A spontaneous gravity prior: newborn chicks prefer stimuli that move against gravity.

At the beginning of life, inexperienced animals use evolutionary-given preferences (predispositions) to decide what stimuli to attend and approach. Stimuli that contain cues of animacy, such as face-like stimuli, biological motion and changes in speed, are particularly attractive across vertebrate taxa. A strong cue of animacy is upward movement against terrestrial gravity, because only animate objects consistently move upward. To test whether upward movement is spontaneously considered attractive already at birth, we tested the early preferences of dark-hatched chicks (Gallus gallus) for upward- versus downward-moving visual stimuli. We found that, without any previous visual experience, chicks consistently exhibited a preference to approach stimuli that move upward, against gravity. A control experiment showed that these preferences are not driven by avoidance of downward stimuli. These results show that newborn animals have a gravity prior that attracts them toward upward movement. Movement against gravity can be used as a cue of animacy to orient early approach responses in the absence of previous visual experience.

Cognition in zero gravity: Effects of non-terrestrial gravity on human behaviour.

As humanity prepares for deep space exploration, understanding the impact of spaceflight on bodily physiology is critical. While the effects of non-terrestrial gravity on the body are well established, little is known about its impact on human behaviour and cognition. Astronauts often describe dramatic alterations in sensorimotor functioning, including orientation, postural control, and balance. Changes in cognitive functioning as well as in socio-affective processing have also been observed. Strikingly, no comprehensive theoretical model exists to outline the impact of non-terrestrial gravity on behaviour. Here, we have reviewed the key literature across the last 10 years and explored the impact of non-terrestrial gravity across three key functional domains: sensorimotor functioning, cognition, and socio-affective processing. We have proposed and preliminary validated a neurocognitive model to account for the effects of non-terrestrial gravity in these domains. Understanding the impact of non-terrestrial gravity on human behaviour has never been timelier and it will help mitigate against risks in both commercial and non-commercial spaceflight.

Human movements do not look the same in a tilted world: Gravitational constraints influence the perception of biological motion.

We investigated whether gravitational constraints influence the interaction of visual, proprioceptive and vestibular cues for Biological Motion Perception (BMP). Participants were asked to distinguish between plausible and random point-light movements, while passively placed in either an upright or a tilted body orientation. Manipulating the body orientation with respect to gravity leads to different gravitational signals transmitted by the visual, proprioceptive, and vestibular systems. Participants were overall faster in distinguishing plausible point-light movements than random movements. Critically, response times for biologically plausible point-light movements - but not for random movements - were significantly prolonged in the tilted body orientation. Our results suggest that BMP depends not only on the spatial-temporal cues embedded in point-light movements but also rely on the congruency between current gravitational signals detected by the sensory systems and our previous knowledge of terrestrial gravity. STATEMENT OF RELEVANCE: As humankind is preparing for a new space age, understanding how gravity influences behaviour and cognition has never been more pressing. All living organisms have evolved to survive in a terrestrial gravitational field. Although we cannot consciously feel gravity, it has an impact in our life: it affects how we move and interact with the external environment. The sensory signals from the vestibular system are continuously combined with visual and proprioceptive cues to help us in maintaining a stable representation of the world. Here we placed participants in a tilted body orientation and were able to determine that a conflict between prior gravitational knowledge and what was actively sensed about gravity affected human Biological Movement Perception. Humans suffer changes in perception under non-terrestrial gravity conditions that may potentially compromise performance during space exploration.

Galvanic Vestibular Stimulation influences risk-taking behaviour.

Risk-taking behaviour is an essential aspect of our interactions with the environment. Here we investigated whether vestibular inputs influence behavioural measurement of risk-taking propensity. We have combined bipolar Galvanic Vestibular Stimulation (GVS) with a well-known and established risk-taking behaviour task, namely the Balloon Analogue Risk Task (BART). A sham stimulation was used to control for non-specific effects. Left-anodal and right-cathodal GVS (L-GVS), which preferentially activates the vestibular projections in the right hemisphere, decreased the willingness to take risk during the BART compared with right-anodal and left-cathodal GVS (R-GVS), which activates the left hemisphere. This proved a specific vestibular effect which depends on GVS polarity. Conversely, no generic vestibular effect, defined as the adjusted average of L-GVS and R-GVS conditions compared to sham, emerged, excluding non-specific vestibular effects. Our results confirmed recent findings of a vestibular contribution to decision-making and strategy control behaviour. We suggest that the vestibular-mediated balancing of risk seeking behaviour is an important element of the brain's capacity to adapt to the environment.

Where is my hand in space? The internal model of gravity influences proprioception.

Knowing where our limbs are in space is crucial for a successful interaction with the external world. Joint position sense (JPS) relies on both cues from muscle spindles and joint mechanoreceptors, as well as the effort required to move. However, JPS may also rely on the perceived external force on the limb, such as the gravitational field. It is well known that the internal model of gravity plays a large role in perception and behaviour. Thus, we have explored whether direct vestibular-gravitational cues could influence JPS. Participants passively estimated the position of their hand while they were upright and therefore aligned with terrestrial gravity, or pitch-tilted 45° backwards from gravity. Overall participants overestimated the position of their hand in both upright and tilted postures; however, the proprioceptive bias was significantly reduced when participants were tilted. Our findings therefore suggest that the internal model of gravity may influence and update JPS in order to allow the organism to interact with the environment.

Which way is down? Visual and tactile verticality perception in expert dancers and non-experts.

Gravity provides an absolute verticality reference for all spatial perception, allowing us to move within and interact effectively with our world. Bayesian inference models explain verticality perception as a combination of online sensory cues with a prior prediction that the head is usually upright. Until now, these Bayesian models have been formulated for judgements of the perceived orientation of visual stimuli. Here, we investigated whether judgements of the verticality of tactile stimuli follow a similar pattern of Bayesian perceptual inference. We also explored whether verticality perception is affected by the postural and balance expertise of dancers. We tested both the subjective visual vertical (SVV) and the subjective tactile vertical (STV) in ballet dancers and non-dancers. A robotic arm traced downward-moving visual or tactile stimuli in separate blocks while participants held their head either upright or tilted 30° to their right. Participants reported whether these stimuli deviated to the left (clockwise) or right (anti-clockwise) of the gravitational vertical. Tilting the head biased the SVV away from the longitudinal head axis (the classical E-effect), consistent with a failure to compensate for the vestibulo-ocular counter-roll reflex. On the contrary, tilting the head biased the STV toward the longitudinal head axis (the classical A-effect), consistent with a strong upright head prior. Critically, tilting the head reduced the precision of verticality perception, particularly for ballet dancers' STV judgements. Head tilt is thought to increase vestibular noise, so ballet dancers seem to be surprisingly susceptible to degradation of vestibular inputs, giving them an inappropriately high weighting in verticality judgements.

Gravity prior in human behaviour: a perceptual or semantic phenomenon?

Humans show a gravitational advantage in perception: we are more precise at judging the speed of downwards-moving than upwards-moving objects, indicating that gravitational acceleration is an internalised prior. However, it is unclear whether this gravity prior is based on purely perceptual cues or whether it can incorporate semantic knowledge. Previous research has used only objects which are known to comply with gravity, possibly confounding semantic and perceptual cues. Here we have addressed this question by asking participants to judge the speed of objects that typically move coherently with gravity (ball) or against it (rocket). Our results showed a perceptual advantage for falling stimuli, irrespective of object identity, suggesting the gravity prior is based on perceptual cues.

Vestibular cognition: State-of-the-art and future directions.

Vestibular information has been traditionally considered as a specialized input for basic orienting behaviours, such as oculo-motor adjustments, postural control and gaze orientation. However, in the past two decades a widespread vestibular network in the human brain has been identified, that goes far beyond the low-level reflex circuits emphasized by earlier work. Because this vestibular cortical network is so widely distributed, it could, in principle, impact multiple neurocognitive functions in health and disease. This paper focuses on the relations between vestibular input, vestibular networks, and vestibular interventions by providing the authors' personal viewpoint on the state-of-the-art of vestibular cognitive neuropsychology, and its potential relevance for neurorehabilitation.

Multisensory Interactions in Virtual Reality: Optic Flow Reduces Vestibular Sensitivity, but Only for Congruent Planes of Motion.

During exposure to Virtual Reality (VR) a sensory conflict may be present, whereby the visual system signals that the user is moving in a certain direction with a certain acceleration, while the vestibular system signals that the user is stationary. In order to reduce this conflict, the brain may down-weight vestibular signals, which may in turn affect vestibular contributions to self-motion perception. Here we investigated whether vestibular perceptual sensitivity is affected by VR exposure. Participants' ability to detect artificial vestibular inputs was measured during optic flow or random motion stimuli on a VR head-mounted display. Sensitivity to vestibular signals was significantly reduced when optic flow stimuli were presented, but importantly this was only the case when both visual and vestibular cues conveyed information on the same plane of self-motion. Our results suggest that the brain dynamically adjusts the weight given to incoming sensory cues for self-motion in VR; however this is dependent on the congruency of visual and vestibular cues.

Vection in virtual reality modulates vestibular-evoked myogenic potentials.

The popularity of virtual reality (VR) has increased rapidly in recent years. While significant technological advancements are apparent, a troublesome problem with VR is that between 20% and 80% of users will experience unpleasant side effects such as nausea, disorientation, blurred vision and headaches-a malady known as Cybersickness. Cybersickness may be caused by a conflict between sensory signals for self-motion: while vision signals that the user is moving in a certain direction with certain acceleration, the vestibular organs provide no corroborating information. To resolve the sensory conflict, vestibular cues may be down-weighted leading to an alteration of how the brain interprets actual vestibular information. This may account for the frequently reported after-effects of VR exposure. Here, we investigated whether exposure to vection in VR modulates vestibular processing. We measured vestibular-evoked myogenic potentials (VEMPs) during brief immersion in a vection-inducing VR environment presented via head-mounted display. We found changes in VEMP asymmetry ratio, with a substantial increase in VEMP amplitude recorded on the left sternocleidomastoid muscle following just one minute of exposure to vection in VR. Our results suggest that exposure to vection in VR modulates vestibular processing, which may explain common after-effects of VR.

Why the whole is more than the sum of its parts: Salience-driven overestimation in aggregated tactile sensations.

Experimental psychology often studies perception analytically, reducing its focus to minimal sensory units, such as thresholds or just noticeable differences in a single stimulus. Here, in contrast, we examine a synthetic aspect: how multiple inputs to a sensory system are aggregated into an overall percept. Participants in three experiments judged the total stimulus intensity for simultaneous electrical shocks to two digits. We tested whether the integration of component somatosensory stimuli into a total percept occurs automatically, or rather depends on the ability to consciously perceive discrepancy among components (Experiment 1), whether the discrepancy among these components influences sensitivity or/and perceptual bias in judging totals (Experiment 2), and whether the salience of each individual component stimulus affects perception of total intensity (Experiment 3). Perceptual aggregation of two simultaneous component events occurred both when participants could perceptually discriminate the two intensities, and also when they could not. Further, the actual discrepancy between the stimuli modulated both participants' sensitivity and perceptual bias: increasing discrepancies produced a systematic and progressive overestimation of total intensity. The degree of this bias depended primarily on the salience of the stronger stimulus in the pair. Overall, our results suggest that important nonlinear mechanisms contribute to sensory aggregation. The mind aggregates component inputs into a coherent and synthetic perceptual experience in a salience-weighted fashion that is not based on simple summation of inputs.

Getting ready for Mars: How the brain perceives new simulated gravitational environments.

On Earth, we are continually exposed to gravity: sensory signals are constantly integrated to form an internal model of gravity. However, it is unclear whether this internal model is fixed to Earth's gravity or whether it can be applied to a new gravitational environment. Under terrestrial gravity, observers show a "gravitational bias" while judging the speed of falling versus rising objects, as they comply with the physical laws of gravity. We investigated whether this gravitational bias may be present when judging the speed of objects moving upwards or downwards in both virtual reality (VR)-simulated Earth gravity (9.81 m/s2) and Mars gravity (3.71 m/s2). Our results highlighted a gravitational bias in both Earth and Mars VR-simulated gravity: the speed of downwards movement was more precisely detected than the speed of upwards movement. Although the internal model of gravity has been built up under terrestrial gravity, it can quickly expand to novel non-terrestrial gravitational environments.

Thermonociceptive interaction: interchannel pain modulation occurs before intrachannel convergence of warmth.

Nonnoxious warmth reduces both perceived pain intensity and the amplitude of EEG markers of pain. However, the spatial properties of thermonociceptive interaction, and the level of sensory processing at which it occurs, remain unclear. We investigated whether interchannel warmth-pain interactions occur before or after intrachannel spatial summation of warmth. Warm stimuli were applied to the fingers of the right hand. Their number and location were manipulated in different conditions. A concomitant noxious test pulse was delivered to the middle finger using a CO2 laser. We replicated the classical suppressive effect of warmth on both perceived pain intensity and EEG markers. Importantly, inhibition of pain was not affected by the location and the number of thermal stimuli, even though they increased the perceived intensity of warmth. Our results therefore suggest that the inhibitory effect of warmth on pain is not somatotopically organized. The results also rule out the possibility that warmth affects nociceptive processing after intrachannel warmth summation. NEW & NOTEWORTHY We used spatial summation of warmth as a model to investigate thermonociceptive interactions. Painful CO2 laser pulses were delivered during different thermal conditions. We found that warmth inhibited pain regardless of its location. Crucially, spatial summation of multiple warm stimuli did not further inhibit pain. These findings suggest that warmth-pain interaction occurs independently of or after spatial summation of warmth.

Gravity modulates behaviour control strategy.

Human behaviour is a trade-off between exploitation of familiar resources and exploration of new ones. In a challenging environment-such as outer space-making the correct decision is vital. On Earth, gravity is always there, and is an important reference for behaviour. Thus, altered gravitational signals may affect behaviour control strategies. Here, we investigated whether changing the body's orientation to the gravitational vector would modulate the balance between routine and novel behaviour. Participants completed a random number generation task while upright or supine. We found decreased randomness when participants were supine. In particular, the degree of equiprobability of pairs of consecutive responses was reduced in the supine orientation. Online gravitational signals may shape the balance between exploitation and exploration, in favour of more stereotyped and routine responses.

The aesthetics of verticality: A gravitational contribution to aesthetic preference.

Verticality plays a fundamental role in the arts, portraying concepts such as power, grandeur, or even morality; however, it is unclear whether people have an aesthetic preference for vertical stimuli. The perception of verticality occurs by integrating vestibular-gravitational input with proprioceptive signals about body posture. Thus, these signals may influence the preference for verticality. Here, we show that people have a genuine aesthetic preference for stimuli aligned with the vertical, and this preference depends on the position of the body relative to the gravitational direction. Observers rated the attractiveness of lines that varied in inclination. Perfectly vertical lines were judged to be more attractive than those inclined clockwise or anticlockwise only when participants held an upright posture. Critically, this preference was not present when their body was tilted away from the gravitational vertical. Our results showed that gravitational signals make a contribution to the perception of attractiveness of environmental objects.

Somatosensory modulation of perceptual vestibular detection.

Vestibular-multisensory interactions are essential for self-motion, navigation and postural stability. Despite evidence suggesting shared brain areas between vestibular and somatosensory inputs, no study has yet investigated whether somatosensory information influences vestibular perception. Here, we used signal detection methods to identify whether somatosensory stimulation might interact with vestibular events in a vestibular detection task. Participants were instructed to detect near-threshold vestibular roll-rotation sensations delivered by galvanic vestibular stimulation in one-half of experimental trials. A vibrotactile signal occurred to the index fingers of both hands in half of the trials, independent of vestibular signals. We found that vibrotactile somatosensory stimulation decreased perceptual vestibular sensitivity. The results are compatible with a gain regulation mechanism between vestibular and somatosensory modalities.

Cybersickness: a Multisensory Integration Perspective.

In the past decade, there has been a rapid advance in Virtual Reality (VR) technology. Key to the user's VR experience are multimodal interactions involving all senses. The human brain must integrate real-time vision, hearing, vestibular and proprioceptive inputs to produce the compelling and captivating feeling of immersion in a VR environment. A serious problem with VR is that users may develop symptoms similar to motion sickness, a malady called cybersickness. At present the underlying cause of cybersickness is not yet fully understood. Cybersickness may be due to a discrepancy between the sensory signals which provide information about the body's orientation and motion: in many VR applications, optic flow elicits an illusory sensation of motion which tells users that they are moving in a certain direction with certain acceleration. However, since users are not actually moving, their proprioceptive and vestibular organs provide no cues of self-motion. These conflicting signals may lead to sensory discrepancies and eventually cybersickness. Here we review the current literature to develop a conceptual scheme for understanding the neural mechanisms of cybersickness. We discuss an approach to cybersickness based on sensory cue integration, focusing on the dynamic re-weighting of visual and vestibular signals for self-motion.

Up, Down, Near, Far: An Online Vestibular Contribution to Distance Judgement.

Whether a visual stimulus seems near or far away depends partly on its vertical elevation. Contrasting theories suggest either that perception of distance could vary with elevation, because of memory of previous upwards efforts in climbing to overcome gravity, or because of fear of falling associated with the downwards direction. The vestibular system provides a fundamental signal for the downward direction of gravity, but the relation between this signal and depth perception remains unexplored. Here we report an experiment on vestibular contributions to depth perception, using Virtual Reality. We asked participants to judge the absolute distance of an object presented on a plane at different elevations during brief artificial vestibular inputs. Relative to distance estimates collected with the object at the level of horizon, participants tended to overestimate distances when the object was presented above the level of horizon and the head was tilted upward and underestimate them when the object was presented below the level of horizon. Interestingly, adding artificial vestibular inputs strengthened these distance biases, showing that online multisensory signals, and not only stored information, contribute to such distance illusions. Our results support the gravity theory of depth perception, and show that vestibular signals make an on-line contribution to the perception of effort, and thus of distance.

Thermal referral: evidence for a thermoceptive uniformity illusion without touch.

When warm thermal stimulators are placed on the ring and index fingers of one hand, and a neutral-temperature stimulator on the middle finger, all three fingers feel warm. This illusion is known as thermal referral (TR). On one interpretation, the heterogenous thermal signals are overridden by homogenous tactile signals. This cross-modal thermo-tactile interaction could reflect a process of object recognition, based on the prior that many objects are thermally homogenous. Interestingly, the illusion was reported to disappear when the middle digit was lifted off the thermal stimulator, suggesting that tactile stimulation is necessary. However, no study has investigated whether purely thermal stimulation might induce TR, without any tactile object to which temperature can be attributed. We used radiant thermal stimulation to deliver purely thermal stimuli, which either were or were not accompanied by simultaneous touch. We found identical TR effects in both the original thermo-tactile condition, and in a purely thermoceptive condition where no tactile object was present. Control experiments ruled out explanations based on poor spatial discrimination of warm signals. Our purely thermoceptive results suggest that TR could reflect low-level organization of the thermoceptive pathway, rather than a cognitive intermodal modulation based on tactile object perception.