise is also investigated. A comparison is made between the case without any background noise and the case with an SNR of 4dBA. Estimates of the thresholds and the slopes, as well as their confidence intervals, are obtained for each level of background noise. When background noise was present, the point of subjective equality (PSE) was higher (19.4° instead of 18.3°) and the slope was steeper (−0.077 instead of −0.062 per degree). Because of the overlap between the confidence intervals, however, it was not possible to statistically differentiate between the two levels of noise. The implications for the sound reproduction in a cinema theater are discussed.

Multi-sensory integration in brainstem and auditory cortex

Tinnitus is the perception of sound in the absence of a physical sound stimulus. It is thought to arise from aberrant neural activity within central auditory pathways that may be influenced by multiple brain centers, including the somatosensory system. Auditory-somatosensory (bimodal) integration occurs in the dorsal cochlear nucleus (DCN), where electrical activation of somatosensory regions alters pyramidal cell spike timing and rates of sound stimuli. Moreover, in conditions of tinnitus, bimodal integration in DCN is enhanced, producing greater spontaneous and sound-driven neural activity, which are neural correlates of tinnitus. In primary auditory cortex (A1), a similar auditory-somatosensory integration has been described in the normal system (Lakatos et al., 2007), where sub-threshold multisensory modulation may be a direct reflection of subcortical multisensory responses (Tyll et al., 2011). The present work utilized simultaneous recordings from both DCN and A1 to directly compare bimodal integration across these separate brain stations of the intact auditory pathway. Four-shank, 32-channel electrodes were placed in DCN and A1 to simultaneously record tone-evoked unit activity in the presence and absence of spinal trigeminal nucleus (Sp5) electrical activation. Bimodal stimulation led to long-lasting facilitation or suppression of single and multi-unit responses to subsequent sound in both DCN and A1. Immediate (bimodal response) and long-lasting (bimodal plasticity) effects of Sp5-tone stimulation were facilitation or suppression of tone-evoked firing rates in DCN and A1 at all Sp5-tone pairing intervals (10, 20, and 40 ms), and greater suppression at 20 ms pairing-intervals for single unit responses. Understanding the complex relationships between DCN and A1 bimodal processing in the normal animal provides the basis for studying its disruption in hearing loss and tinnitus models. This article is part of a Special Issue entitled: Tinnitus Neuroscience.

‘Seeing tunes.’ The role of visual gestures in tune interpretation

AbstractOne of the unresolved questions in audiovisual prosody is the relative contribution of acoustic and visual cues to the expression of prosodic meaning. Though the majority of studies on audiovisual prosody have found a complementary mode of processing whereby sight provides relatively weak and redundant information in comparison with strong auditory cues, other work has found that sight provides information more efficiently than hearing. In Catalan, a pitch range contrast in a rising-falling nuclear configuration conveys a difference between a contrastive focus statement and an echo question. The main goal of this study is to investigate the relative contribution of visual cues in conveying this distinction. Twenty native speakers of Central Catalan participated in two identification tasks in which they had to decide between a focus statement and a question interpretation. Experiment 1 used a pitch range auditory continuum combined with two congruent and incongruent videotapes showing the facial gestures that are characteristic of the two pragmatic meanings. Experiment 2 used the same auditory continuum in combination with another continuum for facial gestures produced using a digital image-morphing technique. The responses and reaction times obtained in both experiments revealed a consistent reliance on visual cues in the listener's decisions, but also a consistent effect of the auditory stimulus. We argue that although facial gestures are the most influential elements that Catalan listeners rely on to decide between contrastive focus and echo question interpretations, bimodal integration with the acoustic cues is necessary for perceptual processing to be accurate and fast. Finally, we discuss the implications of these results for models of audiovisual processing.

Enhanced effectiveness in visuo‐haptic object‐selective brain regions with increasing stimulus salience

The occipital and parietal lobes contain regions that are recruited for both visual and haptic object processing. The purpose of the present study was to characterize the underlying neural mechanisms for bimodal integration of vision and haptics in these visuo-haptic object-selective brain regions to find out whether these brain regions are sites of neuronal or areal convergence. Our sensory conditions consisted of visual-only (V), haptic-only (H), and visuo-haptic (VH), which allowed us to evaluate integration using the superadditivity metric. We also presented each stimulus condition at two different levels of signal-to-noise ratio or salience. The salience manipulation allowed us to assess integration using the rule of inverse effectiveness. We were able to localize previously described visuo-haptic object-selective regions in the lateral occipital cortex (lateral occipital tactile-visual area) and the intraparietal sulcus, and also localized a new region in the left anterior fusiform gyrus. There was no evidence of superadditivity with the VH stimulus at either level of salience in any of the regions. There was, however, a strong effect of salience on multisensory enhancement: the response to the VH stimulus was more enhanced at higher salience across all regions. In other words, the regions showed enhanced integration of the VH stimulus with increasing effectiveness of the unisensory stimuli. We called the effect "enhanced effectiveness." The presence of enhanced effectiveness in visuo-haptic object-selective brain regions demonstrates neuronal convergence of visual and haptic sensory inputs for the purpose of processing object shape.