Changes in perceived peripersonal space following the rubber hand illusion.

Peripersonal space (PPS), the region immediately surrounding the body is essential for bodily protection and goal directed action. Previous studies have suggested that the PPS is anchored to one's own body and in the current study we investigated whether the PPS could be modulated by changes in perceived body ownership. While theoretically important, this anchoring can also have implications for patients with altered body perception. The rubber hand illusion (RHI) is a way to manipulate body ownership. We hypothesized that after induction of a left hand RHI, the perceived space around the body shifts to the right. Sixty-five participants performed a landmark task before and after a left hand RHI. In the landmark task, participants had to determine whether a vertical landmark line was left or right from the center of a horizontal screen. One group of the participants was exposed to synchronous stroking, the other group experienced asynchronous stroking. Results showed a shift in space to the right (e.g. away from the own arm), but only for the 'synchronous stroking' group. These results suggest that the relevant action space becomes linked to the fake hand. Critically, subjective ownership experience did not correlate with this shift, but proprioceptive drift did. This suggests that multisensory integration of bodily information drives this shift in space around the body and not feelings of ownership.

The image features of emotional faces that predict the initial eye movement to a face.

Emotional facial expressions are important visual communication signals that indicate a sender's intent and emotional state to an observer. As such, it is not surprising that reactions to different expressions are thought to be automatic and independent of awareness. What is surprising, is that studies show inconsistent results concerning such automatic reactions, particularly when using different face stimuli. We argue that automatic reactions to facial expressions can be better explained, and better understood, in terms of quantitative descriptions of their low-level image features rather than in terms of the emotional content (e.g. angry) of the expressions. Here, we focused on overall spatial frequency (SF) and localized Histograms of Oriented Gradients (HOG) features. We used machine learning classification to reveal the SF and HOG features that are sufficient for classification of the initial eye movement towards one out of two simultaneously presented faces. Interestingly, the identified features serve as better predictors than the emotional content of the expressions. We therefore propose that our modelling approach can further specify which visual features drive these and other behavioural effects related to emotional expressions, which can help solve the inconsistencies found in this line of research.

The additive nature of the human multisensory evoked pupil response.

Pupillometry has received increased interest for its usefulness in measuring various sensory processes as an alternative to behavioural assessments. This is also apparent for multisensory investigations. Studies of the multisensory pupil response, however, have produced conflicting results. Some studies observed super-additive multisensory pupil responses, indicative of multisensory integration (MSI). Others observed additive multisensory pupil responses even though reaction time (RT) measures were indicative of MSI. Therefore, in the present study, we investigated the nature of the multisensory pupil response by combining methodological approaches of previous studies while using supra-threshold stimuli only. In two experiments we presented auditory and visual stimuli to observers that evoked a(n) (onset) response (be it constriction or dilation) in a simple detection task and a change detection task. In both experiments, the RT data indicated MSI as shown by race model inequality violation. Still, the multisensory pupil response in both experiments could best be explained by linear summation of the unisensory pupil responses. We conclude that the multisensory pupil response for supra-threshold stimuli is additive in nature and cannot be used as a measure of MSI, as only a departure from additivity can unequivocally demonstrate an interaction between the senses.

Body ownership and the absence of touch: approaching the rubber hand inside and outside peri-hand space.

It is widely accepted that the integration of visual and tactile information is a necessity to induce ownership over a rubber hand. This idea has recently been challenged by Ferri et al. (Proc R Soc B 280:1-7, 2013), as they found that sense of ownership was evident by mere expectation of touch. In our study, we aimed to further investigate this finding, by studying whether the mere potential for touch yields a sense of ownership similar in magnitude to that resulting from actually being touched. We conducted two experiments. In the first experiment, our set-up was the classical horizontal set-up (similar to Botvinick and Cohen, Nature 391:756, 1998). Sixty-three individuals were included and performed the classical conditions (synchronous, asynchronous), an approached but not touched (potential for touch), and a 'visual only' condition. In the second experiment, we controlled for differences between the current set-up and the vertical set-up used by Ferri et al. (Proc R Soc B 280:1-7, 2013). Fifteen individuals were included and performed a synchronous and various approaching conditions [i.e., vertical approach, horizontal approach, and a control approach (no hands)]. In our first experiment, we found that approaching the rubber hand neither induced a larger proprioceptive drift nor a stronger subjective sense of ownership than asynchronous stimulation did. Generally, our participants gained most sense of ownership in the synchronous condition, followed by the visual only condition. When using a vertical set-up (second experiment), we confirmed previous suggestions that tactile expectation was able to induce embodiment over a foreign hand, similar in magnitude to actual touch, but only when the real and rubber hand were aligned on the vertical axis, thus along the trajectory of the approaching stimulus. These results indicate that our brain uses bottom-up sensory information, as well as top-down predictions for building a representation of our body.

Audiovisual integration in near and far space: effects of changes in distance and stimulus effectiveness.

A factor that is often not considered in multisensory research is the distance from which information is presented. Interestingly, various studies have shown that the distance at which information is presented can modulate the strength of multisensory interactions. In addition, our everyday multisensory experience in near and far space is rather asymmetrical in terms of retinal image size and stimulus intensity. This asymmetry is the result of the relation between the stimulus-observer distance and its retinal image size and intensity: an object that is further away is generally smaller on the retina as compared to the same object when it is presented nearer. Similarly, auditory intensity decreases as the distance from the observer increases. We investigated how each of these factors alone, and their combination, affected audiovisual integration. Unimodal and bimodal stimuli were presented in near and far space, with and without controlling for distance-dependent changes in retinal image size and intensity. Audiovisual integration was enhanced for stimuli that were presented in far space as compared to near space, but only when the stimuli were not corrected for visual angle and intensity. The same decrease in intensity and retinal size in near space did not enhance audiovisual integration, indicating that these results cannot be explained by changes in stimulus efficacy or an increase in distance alone, but rather by an interaction between these factors. The results are discussed in the context of multisensory experience and spatial uncertainty, and underline the importance of studying multisensory integration in the depth space.

Keep your eyes on development: the behavioral and neurophysiological development of visual mechanisms underlying form processing.

Visual form perception is essential for correct interpretation of, and interaction with, our environment. Form perception depends on visual acuity and processing of specific form characteristics, such as luminance contrast, spatial frequency, color, orientation, depth, and even motion information. As other cognitive processes, form perception matures with age. This paper aims at providing a concise overview of our current understanding of the typical development, from birth to adulthood, of form-characteristic processing, as measured both behaviorally and neurophysiologically. Two main conclusions can be drawn. First, the current literature conveys that for most reviewed characteristics a developmental pattern is apparent. These trajectories are discussed in relation to the organization of the visual system. The second conclusion is that significant gaps in the literature exist for several age-ranges. To complete our understanding of the typical and, by consequence, atypical development of visual mechanisms underlying form processing, future research should uncover these missing segments.

Slow and fast visual motion channels have independent binocular-rivalry stages.

We have previously reported a transparent motion after-effect indicating that the human visual system comprises separate slow and fast motion channels. Here, we report that the presentation of a fast motion in one eye and a slow motion in the other eye does not result in binocular rivalry but in a clear percept of transparent motion. We call this new visual phenomenon 'dichoptic motion transparency' (DMT). So far only the DMT phenomenon and the two motion after-effects (the 'classical' motion after-effect, seen after motion adaptation on a static test pattern, and the dynamic motion after-effect, seen on a dynamic-noise test pattern) appear to isolate the channels completely. The speed ranges of the slow and fast channels overlap strongly and are observer dependent. A model is presented that links after-effect durations of an observer to the probability of rivalry or DMT as a function of dichoptic velocity combinations. Model results support the assumption of two highly independent channels showing only within-channel rivalry, and no rivalry or after-effect interactions between the channels. The finding of two independent motion vision channels, each with a separate rivalry stage and a private line to conscious perception, might be helpful in visualizing or analysing pathways to consciousness.

Spatial structure, contrast polarity and motion integration.

It has been shown that in the initial stages of motion processing, the ON and OFF pathways stay more or less separated. There is evidence that this distinction between motion signals from opposite contrast polarities remains at least partly intact in the integration stage of local motion information. At the same time, interactions between the two systems are also apparent. Here we constructed stimuli that contained a constant number of moving checks. The checks were either assigned only one contrast polarity, or contrast polarity was distributed across the checks either randomly or evenly. We investigated how the spatial configuration of the moving stimulus affected direction discrimination thresholds for the different polarity distributions. Our results provide new evidence for contrast-sign-specific integration of local motion signals within areas of limited size, and inhibitory interactions between these separate ON and OFF motion sensor pools.

Integration after adaptation to transparent motion: static and dynamic test patterns result in different aftereffect directions.

One of the many interesting questions in motion aftereffect (MAE) research is concerned with the location(s) along the pathway of visual processing at which certain perceptual manifestations of this illusory motion originate. One such manifestation is the unidirectionality of the MAE after adaptation to moving plaids or transparent motion. This unidirectionality has led to the suggestion that the origin of this MAE might be a single source (gain control) located at, or beyond areas that are believed to be responsible for the integration of motion signals. In this report we present evidence against this suggestion using a simple experiment. For the same adaptation pattern, which consisted of two orthogonally moving transparent patterns with different speeds, we show that the direction of the resulting unidirectional MAE depends on the nature of the test stimulus. We used two kinds of test patterns: static and dynamic. For exactly the same adaptation conditions, the difference in MAE direction between testing with static and dynamic patterns can be as large as 50 degrees. This finding suggests that this MAE is not just a perceptual manifestation of a passive recovery of adapted motion sensors but an active integrative process using the output of different gain controls. A process which takes place after adaptation. These findings are in line with the idea that there are several sites of adaptation along the pathway of visual motion processing and that the nature of the test pattern determines the fate of our perceptual experience of the MAE.

Integration and segregation of local motion signals: the role of contrast polarity.

In the initial stages of visual processing in primates, more or less separated ON and OFF pathways have been shown to exist. There is ample evidence, that this separation includes the initial stages of motion processing. In the present study, experiments were conducted to investigate whether this ON versus OFF distinction persists into the integration stage of local motion information. We constructed stimuli that consisted of clusters of checks with equal contrast polarity, which could be varied in size, and compared them to stimuli with a random polarity distribution. We found that the ON versus OFF distinction remains partly intact, while interactions between the two systems are also apparent. These interactions prove to be highly correlated with the spatial structure of the stimulus. We propose a mechanism of contrast-sign specific integration of local motion signals, after which these separate ON and OFF pools engage in mutually inhibitory interactions.

Motion aftereffect of combined first-order and second-order motion.

When, after prolonged viewing of a moving stimulus, a stationary (test) pattern is presented to an observer, this results in an illusory movement in the direction opposite to the adapting motion. Typically, this motion aftereffect (MAE) does not occur after adaptation to a second-order motion stimulus (i.e. an equiluminous stimulus where the movement is defined by a contrast or texture border, not by a luminance border). However, a MAE of second-order motion is perceived when, instead of a static test pattern, a dynamic test pattern is used. Here, we investigate whether a second-order motion stimulus does affect the MAE on a static test pattern (sMAE), when second-order motion is presented in combination with first-order motion during adaptation. The results show that this is indeed the case. Although the second-order motion stimulus is too weak to produce a convincing sMAE on its own, its influence on the sMAE is of equal strength to that of the first-order motion component, when they are adapted to simultaneously. The results suggest that the perceptual appearance of the sMAE originates from the site where first-order and second-order motion are integrated.

Local and global factors affecting the coherent motion of gratings presented in multiple apertures.

Using stimuli composed of two independent gratings viewed through multiple apertures, we investigate a number of parameters affecting the integration of locally ambiguous motions into globally coherent motion. In four experiments, we varied local factors (grating spatial frequency, speed, contrast, duty cycle, orientation) and global factors (degree of similarity and common fate between the gratings, and symmetry in the configuration of the grating pattern) and examined their effects on global motion coherence. Our results, confirming accounts offered by previous investigators, indicate that local competition between motion signals generated by contours (ambiguous) and their line terminations (unambiguous) is important in determining global motion coherence in multiple-aperture stimuli. Our results also indicate that global factors can affect perceived coherence independently of local motion signals, suggesting the involvement of higher-level motion areas and a role for non-motion processes such as those involved in pattern and form perception. Comparing motion coherence with other two-dimensional (2-D) stimuli (plaids) shows that 2-D multiple-aperture stimuli are not analogous and that coherence models derived from plaid stimuli do not account for the data.

Aftereffect of high-speed motion.

A visual illusion known as the motion aftereffect is considered to be the perceptual manifestation of motion sensors that are recovering from adaptation. This aftereffect can be obtained for a specific range of adaptation speeds with its magnitude generally peaking for speeds around 3 deg s-1. The classic motion aftereffect is usually measured with a static test pattern. Here, we measured the magnitude of the motion aftereffect for a large range of velocities covering also higher speeds, using both static and dynamic test patterns. The results suggest that at least two (sub)populations of motion-sensitive neurons underlie these motion aftereffects. One population shows itself under static test conditions and is dominant for low adaptation speeds, and the other is prevalent under dynamic test conditions after adaptation to high speeds. The dynamic motion aftereffect can be perceived for adaptation speeds up to three times as fast as the static motion aftereffect. We tested predictions that follow from the hypothesised division in neuronal substrates. We found that for exactly the same adaptation conditions (oppositely directed transparent motion with different speeds), the aftereffect direction differs by 180 degrees depending on the test pattern. The motion aftereffect is opposite to the pattern moving at low speed when the test pattern is static, and opposite to the high-speed pattern for a dynamic test pattern. The determining factor is the combination of adaptation speed and type of test pattern.

Monocular mechanisms determine plaid motion coherence.

Although the neural location of the plaid motion coherence process is not precisely known, the middle temporal (MT) cortical area has been proposed as a likely candidate. This claim rests largely on the neurophysiological findings showing that in response to plaid stimuli, a subgroup of cells in area MT responds to the pattern direction, whereas cells in area V1 respond only to the directions of the component gratings. In Experiment 1, we report that the coherent motion of a plaid pattern can be completely abolished following adaptation to a grating which moves in the plaid direction and has the same spatial period as the plaid features (the so-called "blobs"). Interestingly, we find this phenomenon is monocular: monocular adaptation destroys plaid coherence in the exposed eye but leaves it unaffected in the other eye. Experiment 2 demonstrates that adaptation to a purely binocular (dichoptic) grating does not affect perceived plaid coherence. These data suggest several conclusions: (1) that the mechanism determining plaid coherence responds to the motion of plaid features, (2) that the coherence mechanism is monocular, and thus (3), that it is probably located at a relatively low level in the visual system and peripherally to the binocular mechanisms commonly presumed to underlie two-dimensional (2-D) motion perception. Experiment 3 examines the spatial tuning of the monocular coherence mechanism and our results suggest it is broadly tuned with a preference for lower spatial frequencies. In Experiment 4, we examine whether perceived plaid direction is determined by the motion of the grating components or the features. Our data strongly support a feature-based model.

The perceived direction of textured gratings and their motion aftereffects.

The stimuli in these experiments are square-wave luminance gratings with an array of small random dots covering the high-luminance regions. Owing to the texture, the direction of these gratings, when seen through a circular aperture, is disambiguated because the visual system is provided with an unambiguous motion energy. Thus, the direction of textured gratings can be varied independently of grating orientation. When subjects are required to judge the direction of textured gratings moving obliquely relative to their orientation, they can do so accurately (experiment 1). This is of interest because most studies of one-dimensional motion perception have involved (textureless) luminance-defined since-wave or square-wave gratings, and the perceived direction of these gratings is constrained by the aperture problem to be orthogonal to their orientation. Thus, direction and orientation have often been confounded. Interestingly, when subjects are required to judge the direction of an obliquely moving textured grating during a period of adaptation and then the direction of the motion aftereffect (MAE) immediately following adaptation (experiments 2 and 3), these directions are not directly opposite each other. MAE directions were always more orthogonal to the orientation of the adapting grating than the corresponding direction judgments during adaptation (by as much as 25 degrees). These results are not readily explained by conventional MAE models and possible accounts are considered.