Adaptation to visual numerosity changes neural numerosity selectivity.

Perceiving numerosity, i.e. the set size of a group of items, is an evolutionarily preserved ability found in humans and animals. A useful method to infer the neural underpinnings of a given perceptual property is sensory adaptation. Like other primary perceptual attributes, numerosity is susceptible to adaptation. Recently, we have shown numerosity-selective neural populations with a topographic organization in the human brain. Here, we investigated whether numerosity adaptation can affect the numerosity selectivity of these populations using ultra-high field (7 Tesla) functional magnetic resonance imaging (fMRI). Participants viewed stimuli of changing numerosity (1 to 7 dots), which allowed the mapping of numerosity selectivity. We interleaved a low or high numerosity adapter stimulus with these mapping stimuli, repeatedly presenting 1 or 20 dots respectively to adapt the numerosity-selective neural populations. We analyzed the responses using custom-build population receptive field neural models of numerosity encoding and compared estimated numerosity preferences between adaptation conditions. We replicated our previous studies where we found several topographic maps of numerosity-selective responses. We found that overall, numerosity adaptation altered the preferred numerosities within the numerosity maps, resulting in predominantly attractive biases towards the numerosity of the adapter. The differential biases could be explained by the difference between the unadapted preferred numerosity and the numerosity of the adapter, with attractive biases being observed with higher difference. The results could link perceptual numerosity adaptation effects to changes in neural numerosity selectivity.

Distinct temporal mechanisms modulate numerosity perception.

Our ability to process numerical and temporal information is an evolutionary skill thought to originate from a common magnitude system. In line with a common magnitude system, we have previously shown that adaptation to duration alters numerosity perception. Here, we investigate two hypotheses on how duration influences numerosity perception. A channel-based hypothesis predicts that numerosity perception is influenced by adaptation of onset/offset duration channels which also encode numerosity or wire together with numerosity channels (duration/numerosity channels). Hence, the onset/offset duration of the adapter is driving the effect regardless of the total duration of adaptation. A strength-of-adaptation hypothesis predicts that the effect of duration on numerosity perception is driven by the adaptation of numerosity channels only, with the total duration of adaptation driving the effect regardless of the onset/offset duration of the adapter. We performed two experiments where we manipulated the onset/offset duration of the adapter, the adapter's total presentation time, and the total duration of the adaptation trial. The first experiment tested the effect of adaptation to duration on numerosity discrimination, whereas the second experiment tested the effect of adaptation to numerosity and duration on numerosity discrimination. We found that the effect of adaptation to duration on numerosity perception is primarily driven by adapting duration/numerosity channels, supporting the channel-based hypothesis. In contrast, the effect of adaptation to numerosity on numerosity perception appears to be driven by the total duration of the adaptation trial, supporting the strength-of-adaptation hypothesis. Thus, we show that adaptation of at least two temporal mechanisms influences numerosity perception.

Interaction effects of visual stimulus speed and contrast on postural sway.

Manipulating the characteristics of visual stimuli that simulate self-motion through the environment can affect the resulting postural sway magnitude. In the present study, we address the question whether varying the contrast and speed of a linear translating dot pattern influences medial-lateral postural sway. In a first experiment, we investigated whether the postural sway magnitude increases with increasing dot speed, as was previously demonstrated for expanding and contracting stimuli. In a second experiment, we also manipulated the contrast of the stimuli. For reasons that high-contrast stimuli can be considered 'perceptually' stronger, we expect that higher-contrast stimuli induce more sway than lower-contrast stimuli. The results of the first experiment show that dot speed indeed influences postural sway, although in an unexpected way. For higher speeds, the sway is in the direction of the stimulus motion, yet for lower speeds the sway is in a direction opposite to the stimulus motion. The results of the second experiment show that dot contrast does affect postural sway, but that this depends on the speed of the moving dots. Interestingly, the direction of postural sway induced by a relatively low dot speed (4°/s) depends on dot contrast. Taken together, our results suggest that interactions between the visual, vestibular and proprioceptive system appear to be influenced by an internal representation of the visual stimulus, rather than being influenced by the external visual stimulus characteristics only.

Cross-modal, bidirectional priming in grapheme-color synesthesia.

Grapheme-color synesthetes perceive achromatic graphemes to be inherently colored. In this study grapheme-color synesthetes and non-synesthetes discriminated (1) the color of visual targets presented along with aurally presented digit primes, and (2) the identity of aurally presented digit targets presented with visual color primes. Reaction times to visual color targets were longer when the color of the target was incongruent with the synesthetic percept reported for the prime. Likewise, discriminating aurally presented digit targets took longer when the color of the prime was incongruent with the synesthetic percept for the target. These priming effects were absent in non-synesthetes. We conclude that binding between digits and colors in grapheme-color synesthetes can occur bidirectionally across senses. The results are in line with the idea that synesthesia is the result of linking inducing stimuli (e.g. digits) to synesthetic percepts (colors) at an abstract - supra-modal - conceptual level of processing.

CT-optimal touch and chronic pain experience in Parkinson's Disease; An intervention study.

One of the most underdiagnosed and undertreated non-motor symptoms of Parkinson's Disease is chronic pain. This is generally treated with analgesics which is not always effective and can cause several side-effects. Therefore, new ways to reduce chronic pain are needed. Several experimental studies show that CT-optimal touch can reduce acute pain. However, little is known about the effect of CT-optimal touch on chronic pain. The aim of the current study is to investigate whether CT-optimal touch can reduce the chronic pain experience in Parkinson patients. In this intervention study, 17 Parkinson patients underwent three conditions; no touch, CT-optimal touch and CT non-optimal touch with a duration of one week each. During each touch week, participants received touch from their partners twice a day for 15 minutes. Results show that both types of touch ameliorate the chronic pain experience. Furthermore, it appears that it is slightly more beneficial to apply CT-optimal touch also because it is perceived as more pleasant. Therefore, we argue that CT-optimal touch might be used when immediate pain relief is needed. Importantly, this study shows that CT-optimal touch can reduce chronic pain in Parkinson's Disease and can be administered by a partner which makes it feasible to implement CT-optimal touch as daily routine.

Spatial factors influencing the pain-ameliorating effect of CT-optimal touch: a comparative study for modulating temporal summation of second pain.

Recent studies show that CT-optimal touch, gentle slow stroking of the skin, can reduce pain. However, much is unknown regarding the factors influencing its pain-ameliorating effect, such as tactile attention and touch application site. The current study investigates in 36 healthy individuals, whether CT-optimal touch can reduce temporal summation of second pain (TSSP) compared to CT non-optimal touch and tapping the skin. TSSP refers to activation of the C-nociceptors; by stimulating these fibers a burning and/or tingling sensation can be elicited. All participants underwent three conditions on both the contralateral and ipsilateral side of pain induction. The results show that tapping the skin did not reduce TSSP, meaning that pain reduction through touch cannot be explained by tactile attention effects. CT non-optimal touch only reduced TSSP when applied on the ipsilateral side. Importantly, CT-optimal touch effectively reduced TSSP when applied on the contralateral or ipsilateral side. Furthermore, CT-optimal touch was more effective in reducing TSSP compared to CT non-optimal touch and Tapping. This study shows that that CT-optimal touch can reduce TSSP and this effect appears to be independent of touch application site, which is highly relevant for implementing CT-optimal touch as a treatment.

Decreasing perceived optic flow rigidity increases postural sway.

Optic flow simulating self-motion through the environment can induce postural adjustments in observers. Some studies investigating this phenomenon have used optic flow patterns increasing in speed from center to periphery, whereas others used optic flow patterns with a constant speed. However, altering the speed gradient of an optic flow stimulus changes the perceived rigidity of such a stimulus. Optic flow stimuli that are perceived as rigid can be expected to provide a stronger sensation of self-motion than non-rigid optic flow, and this may well be reflected in the amount of postural sway. The current study, therefore, examined, by manipulating the speed gradient, to what extent the rigidity of an optic flow stimulus influences posture along the anterior-posterior axis. We used radial random dot expanding or contracting optic flow patterns with three different speed profiles (single-speed, linear speed gradient or quadratic speed gradient) that differentially induce the sensation of self-motion. Interestingly, most postural sway was observed for the non-rigid single-speed optic flow pattern, which contained the least self-motion information of the three profiles. Moreover, we found an anisotropy in that contracting optic flow produced more postural sway than expanding optic flow. In addition, the amount of postural sway increased with increasing stimulus speed, but for contracting optic flow only. Taken together, the results of the current study support the view that visual and sensorimotor systems appear to be tailored toward compensating for rigid optic flow stimulation.

Chronic pain relief after receiving affective touch: A single case report.

Affective touch is gentle slow stroking of the skin, which can reduce experimentally induced pain. Our participant, suffering from Parkinson's Disease and chronic pain, received 1 week of non-affective touch and 1 week of affective touch as part of a larger study. Interestingly, after 2 days of receiving affective touch, the participant started to feel less pain. After 7 days, the burning painful sensations fully disappeared. This suggest that affective touch may reduce chronic pain in clinical populations.

Neural basis of affective touch and pain: A novel model suggests possible targets for pain amelioration.

Pain is one of the most common health problems and has a severe impact on quality of life. Yet, a suitable and efficient treatment is still not available for all patient populations suffering from pain. Interestingly, recent research shows that low threshold mechanosensory C-tactile (CT) fibres have a modulatory influence on pain. CT-fibres are activated by slow gentle stroking of the hairy skin, providing a pleasant sensation. Consequently, slow gentle stroking is known as affective touch. Currently, a clear overview of the way affective touch modulates pain, at a neural level, is missing. This review aims to present such an overview. To explain the interaction between affective touch and pain, first the neural basis of the affective touch system and the neural processing of pain will be described. To clarify these systems, a schematic illustration will be provided in every section. Hereafter, a novel model of interactions between affective touch and pain systems will be introduced. Finally, since affective touch might be suitable as a new treatment for chronic pain, possible clinical implications will be discussed.

The role of neural tuning in quantity perception.

Perception of quantities, such as numerosity, timing, and size, is essential for behavior and cognition. Accumulating evidence demonstrates neurons processing quantities are tuned, that is, have a preferred quantity amount, not only for numerosity, but also other quantity dimensions and sensory modalities. We argue that quantity-tuned neurons are fundamental to understanding quantity perception. We illustrate how the properties of quantity-tuned neurons can underlie a range of perceptual phenomena. Furthermore, quantity-tuned neurons are organized in distinct but overlapping topographic maps. We suggest that this overlap in tuning provides the neural basis for perceptual interactions between different quantities, without the need for a common neural representational code.

Suppressed images selectively affect the dominant percept during binocular rivalry.

During binocular rivalry, perception alternates between dissimilar images that are presented dichoptically. It has been argued that perception during the dominance phase of rivalry is unaffected by the suppressed image. Recent evidence suggests, however, that the suppressed image does affect perception of the dominant image, yet the extent and nature of this interaction remain elusive. We hypothesize that this interaction depends on the difference in feature content between the rivaling images. Here, we investigate how sensitivity to probes presented in the image that is currently dominant in perception is affected by the suppressed image. Observers performed a 2AFC discrimination task on oriented probes (Experiment 1) or probes with different motion directions (Experiment 2). Our results show that performance on both orientation and motion direction discrimination was affected by the content of the suppressed image. The strength of interference depended specifically on the difference in feature content (e.g., the difference in orientation) between the probe and the suppressed image. Moreover, the pattern of interference by the suppressed image is qualitatively similar to the situation where this image and the probe are simultaneously visible. We conclude that perception during the dominance phase of rivalry is affected by a suppressed image as if it were visible.

Multiple dimensions in bi-directional synesthesia.

Grapheme-color synesthetes report seeing a specific color when a number is perceived. The reverse, the synesthetic experience of a specific grapheme after the percept of a color is extremely rare. However, recent studies have revealed these interactions at both behavioral and neurophysiological levels. We investigated whether similar neuronal processes (i.e. perceptual and/or attentional) may underlie this bi-directional interaction by measuring event-related potentials (ERPs) during both a number-color and color-number priming task. In addition, we investigated the unitarity of synesthesia by comparing two distinct subtypes of synesthetes, projectors and associators, and assessed whether consistencies between measures (i.e. behavioral and electrophysiological) were present across synesthetes. Our results show longer reaction times for incongruent compared with congruent trials in both tasks. This priming effect is also present in the P3b latency (parietal electrode site) and P3a amplitude (frontal electrode site) of the ERP data. Interestingly, projector and associator synesthetes did not reveal distinct behavioral or electrophysiological patterns. Instead, a dissociation was found when synesthetes were divided in two groups on the basis of their behavioral data. Synesthetes with a large behavioral priming effect revealed ERP modulation at the frontal and parietal electrode sites, whereas synesthetes with a small priming effect revealed a frontal effect only. Together, these results show, for the first time, that similar neural mechanisms underlie bi-directional synesthesia in synesthetes that do not report a synesthetic experience of a grapheme when a color is perceived. In addition, they add support for the notion of the existence of both 'lower' and 'higher' synesthetes.

The development of automated access to symbolic and non-symbolic number knowledge in children: an ERP study.

Infants can visually detect changes in numerosity, which suggests that a (non-symbolic) numerosity system is already present early in life. This non-symbolic system is hypothesized to serve as the basis for the later acquired symbolic system. Little is known about the processes underlying the transition from the non-symbolic to symbolic code. In the current study we investigated the development of automatization of symbolic number processing in children from second (6.0 years) and fourth grade (8.0 years) and adults using a symbolic and non-symbolic size congruency task and event-related potentials (ERPs) as a measure. The comparison between symbolic and non-symbolic size congruency effects (SCEs) allowed us to disentangle processes necessary to perform the task from processes specific to numerosity notation. In contrast to previous studies, second graders already revealed a behavioral symbolic SCE similar to that of adults. In addition, the behavioral SCE increased for symbolic and decreased for non-symbolic notation with increasing age. For all age groups, the ERP data showed that the two magnitudes interfered at a level before selective activation of the response system, for both notations. However, only for the second graders distinct processes were recruited to perform the symbolic size comparison task. This shift in recruited processes for the symbolic task only might reflect the functional specialization of the parietal cortex.

A selective deficit in the appreciation and recognition of brightness: brightness agnosia?

We report a patient with extensive brain damage in the right hemisphere who demonstrated a severe impairment in the appreciation of brightness. Acuity, contrast sensitivity as well as luminance discrimination were normal, suggesting her brightness impairment is not a mere consequence of low-level sensory impairments. The patient was not able to indicate the darker or the lighter of two grey squares, even though she was able to see that they differed. In addition, she could not indicate whether the lights in a room were switched on or off, nor was she able to differentiate between normal greyscale images and inverted greyscale images. As the patient recognised objects, colours, and shapes correctly, the impairment is specific for brightness. As low-level, sensory processing is normal, this specific deficit in the recognition and appreciation of brightness appears to be of a higher, cognitive level, the level of semantic knowledge. This appears to be the first report of 'brightness agnosia'.

Adaptation reveals unbalanced interaction between numerosity and time.

Processing quantities such as the number of objects in a set, size, spatial arrangement and time is an essential means of structuring the external world and preparing for action. The theory of magnitude suggests that number and time, among other continuous magnitudes, are linked by a common cortical metric, and their specialization develops from a single magnitude system. In order to investigate potentially shared neural mechanisms underlying numerosity and time processing, we used visual adaptation, a method which can reveal the existence of a dedicated processing system. We reasoned that cross-adaptation between numerosity and duration would concur with the existence of a common processing mechanism, whereas the absence of cross-adaptation would provide evidence against it. We conducted four experiments using a rapid adaptation protocol where participants adapted to either visual numerosity or visual duration and subsequently performed a numerosity or duration discrimination task. We found that adapting to a low numerosity altered the estimation of the reference numerosity by an average of 5 dots, compared to adapting to a high numerosity. Similarly, adapting to a short duration altered the estimation of the reference duration by an average of 43 msec, compared to adapting to a long duration. In the cross-dimensional adaptation conditions, duration adaptation altered numerosity estimation by an average of 1 dot, whereas there was not sufficient evidence to either support or reject the effect of numerosity adaptation on duration judgments. These results highlight that there are partially overlapping neural mechanisms which are dedicated for processing both numerosity and time.

Disentangling neural structures for processing of high- and low-speed visual motion.

Human psychophysical and electrophysiological evidence suggests at least two separate visual motion pathways, one tuned to a lower and one tuned to a broader and partly overlapping range of higher speeds. It remains unclear whether these two different channels are represented by different cortical areas or by sub-populations within a single area. We recorded evoked potentials at 59 scalp locations to the onset of a slow (3.5 degrees /s) and fast (32 degrees /s) moving test pattern, preceded by either a slow or fast adapting pattern that moved in either the same direction or opposite to the test motion. Baseline potentials were recorded for slow and fast moving test patterns after adaptation to a static pattern. Comparison of adapted responses with baseline responses revealed that the N2 peak around 180 ms after test stimulus onset was modulated by the preceding adaptation. This modulation depended on both direction and speed. Source localization of baseline potentials as well as direction-independent motion adaptation revealed cortical areas activated by fast motion to be more dorsal, medial and posterior compared with neural structures underlying slow motion processing. For both speeds, the direction-dependent component of this adaptation modulation occurred in the same area, located significantly more dorsally compared with neural structures that were adapted in a direction-independent manner. These results demonstrate for the first time the cortical separation of more ventral areas selectively activated by visual motion at low speeds (and not high speeds) and dorsal motion-sensitive cortical areas that are activated by both high and low speeds.

Recognising the forest, but not the trees: an effect of colour on scene perception and recognition.

Colour has been shown to facilitate the recognition of scene images, but only when these images contain natural scenes, for which colour is 'diagnostic'. Here we investigate whether colour can also facilitate memory for scene images, and whether this would hold for natural scenes in particular. In the first experiment participants first studied a set of colour and greyscale natural and man-made scene images. Next, the same images were presented, randomly mixed with a different set. Participants were asked to indicate whether they had seen the images during the study phase. Surprisingly, performance was better for greyscale than for coloured images, and this difference is due to the higher false alarm rate for both natural and man-made coloured scenes. We hypothesized that this increase in false alarm rate was due to a shift from scrutinizing details of the image to recognition of the gist of the (coloured) image. A second experiment, utilizing images without a nameable gist, confirmed this hypothesis as participants now performed equally on greyscale and coloured images. In the final experiment we specifically targeted the more detail-based perception and recognition for greyscale images versus the more gist-based perception and recognition for coloured images with a change detection paradigm. The results show that changes to images are detected faster when image-pairs were presented in greyscale than in colour. This counterintuitive result held for both natural and man-made scenes (but not for scenes without nameable gist) and thus corroborates the shift from more detailed processing of images in greyscale to more gist-based processing of coloured images.

Occlusion and the solution to visual motion ambiguity: Looking beyond the aperture problem.

A horizontally moving grating viewed within a diamond-shaped aperture can be made to appear to move obliquely by introducing appropriate depth-ordering cues (R. O. Duncan, T. D. Albright, & G. R. Stoner, 2000). It is commonly assumed that the depth cues in such displays determine which line terminators are seen as intrinsic to the grating and which are seen as resulting from occlusion and hence extrinsic to the grating. The ambiguous motion of the grating (arising from the aperture problem) is then supposed to be overcome by selectively pooling motion signals arising from the intrinsic terminators with those arising from the grating while discounting the motion of the extrinsic terminators. In our first experiment, we tested the sufficiency of this explanation. Observers reported the direction of motion of ambiguously moving random dots viewed through a diamond-shaped aperture defined by four panels. Binocular disparity was used to simulate occlusion: two panels occluded the virtual surface upon which the dots were positioned and two panels were occluded by that surface. Reports were significantly biased toward the direction of the occluding panels. Since none of the moving features abutted the surrounding panels, none should have been classified as extrinsic and hence this result cannot have relied on terminator classification. In a second experiment, we tested the hypothesis that depth-ordering cues selectively gate the propagation of motion signals so that the representation of the moving surface extends behind the occluders. This was tested by asking observers to report the direction of moving dots viewed through a briefly "opened" probe window within either occluding or occluded panels. Consistent with our hypothesis, evidence of motion propagation was only found for probe windows within occluding panels. Surprisingly, however, this propagation was only observed when the dots in the inducer window moved away from the probe window, suggesting a "pull," and not a "push" mechanism.

Inducing circular vection with tactile stimulation encircling the waist.

In general, moving sensory stimuli (visual and auditory) can induce illusory sensations of self-motion (i.e. vection) in the direction opposite of the sensory stimulation. The aim of the current study was to examine whether tactile stimulation encircling the waist could induce circular vection (around the body's yaw axis) and to examine whether this type of stimulation would influence participants' walking trajectory and balance. We assessed the strength and direction of perceived self-motion while vision was blocked and while either receiving tactile stimulation encircling the waist clockwise or counterclockwise or no tactile stimulation. Additionally, we assessed participants' walking trajectory and balance while receiving these different stimulations. Tactile stimulation encircling the waist was found to lead to self-reported circular vection in a subset of participants. In this subset of participants, circular vection was on average experienced in the same direction as the tactile stimulation. Additionally, perceived rotatory self-motion in participants that reported circular vection correlated with balance (i.e., sway velocity and the standard error of the mean in the medio-lateral dimension). The fact that, in this subset of participants, subjective reports of vection correlated with objective outcome measures indicates that tactile stimulation encircling the waist might indeed be able to induced circular vection.