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.

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.

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.

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.

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.

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.

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.

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.

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.

The vestibular body: Vestibular contributions to bodily representations.

Vestibular signals are integrated with signals from other sensory modalities. This convergence could reflect an important mechanism for maintaining the perception of the body. Here we review the current literature in order to develop a framework for understanding how the vestibular system contributes to body representation. According to recent models, we distinguish between three processes for body representation, and we look at whether vestibular signals might influence each process. These are (i) somatosensation, the primary sensory processing of somatic stimuli, (ii) somatoperception, the processes of constructing percepts and experiences of somatic objects and events and (iii) somatorepresentation, the knowledge about the body as a physical object in the world. Vestibular signals appear to contribute to all three levels in this model of body processing. Thus, the traditional view of the vestibular system as a low-level, dedicated orienting module tends to underestimate the pervasive role of vestibular input in bodily self-awareness.

Caloric vestibular stimulation modulates nociceptive evoked potentials.

Vestibular stimulation has been reported to alleviate central pain. Clinical and physiological studies confirm pervasive interactions between vestibular signals and somatosensory circuits, including nociception. However, the neural mechanisms underlying vestibular-induced analgesia remain unclear, and previous clinical studies cannot rule out explanations based on alternative, non-specific effects such as distraction or placebo. To investigate how vestibular inputs influence nociception, we combined caloric vestibular stimulation (CVS) with psychophysical and electrocortical responses elicited by nociceptive-specific laser stimulation in humans (laser-evoked potentials, LEPs). Cold water CVS applied to the left ear resulted in significantly lower subjective pain intensity for experimental laser pain to the left hand immediately after CVS, relative both to before CVS and to 1 h after CVS. This transient reduction in pain perception was associated with reduced amplitude of all LEP components, including the early N1 wave reflecting the first arrival of nociceptive input to primary somatosensory cortex. We conclude that cold left ear CVS elicits a modulation of both nociceptive processing and pain perception. The analgesic effect induced by CVS could be mediated either by subcortical gating of the ascending nociceptive input, or by direct modulation of the primary somatosensory cortex.

Combining proprioception and touch to compute spatial information.

Localising a tactile stimulus in egocentric space involves integrating information from skin receptors with proprioceptive inputs about body posture. We investigated whether body posture automatically influences tactile spatial judgements, even when it is irrelevant to the task. In Experiment 1, participants received two successive tactile stimuli on the forearm and were asked to indicate whether the first or second touch of the pair was closer to an anatomical body landmark, either the wrist or the elbow. The task was administered in three experimental conditions involving different body postures: canonical body posture with extended forearm and hand pointing distally; a non-canonical body posture with forearm and hand pointing vertically up at 90° and a 'reversed' body posture with the elbow fully flexed at 180°, so that the hand pointed proximally. Thus, our task required localising touch on the skin and then relating skin locations to anatomical body landmarks. Critically, both functions are independent of the posture of the body in space. We nevertheless found reliable effects of body posture: judgement errors increased when the canonical forearm posture was rotated through 180°. These results were further confirmed in Experiment 2, in which stimuli were delivered to the finger. However, additionally reversing the canonical posture of the finger, as well as that of the forearm, so that the finger was restored to its canonical orientation in egocentric space, restored performance to normal levels. Our results confirm an automatic process of localising the body in external space underlying the process of tactile perception. This process appears to involve a combination of proprioceptive and tactile information.

Galvanic vestibular stimulation influences randomness of number generation.

Successful interaction with the external environment requires a balance between novel or exploratory and routine or exploitative behaviours. This distinction is often expressed in terms of location or orientation of the body relative to surrounding space: functions in which the vestibular system plays an important role. However, the distinction can also be applied to novel versus repetitive production of any behaviour or symbol. Here, we investigated whether vestibular inputs contribute to the balance between novel and routine behaviours, independently of their effects on spatial orienting, by assessing effects of galvanic vestibular stimulation (GVS) on a random number generation task. Right-anodal/left-cathodal GVS, which preferentially activates the left cerebral hemisphere decreased the randomness of the sequence, while left-anodal/right-cathodal GVS, which preferentially activates the right hemisphere increased it. GVS did not induce any spatial biases in locations chosen from the number line. Our results suggest that vestibular stimulation of each hemisphere has a specific effect on the balance between novel and routine actions. We found no evidence for effects of non-specific arousal due to GVS on random number generation, and no evidence for effects on number generation consistent with modulation of spatial attention due to GVS.

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.

How the vestibular system modulates tactile perception in normal subjects: a behavioural and physiological study.

Caloric vestibular stimulation (CVS) is a physiological technique demonstrated to transiently improve hemianaesthesia in right brain-damaged patients (Bottini et al. in Exp Brain Res 99(1):164-169, 1994, Nature 376:778-781, 1995, Neurology 65(8):1278-1283, 2005). Recent studies suggest that these effects are based on the anatomical overlapping between vestibular and tactile projections (Bottini et al. in Nature 376:778-781, 1995) in the human brain. However, much less is known about behavioural effects of this manipulation on normal subjects. We aimed to explore tactile perception during left ear CVS in normal subjects. We administered seventeen right-handed normal subjects with different types of tactile stimuli (above and below threshold) during left ear CVS. To further ensure standardized procedure, tactile stimulation was delivered through a tool-developed ad hoc for the experiment. The experiment was divided in 3 conditions: (1) Baseline, (2) PostCVS and (3) Delayed CVS. We found a main effect of stimulus type (F ((2,32)) = 907.712; P = 0.000) and condition (F ((2,32)) = 55.505; P = 0.000). Moreover, post hoc comparisons revealed that below threshold stimuli are most affected by CVS (t ((16)) = -11.213; P = 0.000). Left ear CVS modulates tactile perception also in normal subjects. Moreover, this modulation seems to be selective for below threshold stimuli and not caused by attentive processes. A multisensory phenomenon is possibly the best explanation for this interaction between touch and vestibular systems, corroborated also by the anatomical evidence and by the previous knowledge about interaction with the environment.

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.

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.

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.

Productive symptoms in right brain damage.

PURPOSE OF REVIEW: The purpose of this review is to summarize the more recent studies on productive symptoms from the neuropsychological, neurophysiological and anatomical points of view. The integration of these aspects may provide some clarifications on the cognitive impairments underpinning the main productive disorders, also contributing to better understand the normal functioning of the brain. RECENT FINDINGS: Productive symptoms are closely associated to spatial neglect and are distinguished in relation to the part of space they manifest. The investigation of perseveration in extrapersonal space with different manipulations helps to understand the neuropathological mechanisms underlying this symptom. Anosognosia for hemiplegia and somatoparaphrenia may be considered as disorders of body representation (personal space). Recently it has been proposed that these disorders may be ascribed to an impairment of different levels of motor control. The identification of the anatomical correlates of these two disorders contributes to better understanding of their the cognitive nature. SUMMARY: Productive behaviours have diverse clinical manifestations and may be induced by different mechanisms. Lesional studies are beginning to provide evidence for specific anatomical correlates of these disorders. Further investigations are needed to better understand to what extent productive symptoms can be disentagled from spatial neglect. These attempts may contribute to clarifying the role of the right hemisphere in monitoring spatial cognition.