Subcortical contributions to the sense of body ownership.

The sense of body ownership (i.e. the feeling that our body or its parts belong to us) plays a key role in bodily self-consciousness and is believed to stem from multisensory integration. Experimental paradigms such as the rubber hand illusion have been developed to allow the controlled manipulation of body ownership in laboratory settings, providing effective tools for investigating malleability in the sense of body ownership and the boundaries that distinguish self from other. Neuroimaging studies of body ownership converge on the involvement of several cortical regions, including the premotor cortex and posterior parietal cortex. However, relatively less attention has been paid to subcortical structures that may also contribute to body ownership perception, such as the cerebellum and putamen. Here, on the basis of neuroimaging and neuropsychological observations, we provide an overview of relevant subcortical regions and consider their potential role in generating and maintaining a sense of ownership over the body. We also suggest novel avenues for future research targeting the role of subcortical regions in making sense of the body as our own.

Neural Substrates of Body Ownership and Agency during Voluntary Movement.

Body ownership and the sense of agency are two central aspects of bodily self-consciousness. While multiple neuroimaging studies have investigated the neural correlates of body ownership and agency separately, few studies have investigated the relationship between these two aspects during voluntary movement when such experiences naturally combine. By eliciting the moving rubber hand illusion with active or passive finger movements during functional magnetic resonance imaging, we isolated activations reflecting the sense of body ownership and agency, respectively, as well as their interaction, and assessed their overlap and anatomic segregation. We found that perceived hand ownership was associated with activity in premotor, posterior parietal, and cerebellar regions, whereas the sense of agency over the movements of the hand was related to activity in the dorsal premotor cortex and superior temporal cortex. Moreover, one section of the dorsal premotor cortex showed overlapping activity for ownership and agency, and somatosensory cortical activity reflected the interaction of ownership and agency with higher activity when both agency and ownership were experienced. We further found that activations previously attributed to agency in the left insular cortex and right temporoparietal junction reflected the synchrony or asynchrony of visuoproprioceptive stimuli rather than agency. Collectively, these results reveal the neural bases of agency and ownership during voluntary movement. Although the neural representations of these two experiences are largely distinct, there are interactions and functional neuroanatomical overlap during their combination, which has bearing on theories on bodily self-consciousness.SIGNIFICANCE STATEMENT How does the brain generate the sense of being in control of bodily movement (agency) and the sense that body parts belong to one's body (body ownership)? Using fMRI and a bodily illusion triggered by movement, we found that agency is associated with activity in premotor cortex and temporal cortex, and body ownership with activity in premotor, posterior parietal, and cerebellar regions. The activations reflecting the two sensations were largely distinct, but there was overlap in premotor cortex and an interaction in somatosensory cortex. These findings advance our understanding of the neural bases of and interplay between agency and body ownership during voluntary movement, which has implications for the development of advanced controllable prosthetic limbs that feel like real limbs.

Brief Temporal Perturbations in Somatosensory Reafference Disrupt Perceptual and Neural Attenuation and Increase Supplementary Motor Area-Cerebellar Connectivity.

Intrinsic delays in sensory feedback can be detrimental for motor control. As a compensation strategy, the brain predicts the sensory consequences of movement via a forward model on the basis of a copy of the motor command. Using these predictions, the brain attenuates somatosensory reafference to facilitate the processing of exafferent information. Theoretically, this predictive attenuation is disrupted by (even minimal) temporal errors between the predicted and actual reafference; however, direct evidence of such disruption is lacking as previous neuroimaging studies contrasted nondelayed reafferent input with exafferent input. Here, we combined psychophysics with functional magnetic resonance imaging to test whether subtle perturbations in the timing of somatosensory reafference disrupt its predictive processing. Twenty-eight participants (14 women) generated touches on their left index finger by tapping a sensor with their right index finger. The touches on the left index finger were delivered close to the time of contact of the two fingers or with a temporal perturbation (i.e., 153 ms delay). We found that such a brief temporal perturbation disrupted the attenuation of the somatosensory reafference at both the perceptual and neural levels, leading to greater somatosensory and cerebellar responses and weaker somatosensory connectivity with the cerebellum, proportional to the perceptual changes. We interpret these effects as the failure of the forward model to predictively attenuate the perturbed somatosensory reafference. Moreover, we observed increased connectivity of the supplementary motor area with the cerebellum during the perturbations, which could indicate the communication of the temporal prediction error back to the motor centers.SIGNIFICANCE STATEMENT Our brain receives somatosensory feedback from our movements with a delay. To counteract these delays, motor control theories postulate that the brain predicts the timing of somatosensory consequences of our movements and attenuates sensations received at that time. Thus, a self-generated touch feels weaker than an identical external touch. However, how subtle temporal errors between the predicted and actual somatosensory feedback perturb this predictive attenuation remains unknown. We show that such errors make the otherwise attenuated touch feel stronger, elicit stronger somatosensory responses, weaken cerebellar connectivity with somatosensory areas, and increase this connectivity with motor areas. These findings show that motor and cerebellar areas are fundamental in forming temporal predictions about the sensory consequences of our movements.

Texture congruence modulates perceptual bias but not sensitivity to visuotactile stimulation during the rubber hand illusion.

The sense of body ownership is the feeling that one's body belongs to oneself. To study body ownership, researchers use bodily illusions, such as the rubber hand illusion (RHI), which involves experiencing a visible rubber hand as part of one's body when the rubber hand is stroked simultaneously with the hidden real hand. The RHI is based on a combination of vision, touch, and proprioceptive information following the principles of multisensory integration. It has been posited that texture incongruence between rubber hand and real hand weakens the RHI, but the underlying mechanisms remain poorly understood. To investigate this, we recently developed a novel psychophysical RHI paradigm. Based on fitting psychometric functions, we discovered the RHI resulted in shifts in the point of subjective equality when the rubber hand and the real hand were stroked with matching materials. We analysed these datasets further by using signal detection theory analysis, which distinguishes between the participants' sensitivity to visuotactile stimulation and the associated perceptual bias. We found that texture incongruence influences the RHI's perceptual bias but not its sensitivity to visuotactile stimulation. We observed that the texture congruence bias effect was the strongest in shorter visuotactile asynchronies (50-100 ms) and weaker in longer asynchronies (200 ms). These results suggest texture-related perceptual bias is most prominent when the illusion's sensitivity is at its lowest. Our findings shed light on the intricate interactions between top-down and bottom-up processes in body ownership, the links between body ownership and multisensory integration, and the impact of texture congruence on the RHI.

Causal Inference of Body Ownership in the Posterior Parietal Cortex.

How do we come to sense that a hand in view belongs to our own body or not? Previous studies have suggested that the integration of vision and somatosensation in the frontoparietal areas plays a critical role in the sense of body ownership (i.e., the multisensory perception of limbs and body parts as our own). However, little is known about how these areas implement the multisensory integration process at the computational level and whether activity predicts illusion elicitation in individual participants on a trial-by-trial basis. To address these questions, we used functional magnetic resonance imaging and a rubber hand illusion-detection task and fitted the registered neural responses to a Bayesian causal inference model of body ownership. Thirty healthy human participants (male and female) performed 12 s trials with varying degrees of asynchronously delivered visual and tactile stimuli of a rubber hand (in view) and a (hidden) real hand. After the 12 s period, participants had to judge whether the rubber hand felt like their own. As hypothesized, activity in the premotor and posterior parietal cortices was related to illusion elicitation at the level of individual participants and trials. Importantly, activity in the posterior parietal cortex fit the predicted probability of illusion emergence of the Bayesian causal inference model based on each participant's behavioral response profile. Our findings suggest an important role for the posterior parietal cortex in implementing Bayesian causal inference of body ownership and reveal how trial-by-trial variations in neural signatures of multisensory integration relate to the elicitation of the rubber hand illusion.SIGNIFICANCE STATEMENT How does the brain create a coherent perceptual experience of one's own body based on information from the different senses? We examined how the likelihood of eliciting a classical bodily illusion that depends on vision and touch-the rubber hand illusion-is related to neural activity measured by functional magnetic resonance imaging. We found that trial-by-trial variations in the neural signal in the posterior parietal cortex, a well known center for sensory integration, fitted a statistical function that describes how likely it is that the brain infers that a rubber hand is one's own given the available visual and tactile evidence. Thus, probabilistic analysis of sensory information in the parietal lobe underlies our unitary sense of bodily self.

Synchronous motor imagery and visual feedback of finger movement elicit the moving rubber hand illusion, at least in illusion-susceptible individuals.

Recent evidence suggests that imagined auditory and visual sensory stimuli can be integrated with real sensory information from a different sensory modality to change the perception of external events via cross-modal multisensory integration mechanisms. Here, we explored whether imagined voluntary movements can integrate visual and proprioceptive cues to change how we perceive our own limbs in space. Participants viewed a robotic hand wearing a glove repetitively moving its right index finger up and down at a frequency of 1 Hz, while they imagined executing the corresponding movements synchronously or asynchronously (kinesthetic-motor imagery); electromyography (EMG) from the participants' right index flexor muscle confirmed that the participants kept their hand relaxed while imagining the movements. The questionnaire results revealed that the synchronously imagined movements elicited illusory ownership and a sense of agency over the moving robotic hand-the moving rubber hand illusion-compared with asynchronously imagined movements; individuals who affirmed experiencing the illusion with real synchronous movement also did so with synchronous imagined movements. The results from a proprioceptive drift task further demonstrated a shift in the perceived location of the participants' real hand toward the robotic hand in the synchronous versus the asynchronous motor imagery condition. These results suggest that kinesthetic motor imagery can be used to replace veridical congruent somatosensory feedback from a moving finger in the moving rubber hand illusion to trigger illusory body ownership and agency, but only if the temporal congruence rule of the illusion is obeyed. This observation extends previous studies on the integration of mental imagery and sensory perception to the case of multisensory bodily awareness, which has potentially important implications for research into embodiment of brain-computer interface controlled robotic prostheses and computer-generated limbs in virtual reality.

Modeling affective touch pleasantness across skin types at the individual level reveals a reliable and stable basic function.

Touch is perceived most pleasant when delivered at velocities known to optimally activate the C-tactile afferent system. At the group level, pleasantness ratings of touch delivered at velocities in the range between 0.3 and 30 cm/s follow an inverted-U shape curve, with maximum pleasantness between 1 and 10 cm/s. However, the prevalence, reliability, and stability of this function at the individual level and across skin types based on hair density remains unknown. Here, we tested a range of seven velocities (0.3, 1, 3, 6, 9, 18, and 27 cm/s) delivered with a soft brush, on both hairy (forearm and dorsal hand) and nonhairy skin (palm) in 123 participants. Our results suggest that the relationship between pleasantness and velocity of touch is significantly best described by a negative quadratic model at the individual level in the majority of participants both on hairy (67.1%) and nonhairy (62.6%) skin, a larger extent than previously reported. Higher interoceptive accuracy and self-reported depression were related to a better fit of the quadratic model and the steepness of the curve, respectively. The prevalence of the quadratic model at the individual level was stable across body sites (62.6%, experiment 1), across two experimental sessions (73%-78%, experiment 2), and regardless of the number of repetitions of each velocity (experiment 3). Thus, the individual perception of tactile pleasantness follows a characteristic velocity-dependent function across skin types and shows trait characteristics. Future studies can investigate further the possibility to use affective touch as a behavioral biomarker for mental health disorders.NEW & NOTEWORTHY Touch is perceived as most pleasant when delivered at slow, caress-like velocities, known to activate C-tactile afferents. At the group level, tactile pleasantness and velocity of touch show a reliable pattern of relationship on hairy skin. Here, we found that the perception of tactile pleasantness follows a consistent pattern also at the individual level, across skin types and testing sessions. However, individual differences in interoceptive abilities and self-reported depression do play a role.

Functional Connectivity between the Cerebellum and Somatosensory Areas Implements the Attenuation of Self-Generated Touch.

Since the early 1970s, numerous behavioral studies have shown that self-generated touch feels less intense and less ticklish than the same touch applied externally. Computational motor control theories have suggested that cerebellar internal models predict the somatosensory consequences of our movements and that these predictions attenuate the perception of the actual touch. Despite this influential theoretical framework, little is known about the neural basis of this predictive attenuation. This is due to the limited number of neuroimaging studies, the presence of conflicting results about the role and the location of cerebellar activity, and the lack of behavioral measures accompanying the neural findings. Here, we combined psychophysics with fMRI to detect the neural processes underlying somatosensory attenuation in male and female healthy human participants. Activity in bilateral secondary somatosensory areas was attenuated when the touch was presented during a self-generated movement (self-generated touch) than in the absence of movement (external touch). An additional attenuation effect was observed in the cerebellum that is ipsilateral to the passive limb receiving the touch. Importantly, we further found that the degree of functional connectivity between the ipsilateral cerebellum and the contralateral primary and bilateral secondary somatosensory areas was linearly and positively related to the degree of behaviorally assessed attenuation; that is, the more participants perceptually attenuated their self-generated touches, the stronger this corticocerebellar coupling. Collectively, these results suggest that the ipsilateral cerebellum is fundamental in predicting self-generated touch and that this structure implements somatosensory attenuation via its functional connectivity with somatosensory areas.SIGNIFICANCE STATEMENT When we touch our hand with the other, the resulting sensation feels less intense than when another person or a machine touches our hand with the same intensity. Early computational motor control theories have proposed that the cerebellum predicts and cancels the sensory consequences of our movements; however, the neural correlates of this cancelation remain unknown. By means of fMRI, we show that the more participants attenuate the perception of their self-generated touch, the stronger the functional connectivity between the cerebellum and the somatosensory cortical areas. This provides conclusive evidence about the role of the cerebellum in predicting the sensory feedback of our movements and in attenuating the associated percepts via its connections to early somatosensory areas.

Little evidence for an effect of the rubber hand illusion on basic movement.

Body ownership refers to the distinct sensation that our observed body belongs to us, which is believed to stem from multisensory integration. This is commonly shown through the rubber hand illusion (RHI), which induces a sense of ownership over a false limb. Whilst the RHI may interfere with object-directed action and alter motor cortical activity, it is not yet clear whether a sense of ownership over an artificial hand has functional consequences for movement production per se. As such, we performed two motion-tracking experiments (n = 117) to examine the effects of the RHI on the reaction time, acceleration, and velocity of rapid index finger abduction. We observed little convincing evidence that the induction of the RHI altered these kinematic variables. Moreover, the subjective sensations of rubber hand ownership, referral of touch, and agency did not convincingly correlate with kinematic variables, and nor did proprioceptive drift, suggesting that changes in body representation elicited by the RHI may not influence basic movement. Whilst experiment 1 suggested that individuals reporting a greater sensation of the real hand disappearing performed movements with smaller acceleration and velocity following illusion induction, we did not replicate this effect in a second experiment, suggesting that these effects may be small or not particularly robust. Overall, these results indicate that manipulating the conscious experience of body ownership has little impact on basic motor control, at least in the RHI with healthy participants.

No evidence for somatosensory attenuation during action observation of self-touch.

The discovery of mirror neurons in the macaque brain in the 1990s triggered investigations on putative human mirror neurons and their potential functionality. The leading proposed function has been action understanding: Accordingly, we understand the actions of others by 'simulating' them in our own motor system through a direct matching of the visual information to our own motor programmes. Furthermore, it has been proposed that this simulation involves the prediction of the sensory consequences of the observed action, similar to the prediction of the sensory consequences of our executed actions. Here, we tested this proposal by quantifying somatosensory attenuation behaviourally during action observation. Somatosensory attenuation manifests during voluntary action and refers to the perception of self-generated touches as less intense than identical externally generated touches because the self-generated touches are predicted from the motor command. Therefore, we reasoned that if an observer simulates the observed action and, thus, he/she predicts its somatosensory consequences, then he/she should attenuate tactile stimuli simultaneously delivered to his/her corresponding body part. In three separate experiments, we found a systematic attenuation of touches during executed self-touch actions, but we found no evidence for attenuation when such actions were observed. Failure to observe somatosensory attenuation during observation of self-touch is not compatible with the hypothesis that the putative human mirror neuron system automatically predicts the sensory consequences of the observed action. In contrast, our findings emphasize a sharp distinction between the motor representations of self and others.

Direct Electrophysiological Correlates of Body Ownership in Human Cerebral Cortex.

Over the past decade, numerous neuroimaging studies based on hemodynamic markers of brain activity have examined the feeling of body ownership using perceptual body-illusions in humans. However, the direct electrophysiological correlates of body ownership at the cortical level remain unexplored. To address this, we studied the rubber hand illusion in 5 patients (3 males and 2 females) implanted with intracranial electrodes measuring cortical surface potentials. Increased high-γ (70-200 Hz) activity, an index of neuronal firing rate, in premotor and intraparietal cortices reflected the feeling of ownership. In both areas, high-γ increases were intimately coupled with the subjective illusion onset and sustained both during and in-between touches. However, intraparietal activity was modulated by tactile stimulation to a higher degree than the premotor cortex through effective connectivity with the hand-somatosensory cortex, which suggests different functional roles. These findings constitute the first intracranial electrophysiological characterization of the rubber hand illusion and extend our understanding of the dynamic mechanisms of body ownership.

Body ownership determines the attenuation of self-generated tactile sensations.

Self-perception depends on the brain's abilities to differentiate our body from the environment and to distinguish between the sensations generated as a consequence of voluntary movement and those arising from events in the external world. The first process refers to the sense of ownership of our body and relies on the dynamic integration of multisensory (afferent) signals. The second process depends on internal forward models that use (efferent) information from our motor commands to predict and attenuate the sensory consequences of our movements. However, the relationship between body ownership and sensory attenuation driven by the forward models remains unknown. To address this issue, we combined the rubber hand illusion, which allows experimental manipulation of body ownership, and the force-matching paradigm, which allows psychophysical quantification of somatosensory attenuation. We found that a rubber right hand pressing on the left index finger produced somatosensory attenuation but only when the model hand felt like one's own (illusory self-touch); reversely, the attenuation that was expected to occur during actual self-touch with the real hands was reduced when the participants simultaneously experienced ownership of a rubber right hand that was placed at a distance from their left hand. These results demonstrate that the sense of body ownership determines somatosensory attenuation. From a theoretical perspective, our results are important because they suggest that body ownership updates the internal representation of body state that provides the input to the forward model generating sensory predictions during voluntary action.

Ownership of an artificial limb induced by electrical brain stimulation.

Replacing the function of a missing or paralyzed limb with a prosthetic device that acts and feels like one's own limb is a major goal in applied neuroscience. Recent studies in nonhuman primates have shown that motor control and sensory feedback can be achieved by connecting sensors in a robotic arm to electrodes implanted in the brain. However, it remains unknown whether electrical brain stimulation can be used to create a sense of ownership of an artificial limb. In this study on two human subjects, we show that ownership of an artificial hand can be induced via the electrical stimulation of the hand section of the somatosensory (SI) cortex in synchrony with touches applied to a rubber hand. Importantly, the illusion was not elicited when the electrical stimulation was delivered asynchronously or to a portion of the SI cortex representing a body part other than the hand, suggesting that multisensory integration according to basic spatial and temporal congruence rules is the underlying mechanism of the illusion. These findings show that the brain is capable of integrating "natural" visual input and direct cortical-somatosensory stimulation to create the multisensory perception that an artificial limb belongs to one's own body. Thus, they serve as a proof of concept that electrical brain stimulation can be used to "bypass" the peripheral nervous system to induce multisensory illusions and ownership of artificial body parts, which has important implications for patients who lack peripheral sensory input due to spinal cord or nerve lesions.

Mental Imagery Induces Cross-Modal Sensory Plasticity and Changes Future Auditory Perception.

Can what we imagine in our minds change how we perceive the world in the future? A continuous process of multisensory integration and recalibration is responsible for maintaining a correspondence between the senses (e.g., vision, touch, audition) and, ultimately, a stable and coherent perception of our environment. This process depends on the plasticity of our sensory systems. The so-called ventriloquism aftereffect-a shift in the perceived localization of sounds presented alone after repeated exposure to spatially mismatched auditory and visual stimuli-is a clear example of this type of plasticity in the audiovisual domain. In a series of six studies with 24 participants each, we investigated an imagery-induced ventriloquism aftereffect in which imagining a visual stimulus elicits the same frequency-specific auditory aftereffect as actually seeing one. These results demonstrate that mental imagery can recalibrate the senses and induce the same cross-modal sensory plasticity as real sensory stimuli.

Recalibration of hand position sense during unconscious active and passive movement.

Precise knowledge of one's limbs' position in space is fundamental for goal-directed action. The brain's representation of the body in space is thought to be generated through a process of multisensory integration of visual, tactile and proprioceptive signals. In this study, we devised a setup that allowed us to displace participants' right hand without their subjective awareness. We accomplished this task by instructing the participants to view a live video feed of their hand from the first-person perspective. In the active condition, we used a sensorimotor illusion that caused the participants to actively but unknowingly displace their unseen right hand to a location 8 cm lateral to the image of their hand. In the passive condition, we mechanically displaced the participants' hand-at a slow, unnoticeable velocity-to the same location. We found that during active displacement, the participants indicated that the location of their hand was closer to the digital image of the hand rather than the veridical location of the hand, as compared with the passive condition, in which the participants indicated that the locations of their hand were closer to the actual location. These results indicated that, compared with passive displacement, active movements cause greater recalibration of the hand's spatial position and that the boosted spatial recalibration of hand position sense in the active task is driven by error-based sensorimotor corrections. These results have bearing on the perceptual mechanisms of recalibration of perceived limb location.

Dissociating the Neural Basis of Conceptual Self-Awareness from Perceptual Awareness and Unaware Self-Processing.

Conceptual self-awareness is a mental state in which the content of one's consciousness refers to a particular aspect of semantic knowledge about oneself. This form of consciousness plays a crucial role in shaping human behavior; however, little is known about its neural basis. Here, we use functional magnetic resonance imaging (fMRI) and a visual masked priming paradigm to dissociate the neural responses related to the awareness of semantic autobiographical information (one's own name, surname, etc.) from the awareness of information related to any visual stimulus (perceptual awareness), as well as from the unaware processing of self-relevant stimuli. To detect brain activity that is highly selective for self-relevant information, we used the blood-oxygen-level-dependent (BOLD) adaptation approach, which goes beyond the spatial limitations of conventional fMRI. We found that self-awareness was associated with BOLD adaptation in the medial frontopolar-retrosplenial areas, whereas perceptual awareness and unaware self-processing were associated with BOLD adaptation in the lateral fronto-parietal areas and the inferior temporal cortex, respectively. Thus, using a direct manipulation of conscious awareness we demonstrate for the first time that the neural basis of conceptual self-awareness is neuroanatomically distinct from the network mediating perceptual awareness of the sensory environment or unaware processing of self-related stimuli.

Illusory Obesity Triggers Body Dissatisfaction Responses in the Insula and Anterior Cingulate Cortex.

In today's Western society, concerns regarding body size and negative feelings toward one's body are all too common. However, little is known about the neural mechanisms underlying negative feelings toward the body and how they relate to body perception and eating-disorder pathology. Here, we used multisensory illusions to elicit illusory ownership of obese and slim bodies during functional magnetic resonance imaging. The results implicate the anterior insula and the anterior cingulate cortex in the development of negative feelings toward the body through functional interactions with the posterior parietal cortex, which mediates perceived obesity. Moreover, cingulate neural responses were modulated by nonclinical eating-disorder psychopathology and were attenuated in females. These results reveal how perceptual and affective body representations interact in the human brain and may help explain the neurobiological underpinnings of eating-disorder vulnerability in women.

Out-of-body-induced hippocampal amnesia.

Theoretical models have suggested an association between the ongoing experience of the world from the perspective of one's own body and hippocampus-based episodic memory. This link has been supported by clinical reports of long-term episodic memory impairments in psychiatric conditions with dissociative symptoms, in which individuals feel detached from themselves as if having an out-of-body experience. Here, we introduce an experimental approach to examine the necessary role of perceiving the world from the perspective of one's own body for the successful episodic encoding of real-life events. While participants were involved in a social interaction, an out-of-body illusion was elicited, in which the sense of bodily self was displaced from the real body to the other end of the testing room. This condition was compared with a well-matched in-body illusion condition, in which the sense of bodily self was colocalized with the real body. In separate recall sessions, performed ∼1 wk later, we assessed the participants' episodic memory of these events. The results revealed an episodic recollection deficit for events encoded out-of-body compared with in-body. Functional magnetic resonance imaging indicated that this impairment was specifically associated with activity changes in the posterior hippocampus. Collectively, these findings show that efficient hippocampus-based episodic-memory encoding requires a first-person perspective of the natural spatial relationship between the body and the world. Our observations have important implications for theoretical models of episodic memory, neurocognitive models of self, embodied cognition, and clinical research into memory deficits in psychiatric disorders.

The fusion of mental imagery and sensation in the temporal association cortex.

It is well understood that the brain integrates information that is provided to our different senses to generate a coherent multisensory percept of the world around us (Stein and Stanford, 2008), but how does the brain handle concurrent sensory information from our mind and the external world? Recent behavioral experiments have found that mental imagery--the internal representation of sensory stimuli in one's mind--can also lead to integrated multisensory perception (Berger and Ehrsson, 2013); however, the neural mechanisms of this process have not yet been explored. Here, using functional magnetic resonance imaging and an adapted version of a well known multisensory illusion (i.e., the ventriloquist illusion; Howard and Templeton, 1966), we investigated the neural basis of mental imagery-induced multisensory perception in humans. We found that simultaneous visual mental imagery and auditory stimulation led to an illusory translocation of auditory stimuli and was associated with increased activity in the left superior temporal sulcus (L. STS), a key site for the integration of real audiovisual stimuli (Beauchamp et al., 2004a, 2010; Driver and Noesselt, 2008; Ghazanfar et al., 2008; Dahl et al., 2009). This imagery-induced ventriloquist illusion was also associated with increased effective connectivity between the L. STS and the auditory cortex. These findings suggest an important role of the temporal association cortex in integrating imagined visual stimuli with real auditory stimuli, and further suggest that connectivity between the STS and auditory cortex plays a modulatory role in spatially localizing auditory stimuli in the presence of imagined visual stimuli.