No abstract

The rubber hand illusion (RHI) is an enigmatic illusion that creates a feeling of owning an artificial limb. Enthusiasts of this paradigm assert that it operationalizes bodily self-awareness, but there are reasons to doubt such a clear link. Because little is known about other functional contributions to the RHI, including effects of context-dependent visual processing and cognitive control or the ability to resolve intermodal conflict, we carried out two complementary experiments. In the first, we examined the relationships between the RHI and (1) body awareness, as assessed by the Body Perception Questionnaire (BPQ); (2) context-dependent visual processing, as assessed by the rod-and-frame test (RFT); and (3) conflict resolution, as assessed by the Stroop test. We found a significant positive correlation between the RHI-associated proprioceptive drift and context-dependent visual processing on the RFT, but not between the RHI and body awareness on the BPQ. In the second experiment, we examined the RHI in advanced yoga practitioners with an embodied lifestyle and a heightened sense of their own body in space. They succumbed to the illusion just as much as did yoga-naïve control participants, despite significantly greater body awareness on the BPQ. These findings suggest that susceptibility to the RHI and awareness of one's own body are at least partially independent processes.

The primary motor cortex (M1) is highly influenced by premotor/motor areas both within and across hemispheres. Dual site transcranial magnetic stimulation (dsTMS) has revealed interhemispheric interactions mainly at early latencies. Here, we used dsTMS to systematically investigate long-latency causal interactions between right-hemisphere motor areas and the left M1 (lM1). We stimulated lM1 using a suprathreshold test stimulus (TS) to elicit motor-evoked potentials (MEPs) in the right hand. Either a suprathreshold or a subthreshold conditioning stimulus (CS) was applied over the right M1 (rM1), the right ventral premotor cortex (rPMv), the right dorsal premotor cortex (rPMd) or the supplementary motor area (SMA) prior to the TS at various CS-TS inter-stimulus intervals (ISIs: 40–150 ms). The CS strongly affected lM1 excitability depending on ISI, CS site and intensity. Inhibitory effects were observed independently of CS intensity when conditioning PMv, rM1 and SMA at a 40-ms ISI, with larger effects after PMv conditioning. Inhibition was observed with suprathreshold PMv and rM1 conditioning at a 150-ms ISI, while site-specific, intensity-dependent facilitation was detected at an 80-ms ISI. Thus, long-latency interhemispheric interactions, likely reflecting indirect cortico-cortical/cortico-subcortical pathways, cannot be reduced to nonspecific activation across motor structures. Instead, they reflect intensity-dependent, connection- and time-specific mechanisms.

Perception of the visual vertical is strongly based on our ability to match visual inflow with vestibular, proprioceptive, tactile, and even visceral information that contributes to maintaining an internal representation of the vertical. An important cortical region implicated in multisensory integration is the right temporoparietal junction (rTPJ), which also is involved in higher order forms of body- and space-related cognition. To test whether this region integrates body-related multisensory information necessary for establishing the subjective visual vertical, we combined a psychophysical task (the rod-and-frame test) with transient inhibition of the rTPJ via continuous theta burst stimulation (cTBS). A Gabor patch visual detection task was used as a control visual task. cTBS of early visual cortex (V1-V3) was used to test whether early visual cortices played any role in verticality estimation. We show that inhibition of rTPJ activity selectively impairs the ability to evaluate the rod's verticality when no contextual visual information, such as a frame surrounding the rod, is provided. Conversely, transient inhibition of V1-V3 selectively disrupts the ability to visually detect Gabor patch orientation. This anatomofunctional dissociation supports the idea that the rTPJ plays a causal role in integrating egocentric sensory information encoded in different reference systems (i.e., vestibular and somatic) to maintain an internal representation of verticality.

The primary motor cortex (M1) is strongly influenced by several frontal regions. Dual-site transcranial magnetic stimulation (dsTMS) has highlighted the timing of early (<40 ms) prefrontal/premotor influences over M1. Here we used dsTMS to investigate, for the first time, longer-latency causal interactions of the posterior inferior frontal gyrus (pIFG) and pre-supplementary motor area (pre-SMA) with M1 at rest. A suprathreshold test stimulus (TS) was applied over M1 producing a motor-evoked potential (MEP) in the relaxed hand. Either a subthreshold or a suprathreshold conditioning stimulus (CS) was administered over ipsilateral pIFG/pre-SMA sites before the TS at different CS-TS inter-stimulus intervals (ISIs: 40–150 ms). Independently of intensity, CS over pIFG and pre-SMA (but not over a control site) inhibited MEPs at an ISI of 40 ms. The CS over pIFG produced a second peak of inhibition at an ISI of 150 ms. Additionally, facilitatory modulations were found at an ISI of 60 ms, with supra- but not subthreshold CS intensities. These findings suggest differential modulatory roles of pIFG and pre-SMA in M1 excitability. In particular, the pIFG –but not the pre-SMA– exerts intensity-dependent modulatory influences over M1 within the explored time window of 40-150 ms, evidencing fine-tuned control of M1 output.

In the rod and frame test (RFT), participants are asked to set a tilted visual linear marker (i.e., a rod), embedded in a square, to the subjective vertical, irrespective of the surrounding frame. People not influenced by the frame tilt are defined as field-independent, while people biased in their rod verticality perception are field-dependent. Performing RFT requires the integration of proprioceptive, vestibular and visual signals with the latter accounting for field-dependency. Studies indicate that motor experts in body-related, balance-improving disciplines tend to be field-independent, i.e., better at verticality perception, suggesting that proprioceptive and vestibular expertise acquired by such exercise may weaken the influence of irrelevant visual signals. What remains unknown is whether the effect of body-related expertise in weighting perceptual information might also be mediated by personality traits, in particular those indexing self-focusing abilities. To explore this issue, we tested field-dependency in a class of body experts, namely yoga practitioners and in non-expert participants. Moreover we explored any link between performance on RFT and self-transcendence (ST), a complex personality construct, which refers to tendency to experience spiritual feelings and ideas. As expected, yoga practitioners (i) were more accurate in assessing the rod's verticality on the RFT, and (ii) expressed significantly higher ST. Interestingly, the performance in these two tests was negatively correlated. More specifically, when asked to provide verticality judgments, highly self-transcendent yoga practitioners were significantly less influenced by a misleading visual context. Our results suggest that being highly self-transcendent may enable yoga practitioners to optimize verticality judgment tasks by relying more on internal (vestibular and proprioceptive) signals coming from their own body, rather than on exteroceptive, visual cues.

Cortico-cortical paired associative stimulation (ccPAS) is an effective transcranial magnetic stimulation (TMS) method for inducing associative plasticity between interconnected brain areas in humans. Prior ccPAS studies have focused on protocol’s aftereffects. Here, we investigated physiological changes induced “online” during ccPAS administration. We tested 109 participants receiving ccPAS over left ventral premotor cortex (PMv) and primary motor cortex (M1) using a standard procedure (90 paired-pulses with 8-ms interstimulus interval, repeated at 0.1 Hz frequency). On each paired-pulse, we recorded a motor-evoked potential (MEP) to continuously trace the emergence of corticomotor changes. Participant receiving forward-ccPAS (on each pair, a first TMS pulse was administered over PMv, second over M1, i.e., PMv-to-M1) showed a gradual and linear increase in MEP size that did not reach a plateau at the end of the protocol and was greater in participants with low motor threshold. Participants receiving reverse-ccPAS (i.e., M1-to-PMv) showed a trend toward inhibition. Our study highlights the facilitatory and inhibitory modulations that occur during ccPAS administration and suggest that online MEP monitoring could provide insights into the malleability of the motor system and protocol’s effectiveness. Our findings open interesting prospects about ccPAS potential optimization in experimental and clinical settings.

These results might compellingly suggest that, despite high compassion, heightened bodily self-awareness might increase a self-centred perspective and limit altruistic acts in advanced yoga practitioners.