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International audience; We tested whether the acquisition of grapheme-color synesthesia during childhood is related to difficulties in written language learning by measuring whether it is more frequent in 79 children receiving speech and language therapy for such difficulties than in the general population of children (1.3%). By using criteria as similar as possible to those used in the reference study (Simner et al., 2009), we did not identify any synesthete (Bayesian 95% credible interval [0, 4.5]% for a flat prior). The odds of the null model (no difference between 0/79 and 1.3%) over alternative models is 28 (Bayes Factor). A higher prevalence of grapheme-color synesthetes among children with learning difficulties is therefore very unlikely, questioning the hypothesis of a link between synesthesia and difficulties in language acquisition. We also describe the difficulty of diagnosing synesthesia in children and discuss the need for new approaches to do so.

Nabokov belongs to a small “family” of great artists also known as synesthetes, suggesting a link between synesthesia and creativity. Moreover, Nabokov’s report of precise synesthetic associations between letters and colors already in his early childhood suggests that synesthesia is a gift, possibly innate. Could we suppose that the brain of Nabokov was special because of synesthesia? Could synesthesia relate to his metaphoric creativity, his strong visual mental imagery and memory or even some aspects of his personality? Here we show that the synesthetic experiences described by Nabokov are similar to those of synesthetes studied by scientists for the last twenty years. We review what cognitive neuroscience has learnt so far about the possible mechanisms of synesthesia and whether synesthetes display any special characteristics beyond synesthesia. We show that little is known and that there is little support to naturalize Nabokov’s creative process on the basis of him having synesthesia.

The aim of the present study was to uncover a possible common neural organizing principle in spoken and written communication, through the coupling of perceptual and motor representations. In order to identify possible shared neural substrates for processing the basic units of spoken and written language, a sparse sampling fMRI acquisition protocol was performed on the same subjects in two experimental sessions with similar sets of letters being read and written and of phonemes being heard and orally produced. We found evidence of common premotor regions activated in spoken and written language, both in perception and in production. The location of those brain regions was confined to the left lateral and medial frontal cortices, at locations corresponding to the premotor cortex, inferior frontal cortex and supplementary motor area. Interestingly, the speaking and writing tasks also appeared to be controlled by largely overlapping networks, possibly indicating some domain general cognitive processing. Finally, the spatial distribution of individual activation peaks further showed more dorsal and more left-lateralized premotor activations in written than in spoken language.

International audience; Perception is sometimes bistable, switching between two possible interpretations. Levelt developed several propositions to explain bistable perception in binocular rivalry, based on a model of competing neural populations connected through reciprocal inhibition. Here we test Levelt's laws with bistable plaid motion. Plaids are typically tristable, either a coherent pattern, transparent with one component in front, or transparent with the opposite depth order. In Experiment 1, we use a large angle between component directions to prevent plaid coherence, limiting the ambiguity to alternations of grating depth order. Similar to increasing contrast in binocular rivalry, increasing component speed led to higher switch rates (analogous to Levelt's fourth proposition). In Experiment 2, we used occlusion cues to prevent one depth order and limit bistability to one transparent depth order alternating with coherence. Increasing grating speed shortened coherent motion periods but left transparent periods largely unchanged (analogous to Levelt's second proposition). Switch dynamics showed no correlation between the experiments. These data suggest that plaid component speed acts like contrast in binocular rivalry to vary switch dynamics through a mutual inhibition model. The lack of correlation between both experiments suggests reciprocal inhibition mediates bistability between a variety of neural populations across the visual system.

Several publications have reported structural changes in the brain of synesthetes compared to controls, either local differences or differences in connectivity. In the present study, we pursued this quest for structural brain differences that might support the subjective experience of synaesthesia. In particular, for the first time in this field, we investigated brain folding in comparing 45 sulcal shapes in each hemisphere of control and grapheme-color synesthete populations. To overcome flaws relative to data interpretation based only on p-values, common in the synesthesia literature, we report confidence intervals of effect sizes. Moreover, our statistical maps are displayed without introducing the classical, but misleading, p-value level threshold. We adopt such a methodological procedure to facilitate appropriate data interpretation and promote the New Statistics approach. Based on structural or diffusion magnetic resonance imaging data, we did not find any strong cerebral anomaly, in sulci, tissue volume, tissue density or fiber organization that could support synesthetic color experience. Finally, by sharing our complete datasets, we strongly support the multi-center construction of a sufficient large dataset repository for detecting, if any, subtle brain differences that may help understanding how a subjective experience, such as synesthesia, is mentally constructed.

Martinho and Kacelnik’s (Reports, 15 July 2016, p. 286) finding that mallard ducklings can deal with abstract concepts is important for understanding the evolution of cognition. However, a statistically more robust analysis of the data calls their conclusions into question. This example brings to light the risk of drawing too strong an inference by relying solely on P values.

Synesthetes, a small fraction of the population, experience systematic, additional associations. For example, they may arbitrarily associate a specific color to each letter or number. Synesthesia has offered for the last ten years to cognitive science a unique opportunity to study the neural bases of subjective experience, drawing on individual differences just like in neuropsychology, but with healthy people. Here we review the current knowledge and propose a new theory, the "palimpsest hypothesis", a variant of the recycling hypothesis for reading. The neural development of written language expertise (a recent cultural invention acquired without any genetic modification) requires indeed the recycling of brain regions predisposed to expertise acquisition into reading regions. The palimpsest hypothesis supposes that for synesthetes recycling involves neuronal networks that were already specialized for color perception. Synesthetic colors would be the remains of this former expertise.

Published studies using functional and structural MRI include many errors in the way data are analyzed and conclusions reported. This was observed when working on a comprehensive review of the neural bases of synesthesia, but these errors are probably endemic to neuroimaging studies. All studies reviewed had based their conclusions using Null Hypothesis Significance Tests (NHST). NHST have yet been criticized since their inception because they are more appropriate for taking decisions related to a Null hypothesis (like in manufacturing) than for making inferences about behavioral and neuronal processes. Here I focus on a few key problems of NHST related to brain imaging techniques, and explain why or when we should not rely on “significance” tests. I also observed that, often, the ill-posed logic of NHST was even not correctly applied, and describe what I identified as common mistakes or at least problematic practices in published papers, in light of what could be considered as the very basics of statistical inference. MRI statistics also involve much more complex issues than standard statistical inference. Analysis pipelines vary a lot between studies, even for those using the same software, and there is no consensus which pipeline is the best. I propose a synthetic view of the logic behind the possible methodological choices, and warn against the usage and interpretation of two statistical methods popular in brain imaging studies, the false discovery rate (FDR) procedure and permutation tests. I suggest that current models for the analysis of brain imaging data suffer from serious limitations and call for a revision taking into account the “new statistics” (confidence intervals) logic.

SummaryWhen dealing with natural scenes, sensory systems have to process an often messy and ambiguous flow of information. A stable perceptual organization nevertheless has to be achieved in order to guide behavior. The neural mechanisms involved can be highlighted by intrinsically ambiguous situations. In such cases, bistable perception occurs: distinct interpretations of the unchanging stimulus alternate spontaneously in the mind of the observer [1]. Bistable stimuli have been used extensively for more than two centuries to study visual perception [2]. Here we demonstrate that bistable perception also occurs in the auditory modality. We compared the temporal dynamics of percept alternations observed during auditory streaming [3, 4] with those observed for visual plaids [5, 6] and the susceptibilities of both modalities to volitional control. Strong similarities indicate that auditory and visual alternations share common principles of perceptual bistability. The absence of correlation across modalities for subject-specific biases, however, suggests that these common principles are implemented at least partly independently across sensory modalities. We propose that visual and auditory perceptual organization could rely on distributed but functionally similar neural competition mechanisms aimed at resolving sensory ambiguities.

Prolonged observations of moving plaids lead to bi-stable alternations between coherency and transparency. However, most studies of plaids used brief presentations and a 2AFC between the two interpretations, thus overlooking the dynamical aspect of plaid perception. In other domains, most notably binocular rivalry, it was shown that the dynamics of the bi-stable alternations reveal important insights about the underlying mechanisms. Here we develop methods to study the dynamics of plaid perception. Observers continually indicated their percept (coherency or transparency) during presentations that lasted 1–5 min. Two measures of the relative strength of the coherency percept were derived from those data: C=½C þ T � , the relative time spent seeing coherency, and RTtransp, the response time to report transparency. Those measures are independent of each other yet tightly correlated, and both show systematic relations to manipulations of plaid parameters. Furthermore, the two measures are sensitive to manipulations in wide parametric regimes, including ranges where brief-presentation methods suffer from ‘‘ceiling’’ and ‘‘floor’’ effects. We conclude that studying the dynamics of bi-stability in plaids can provide new and unsuspected findings about motion integration and segmentation. � 2002 Elsevier Science Ltd. All rights reserved.

Contrast-dependent changes in spatial summation and contextual modulation of primary visual cortex (V1) neuron responses to stimulation of their receptive field reveal long-distance integration of visual signals within V1, well beyond the classical receptive field (cRF) of single neurons. To identify the cortical circuits mediating these long-distance computations, we have used a combination of anatomical and physiological recording methods to determine the spatial scale and retinotopic logic of intra-areal V1 horizontal connections and inter-areal feedback connections to V1. We have then compared the spatial scales of these connectional systems to the spatial dimensions of the cRF, spatial summation field (SF), and modulatory surround field of macaque V1 neurons. We find that monosynaptic horizontal connections within area V1 are of an appropriate spatial scale to mediate interactions within the SF of V1 neurons and to underlie contrast-dependent changes in SF size. Contrary to common beliefs, these connections cannot fully account for the dimensions of the surround field. The spatial scale of feedback circuits from extrastriate cortex to V1 is, instead, commensurate with the full spatial range of center-surround interactions. Thus these connections could represent an anatomical substrate for contextual modulation and global-to-local integration of visual signals. Feedback projections connect corresponding and equal-sized regions of the visual field in striate and extrastriate cortices and cover anisotropic parts of visual space, unlike V1 horizontal connections that are isotropic in the macaque. V1 isotropic connectivity demonstrates that anisotropic horizontal connections are not necessary to generate orientation selectivity. Anisotropic feedback connections may play a role in contour completion.

Grapheme-color synesthesia, the idiosyncratic, arbitrary association of colors to letters or numbers, develops in childhood once reading is mastered. Because language processing is strongly left-lateralized in most individuals, we hypothesized that grapheme-color synesthesia could be left-lateralized as well. We used synesthetic versions of the Stroop test with colored letters and numbers presented either in the right or the left visual field of thirty-four synesthetes. Interference by synesthetic colors was stronger for stimuli in the right hemifield (first experiment, color naming task). Synesthetes were also faster in the right hemifield when naming the synesthetic color of graphemes (second experiment). Overall, the lateralization effect was 7 ms (the 95% confidence interval was [1.5 12] ms), a delay compatible with an additional callosal transfer for stimuli presented in the left hemifield. Though weak, this effect suggests that the association of synesthetic colors to graphemes may be preferentially processed in the left hemisphere. We speculate that this left-lateralization could be a landmark of synesthetic grapheme-color associations, if not found for color associations learnt by non-synesthete adults.

We study the dynamics of perceptual switching in ambiguous visual scenes that admit more than two interpretations/percepts to gain insight into the dynamics of perceptual multistability and its underlying neural mechanisms. We focus on visual plaids that are tristable and we present both experimental and computational results. We develop a firing-rate model based on mutual inhibition and adaptation that involves stochastic dynamics of multiple-attractor systems. The model can account for the dynamic properties (transition probabilities, distributions of percept durations, etc.) observed in the experiments. Noise and adaptation have both been shown to play roles in the dynamics of bistable perception. Here, tristable perception allows us to specify the roles of noise and adaptation in our model. Noise is critical in considering the time of a switch. On the other hand, adaptation mechanisms are critical in considering perceptual choice (in tristable perception, each time a percept ends, there is a possible choice between two new percepts).

It is often assumed that the action of cortical feedback connections is slow and modulatory, whereas feedforward connections carry a rapid drive to their target neurons. Recent results from our laboratory showed a very rapid effect of feedback connections on the visual responses of neurons in lower order areas. We wanted to determine whether such a rapid action is mediated by fast conducting axons. Using electrical stimulation, we compared the conduction velocities along feedforward and feedback axons between areas V1 and V2 of the macaque monkey. We conclude that feedback and feedforward connections between V1 and V2 have comparable fast conduction velocities (around 3.5 m/s).

Object. In this paper the authors report on the conception and adjustment of a microelectrode used to obtain unitary recordings in the human spinal cord. Methods. To overcome the difficulties related to intraoperative pulsations of the spinal cord, the authors opted to use a floating microelectrode. Because the recordings are obtained most often from spontaneous activities, it is difficult, with a single microelectrode, to separate spikes from electrical artifacts that are related to the switching of devices. Consequently, the authors designed a dual microelectrode made of two tungsten-in-glass—attached microelectrodes separated by 300 µm. Because the two electrodes cannot obtain recordings in the same neuron, it is possible to distinguish unambiguously spikes (recorded on one tip) from electrical artifacts (recorded simultaneously on the two tips). The dual microelectrode is 2 cm long, with a 20-µm tip length, and 800 to 1200—Ohms impedance. This microelectrode can be implanted “free hand,” in the dorsal horn, by using a microsurgical forceps under a surgical microscope. The data analysis is performed off-line with spike sorter hardware. In the dorsal horns in 17 patients who were selected to undergo a dorsal root entry zone (DREZ) rhizotomy to treat various pathological conditions, unitary recordings were obtained using this double microelectrode. The authors recorded 57 neurons in good conditions of stability and isolation. Conclusions. The microelectrode described in this paper was successfully used to obtain recordings in neurons in more than 85% of the patients. This simplified, floating double microelectrode can therefore be considered for use in microsurgical DREZ rhizotomy to obtain unitary recordings in the human spinal dorsal horn.

Inactivation by GABA is a powerful tool for studying the function of specific cortical regions. It is especially useful in electrophysiology, because inactivation is reversible within short time periods, and because the extent of the inactivated region can be accurately controlled. Iontophoresis of GABA inactivates neurons up to 300 microm around the micropipette. Pressure injection of GABA inactivates neurons further away, but the spatial and temporal characteristics of inactivation by this method have been poorly studied. In order to address this question, we built devices made of micropipettes and microelectrodes glued at various distances. We experienced that repetition of small injections of 100 mM GABA inactivate cortex in a more homogenous way than bolus injections. Diffusion of GABA after pressure injection does not seem to follow a point spread diffusion model as in the case of iontophoresis: GABA probably goes up along the micropipette shaft, and the volume of inactivation has an ellipsoidal form. In order to precisely determine the extent of the inactivated region, we built a mathematical model to fit the experimental data of inactivations obtained above and below the pipette tip. The model provides estimates of the inactivated region for volumes smaller than 60 nl of GABA 100 mM. Limits of inactivation are between 250 and 500 microm lateral to the tip of the pipette. The geometry of inactivation is difficult to predict beyond 60 nl and it seems hazardous to try to inactivate neurons beyond 800 microm with pressure injections of GABA 100 mM.

International audience; : The sense of agency depends on some internal cues that derive from action control, as well as external cues like contextual information and prior information (degree of contingency between an action and is effect). We assessed whether external agency cues are combined with internal agency cues to affect the sense of agency. In two experiments participants performed a movement (button press) that elicited, after a varying delay, an effect (ball appearing on a screen), and reported their sense of agency over the effect (full, partial or no-agency) while internal cues (premotor information) and external cues (contextual and prior information) were manipulated. We assessed the effect of agency cues on the delays at which the sense of agency varied. The delays were increased with premotor signals but were decreased with contextual information. These findings favour a model of integration of internal and external agency cues over time.

International audience; A fundamental question in the field of synesthesia is whether it is associated with other cognitive phenomena. The current study examined synesthesia's connections with phenomenal traits of mirror-touch and ticker tape experiences, as well as the representation of the three phenomena in the population, across gender and domain of work/study. Mirror-touch is the automatic, involuntary experience of tactile sensation on one's own body when others are being touched. For example, seeing another person's arm being stroked can evoke physical touch sensation on one's own arm. Ticker tape is the automatic visualization of spoken words or thoughts, such as a teleprompter. For example, when spoken to, a ticker taper might see mentally the spoken words displayed in front of his face or as coming out of the speaker's mouth. To explore synesthesia's associations with these phenomena, a diverse group (n = 3743) was systematically recruited from eight universities and one public museum in France to complete an online screening. Of the 1017 eligible respondents, synesthetes (across all subtypes) reported higher rates of mirror-touch and ticker tape than non-synesthetes, suggesting that synesthesia is associated with these phenomenal traits. However, effect sizes were small and we could not rule out that response bias influenced these associations. Mirror-touch and ticker tape were independent. No differences were found across gender or domain of work and study in prevalence of synesthesia, mirror-touch or ticker tape. The prevalence of ticker tape, unknown so far, was estimated at about 7%, an intermediate rate between estimates of grapheme-color (2-4%) and sequence-space synesthesia (9-14%). Within synesthesia, grapheme-personification, also called ordinal-linguistic personification (OLP) was the most common subtype and was estimated around 12%. Co-occurences of the different types of synesthesia were higher than chance, though at the level of small effect sizes.

International audience; We are usually unaware of the brief but large illumination changes caused by blinks, presumably because of blink suppression mechanisms. In fMRI however, increase of the BOLD signal was reported in the visual cortex, e.g. during blocks of voluntary blinks (Bristow, Frith and Rees, 2005) or after spontaneous blinks recorded during the prolonged fixation of a static stimulus (Tse, Baumgartner and Greenlee, 2010). We tested whether such activation, possibly related to illumination changes, was also present during standard fMRI retinotopic and visual experiments and was large enough to contaminate the BOLD signal we are interested in. We monitored in a 3T scanner the eyeblinks of 14 subjects who observed three different types of visual stimuli, including periodic rotating wedges and contracting/expanding rings, event-related Mondrians and graphemes, while fixating. We performed event-related analyses on the set of detected spontaneous blinks. We observed large and widespread BOLD responses related to blinks in the visual cortex of every subject and whatever the visual stimulus. The magnitude of the modulation was comparable to visual stimulation. However, blink-related activations lay mostly in the anterior parts of retinotopic visual areas, coding the periphery of the visual field well beyond the extent of our stimuli. Blinks therefore represent an important source of BOLD variations in the visual cortex and a troublesome source of noise since any correlation, even weak, between the distribution of blinks and a tested protocol could trigger artifactual activities. However, the typical signature of blinks along the anterior calcarine and the parieto-occipital sulcus allows identifying, even in the absence of eyetracking, fMRI protocols possibly contaminated by a heterogeneous distribution of blinks.

International audience; Auditory streaming and visual plaids have been used extensively to study perceptual organization in each modality. Both stimuli can produce bistable alternations between grouped (one object) and split (two objects) interpretations. They also share two peculiar features: (i) at the onset of stimulus presentation, organization starts with a systematic bias towards the grouped interpretation; (ii) this first percept has 'inertia'; it lasts longer than the subsequent ones. As a result, the probability of forming different objects builds up over time, a landmark of both behavioural and neurophysiological data on auditory streaming. Here we show that first percept bias and inertia are independent. In plaid perception, inertia is due to a depth ordering ambiguity in the transparent (split) interpretation that makes plaid perception tristable rather than bistable: experimental manipulations removing the depth ambiguity suppressed inertia. However, the first percept bias persisted. We attempted a similar manipulation for auditory streaming by introducing level differences between streams, to bias which stream would appear in the perceptual foreground. Here both inertia and first percept bias persisted. We thus argue that the critical common feature of the onset of perceptual organization is the grouping bias, which may be related to the transition from temporally/spatially local to temporally/spatially global computation.

International audience; This paper introduces the basic phenomenon of multistability, which occurs when a single physical stimulus produces alternations between different subjective percepts. The phenomenon was first discovered when different images were presented to the two eyes; the perceived image was dominated at any one time by the image presented to one eye, but the percept "flipped" unpredictably, an effect called binocular rivalry. Multistability was later discovered to occur for a wide range of visual stimuli, including those involving apparent motion, and to occur for auditory, tactile and olfactory stimuli. The key features of stimuli leading to multistability are: (i) they have more than one plausible perceptual organisation; (ii) these organisations are not compatible with each other. Most cases of multistability are based on competition in selecting and binding stimulus information. Hence, multistability can be used as a method for studying the binding process in various perceptual modalities. The binding issue is considered here in three different directions: binding sensory information into perceptual scenes for defining perceptual objects in each modality; binding sensory inputs across modalities for addressing questions about multisensory perceptual organisation in the brain; and binding our knowledge of neural processing in various sensory modalities to let emerge new questions, new methods and new models. The paper concludes with an outline of the papers in this issue and with some thoughts about open directions and avenues for further research.

International audience; The subjective experience of color by synesthetes when viewing achromatic letters and numbers supposedly relates to real color experience, as exemplified by the recruitment of the V4 color center observed in some brain imaging studies. Phenomenological reports and psychophysics tests indicate, however, that both experiences are different. Using functional magnetic resonance imaging, we tried to precise the degree of coactivation by real and synesthetic colors, by evaluating each color center individually, and applying adaptation protocols across real and synesthetic colors. We also looked for structural differences between synesthetes and nonsynesthetes. In 10 synesthetes, we found that color areas and retinotopic areas were not activated by synesthetic colors, whatever the strength of synesthetic associations measured objectively for each subject. Voxel-based morphometry revealed no white matter (WM) or gray matter difference in those regions when compared with 25 control subjects. But synesthetes had more WM in the retrosplenial cortex bilaterally. The joint coding of real and synesthetic colors, if it exists, must therefore be distributed rather than localized in the visual cortex. Alternatively, the key to synesthetic color experience might not lie in the color system.

What is the cause of perceptual switching of ambiguous stimuli? Although local competition between stimulus-specific representations can explain bistable perception dynamics (1), Einhauser et al. (2) argue that norepinephrine (NE) release in the locus coeruleus (a nonperceptual structure) could play a critical role in perceptual bistability. They reported an NE-mediated increase in pupil diameter coinciding with perceptual switching. Remarkably, the relative amount of dilation before switching was a weak but significant predictor of the duration of the subsequent percept. We questioned these results in two independent experiments on bistable perception (3, 4). Using Einhauser et al.'s method (2), we replicated both results. However, we contest that the positive correlation between pupil dilation and relative percept duration is predictive, because both measures are not independent of each other: For percepts shorter than 3 s (in our dataset, median = 1,840 ms, and n = 940), switch-related pupil dilation can last until the next switch. Meanwhile, given the overall distribution of percept durations, we observed that the next percept lasted longer (median = 4,095 ms, and t = −30). We verified that removing these short percepts completely abolished the positive correlation. We conclude that this correlation, which is not reliable across subjects, is most likely an artifact. However, pupil response around perceptual switching is intriguing. We conducted a series of experiments to dissect the perceptual and motor components of this response, and we observed a large influence of the motor command (3), contrary to the suggestion made by Einhauser et al. (2).

International audience; Ambiguous stimuli can produce spontaneous perceptual alternations in the mind of the observer, even though the stimulus itself remains the same. Common features in the temporal dynamics of bistability have been observed for various types of stimuli, both visual and auditory. This raises the question of whether bistable perception results from stereotyped, local competition between stimulus-specific representations or whether it is triggered by some central, supramodal mechanism. We tested the distributed versus centralized hypothesis by asking observers to simultaneously monitor their bistable perception of ambiguous auditory and visual stimuli. Strong interactions between auditory and visual perceptual switches would indicate a central decision mechanism. We used streaming stimuli in the auditory modality and either plaids or apparent motion stimuli in the visual modality. The use of two different sensory modalities allowed the distinction of contextual interactions due to the similarity between stimuli from interactions linked to perceptual decision itself. The longterm dynamics of bistable perception were identical in unimodal and bimodal presentations for all types of stimuli. Surprisingly, even strong short-term cross-modal interactions, when present, did not alter these dynamics. We conclude that bistability can co-occur independently in different sensory modalities. This observation supports models of distributed competition for perceptual decision and awareness.

Plaids are ambiguous stimuli that can be perceived either as a coherent pattern moving rigidly or as two gratings sliding over each other. Here we report a new factor that affects the relative strength of coherency versus transparency: the global direction of motion of the plaid. Plaids moving in oblique directions are perceived as sliding more frequently than plaids moving in cardinal directions. We term this the oblique plaid effect. There is also a difference between the two cardinal directions: for most observers, plaids moving in horizontal directions cohere more than plaids moving in vertical directions. Two measures were used to quantify the relative strength of coherency vs. transparency: C/[C + T ] and RTtransp. Those measures were derived from dynamics data obtained in longduration trials (>1 min) where observers continually indicated their percept. The perception of plaids is bi-stable: over time it alternates between coherency and transparency, and the dynamics data reveal the relative strength of the two interpretations [Vision Research 43 (2003) 531]. C/[C + T ] is the relative cumulative time spent perceiving coherency; RTtransp is the time between stimulus onset and the first report of transparency. The dynamics-based measures quantify the relative strength of coherency over a wider range of parameters than brief-presentation 2AFC methods, and exposed an oblique plaid effect in the entire range tested. There was no interaction between the effect of the global direction of motion and the effect of gratings’ orientations. Thus, the oblique plaid effect is due to anisotropies inherent to motion mechanisms, not a bi-product of orientation anisotropies. The strong effect of a plaid’s global direction on its tendency to cohere imposes new and important constraints on models of motion integration and transparency. Models that rely solely on relative differences in directions and/or orientations in the stimulus cannot predict our results. Instead, models should take into account anisotropies in the neuronal populations that represent the coherent percept (integrated motion) and those that represent the transparent percept (segmented motion). Furthermore, the oblique plaid effect could be used to test whether neuronal populations supposed to be involved in plaid perception display tuning biases in favor of cardinal directions. � 2003 Elsevier Ltd. All rights reserved.

We analyzed the extracellular responses of 70 V1 neurons (recorded in 3 anesthetized macaque monkeys) to a single oriented line segment (or bar) placed within the cell classical receptive field (RF), or center of the RF. These responses could be modulated when rings of bars were placed entirely outside, but around the RF (the “near” surround region), as described in previous studies. Suppression was the main effect. The response was enhanced for 12 neurons when orthogonal bars in the surround were presented instead of bars having the same orientation as the center bar. This orientation contrast property is possibly involved in the mediation of perceptual pop-out. The enhancement was delayed compared with the onset of the response by about 40 ms. We also observed a suppression originating specifically from the flanks of the surround. This “side-inhibition,” significant for nine neurons, was delayed by about 20 ms. We tested whether these center/surround interactions in V1 depend on feedback connections from area V2. V2 was inactivated by GABA injections. We used devices made of six micropipettes to inactivate the convergent zone from V2 to V1. We could reliably inactivate a 2- to 4-mm-wide region of V2. Inactivation of V2 had no effect on the center/surround interactions of V1 neurons, even those that were delayed. Therefore the center/surround interactions of V1 neurons that might be involved in pop-out do not appear to depend on feedback connections from V2, at least in the anesthetized monkey. We conclude that these properties are probably shaped by long-range connections within V1 or depend on other feedback connections. The main effect of V2 inactivation was a decrease of the response to the single bar for about 10% of V1 neurons. The decrease was delayed by

We previously showed that feedback connections from MT play a role in figure/ground segmentation. Figure/ground coding has been described at the V1 level in the late part of the neuronal responses to visual stimuli, and it has been suggested that these late modulations depend on feedback connections. In the present work we tested whether it actually takes time for this information to be fed back to lower order areas. We analyzed the extracellular responses of 169 V1, V2, and V3 neurons that we recorded in two anesthetized macaque monkeys. MT was inactivated by cooling. We studied the time course of the responses of the neurons that were significantly affected by the inactivation of MT to see whether the effects were delayed relative to the onset of the response. We first measured the time course of the feedback influences from MT on V1, V2, and V3 neurons tested with moving stimuli. For the large majority of the 51 neurons for which the response decreased, the effect was present from the beginning of the response. In the responses averaged after normalization, the decrease of response was significant in the first 10-ms bin of response. A similar result was found for six neurons for which the response significantly increased when MT was inactivated. We then looked at the time course of the responses to flashed stimuli (95 neurons). We observed 15 significant decreases of response and 14 significant increases. In both populations, the effects were significant within the first 10 ms of response. For some neurons with increased responses we even observed a shorter latency when MT was inactivated. We measured the latency of the response to the flashed stimuli. We found that even the earliest responding neurons were affected early by the feedback from MT. This was true for the response to flashed and to moving stimuli. These results show that feedback connections are recruited very early for the treatment of visual information. It further indicates that the presence or absence of feedback effects cannot be deduced from the time course of the response modulations.

The results of a previous study [Hupe et al. (1998) Nature, 394: 784–787] led us to conclude that feedback connections are important for differentiating a figure from the background, particularly in the case of low salience stimuli. This conclusion was principally based on the observation in area V3 neurons that inactivating MT by cooling led to a severe weakening of the center response and of the center-surround interactions, and that these effects were particularly strong for low salience stimuli. In the present paper, we first show that the results extend to areas V1 and V2. In particular, the inhibitory center-surround interactions in areas V1, V2 and V3 disappear almost completely in the absence of feedback input from MT for low salience stimuli, whereas the effects are much more limited for stimuli of middle and high salience. We then compare the results obtained in studies of feedback connections from MT to those obtained in a study of the feedback action of area V2 onto V1 neurons [Hupe et al. (2001) J. Neurophysiol., 85: 146–163], in which the same effects were observed on the center mechanism (decrease in response), but no effects were seen on the center-surround interactions. We conclude that feedback connections act in a non-linear fashion to boost the gain of the center mechanism and that they combine with horizontal connections to generate the center-surround interactions.

A single visual stimulus activates neurons in many different cortical areas. A major challenge in cortical physiology is to understand how the neural activity in these numerous active zones leads to a unified percept of the visual scene. The anatomical basis for these interactions is the dense network of connections that link the visual areas. Within this network, feedforward connections transmit signals from lower-order areas such as V1 or V2 to higher-order areas. In addition, there is a dense web of feedback connections which, despite their anatomical prominence1,2,3,4, remain functionally mysterious5,6,7,8. Here we show, using reversible inactivation of a higher-order area (monkey area V5/MT), that feedback connections serve to amplify and focus activity of neurons in lower-order areas, and that they are important in the differentiation of figure from ground, particularly in the case of stimuli of low visibility. More specifically, we show that feedback connections facilitate responses to objects moving within the classical receptive field; enhance suppression evoked by background stimuli in the surrounding region; and have the strongest effects for stimuli of low salience.

The role of feedback connections from area V2 to V1 was studied by reversible inactivation. When V2 was inactivated the responses of some V1 neurons to stimulation of the surround region were increased while responses to center stimulation were unchanged or decreased. Latencies to small flashing stimuli were also compared in areas V1 and V2. The distributions in the two areas overlap largely, with a 10 ms shift between the two. Neurons of V1 and V2 that are driven by the magnocellular layers of the LGN are activated 20 ms earlier than neurons of the parvocellular stream.

When observers view for extended time an ambiguous visual scene with two or more different interpretations they report switching between different perceptions. We focus on a classical paradigmatic stimulus, the visual plaids, consisting of two superimposed drifting gratings with transparent intersections [1,2]. For visual plaids, tristable perception is experienced: one coherent percept (the gratings move together as a single pattern) and two transparent percepts (the gratings slide across one another) with alternating depth order [3]. In order to decipher the complex mechanisms of tristable perception, we gathered a large amount of psychophysical data on tristable plaids and developed a neural network, firing rate model of interaction between neural populations that could account for the experimental results. Nine subjects reported continuously the three possible percepts for 3 sessions of 10 stimuli each (3 minutes per stimulus). The angle between the vectors normal to gratings was equal to 80, 100 or 120 degrees. We collected enough percepts to compute statistics for each subject and parameter without collapsing data [4]. As opposed to bistable stimuli where the only possibility is the alternation between the two percepts, in the tristable case the results show that the next percept probability depends on the previous percepts. Indeed, the sequence of perceptual switches confirms that switches between two transparency states are typically interleaved by a coherent percept, especially for values of the angle equal to 80 and 100 [3,4]. Moreover, by examining triplets consisting of two transparent percepts interleaved by a coherent one, we observed that the probability of the two transparent percepts in the triplet having the same depth pattern decreases as the duration of the coherent percept shortens. These trends suggest that adaptation is implicated in perceptual alternations. For bistable alternations correlations are absent or insignificant. We propose inhibition-based competition along with adaptation and noise in a multi-state framework as plausible mechanisms for the dynamics of perceptual switching. Our model is a firing rate model consisting of three mutually coupled populations of cells, each one encoding a different percept. It is based on the firing rate models for alternations during perceptual bistability [5]. We can explain the dependence in perceptual history by introducing an inhibition imbalance in the interactions between neural populations (the two transparent percepts inhibit each other more strongly than they inhibit the coherent state, making the latter more dominant and more likely to switch to). Adjusting the relative strength of adaptation and noise we can account for the dominance duration distributions and the switching probability between depth percepts as a function of the coherent percept duration. Finally, we consider other possible architectures for the model and we show that a non-hierarchical architecture where motion is encoded together with depth fits better with the experimental results.

International audience; Pupil size not only varies to changes in illumination but is also modulated by several cognitive factors, making it a potentially versatile physiological marker of cortical states. We recorded pupil dynamics while subjects continuously reported their bistable perception of ambiguous moving stimuli, plaids, and partially occluded rotating diamonds. We observed small (about 5% of surface change on average) but reliable pupil dilation around (-300 ms to 1.5 s) the button presses indicating the changes of percepts. We found that 70% of pupil dilation could be accounted for by the motor response. The remaining perceptual component was similar for spontaneously occurring transitions and transitions triggered by physical stimulus manipulations. Moreover, the amplitude of pupil modulation in the spontaneous condition was unrelated to the duration of each perceptual state. It is therefore unlikely that the mechanisms of endogenous perceptual bistability reflect in the pupil. In addition, we measured a clear constriction of the pupil after blinks (about 8% of surface change on average). As pupil changes have the potential to entail retino-cortical activity, their monitoring in studies of visual processing could prove worthwhile.