The Role of Unimodal Feedback Pathways in Gender Perception During Activation of Voice and Face Areas.

Cross-modal effects provide a model framework for investigating hierarchical inter-areal processing, particularly, under conditions where unimodal cortical areas receive contextual feedback from other modalities. Here, using complementary behavioral and brain imaging techniques, we investigated the functional networks participating in face and voice processing during gender perception, a high-level feature of voice and face perception. Within the framework of a signal detection decision model, Maximum likelihood conjoint measurement (MLCM) was used to estimate the contributions of the face and voice to gender comparisons between pairs of audio-visual stimuli in which the face and voice were independently modulated. Top-down contributions were varied by instructing participants to make judgments based on the gender of either the face, the voice or both modalities (N = 12 for each task). Estimated face and voice contributions to the judgments of the stimulus pairs were not independent; both contributed to all tasks, but their respective weights varied over a 40-fold range due to top-down influences. Models that best described the modal contributions required the inclusion of two different top-down interactions: (i) an interaction that depended on gender congruence across modalities (i.e., difference between face and voice modalities for each stimulus); (ii) an interaction that depended on the within modalities' gender magnitude. The significance of these interactions was task dependent. Specifically, gender congruence interaction was significant for the face and voice tasks while the gender magnitude interaction was significant for the face and stimulus tasks. Subsequently, we used the same stimuli and related tasks in a functional magnetic resonance imaging (fMRI) paradigm (N = 12) to explore the neural correlates of these perceptual processes, analyzed with Dynamic Causal Modeling (DCM) and Bayesian Model Selection. Results revealed changes in effective connectivity between the unimodal Fusiform Face Area (FFA) and Temporal Voice Area (TVA) in a fashion that paralleled the face and voice behavioral interactions observed in the psychophysical data. These findings explore the role in perception of multiple unimodal parallel feedback pathways.

Neural circuits for long-range color filling-in.

Surface color appearance depends on both local surface chromaticity and global context. How are these inter-dependencies supported by cortical networks? Combining functional imaging and psychophysics, we examined if color from long-range filling-in engages distinct pathways from responses caused by a field of uniform chromaticity. We find that color from filling-in is best classified and best correlated with appearance by two dorsal areas, V3A and V3B/KO. In contrast, a field of uniform chromaticity is best classified by ventral areas hV4 and LO. Dynamic causal modeling revealed feedback modulation from area V3A to areas V1 and LO for filling-in, contrasting with feedback from LO modulating areas V1 and V3A for a matched uniform chromaticity. These results indicate a dorsal stream role in color filling-in via feedback modulation of area V1 coupled with a cross-stream modulation of ventral areas suggesting that local and contextual influences on color appearance engage distinct neural networks.

Effects of background and contour luminance on the hue and brightness of the Watercolor effect.

Conjoint measurement was used to investigate the joint influences of the luminance of the background and the inner contour on hue- and brightness filling-in for a stimulus configuration generating a water-color effect (WCE), i.e., a wiggly bi-chromatic contour enclosing a region with the lower luminance component on the exterior. Two stimuli with the background and inner contour luminances covarying independently were successively presented, and in separate experiments, the observer judged which member of the pair's interior regions contained a stronger hue or was brighter. Braided-contour control stimuli that generated little or no perceptual filling-in were also used to assess whether observers were judging the interior regions and not the contours themselves. Three nested models of the contributions of the background and inner contour to the judgments were fit to the data by maximum likelihood and evaluated by likelihood ratio tests. Both stimulus components contributed to both the hue and brightness of the interior region with increasing luminance of the inner contour generating an assimilative filling-in for the hue judgments but a contrast effect for the brightness judgments. Control analyses showed negligible effects for the order of the luminance of the background or inner contour on the judgments. An additive contribution of both components was rejected in favor of a saturated model in which the responses depended on the levels of both stimulus components. For the hue judgments, increased background luminance led to greater hue filling-in at higher luminances of the interior contour. For the brightness judgments, the higher background luminance generated less brightness filling-in at higher luminances of the interior contour. The results indicate different effects of the inner contour and background on the induction of the brightness and coloration percepts of the WCE, suggesting that they are mediated by different mechanisms.

Increasing Attentional Load Boosts Saccadic Adaptation.

PURPOSE: Visual exploration relies on saccadic eye movements and attention processes. Saccadic adaptation mechanisms, which calibrate the oculomotor commands to continuously maintain the accuracy of saccades, have been suggested to act at downstream (motor) and upstream (visuoattentional) levels of visuomotor transformation. Conversely, whether attention can directly affect saccadic adaptation remains unknown. To answer this question, we manipulated the level of attention engaged in a visual discrimination task performed during saccadic adaptation. METHODS: Participants performed low or high attention demanding orientation discrimination tasks on largely or faintly oriented Gabor patches, respectively, which served as targets for reactive saccades. Gabor patches systematically jumped backward during eye motion to elicit an adaptive shortening of saccades, and replaced 50 msec later (100 msec in two subjects) by a mask. Subjects judged whether Gabors' orientation was "nearly horizontal" versus "nearly vertical" (low attention demanding) or "slightly left" versus "slightly right" (high attention demanding), or made no discrimination (control task). RESULTS: We found that the build-up and the retention of adaptation of reactive saccades were larger in the "high attention demanding" condition than in the "low attention demanding" and the no-discrimination control conditions. CONCLUSIONS: These results indicate that increasing the level of attention to the perceptual processing of otherwise identical targets boosts saccadic adaptation, and suggest that saccadic adaptation mechanisms and attentional load effects may functionally share common neural substrates.

Spatial selectivity of the watercolor effect.

The spatial selectivity of the watercolor effect (WCE) was assessed by measuring its strength as a function of the luminance contrast of its inducing contours for different spatial configurations, using a maximum likelihood scaling procedure. The approach has previously been demonstrated to provide an efficient method for investigating the WCE as well as other perceptual dimensions. We show that the strength is narrowly tuned to the width of the contour, that it is optimal when its pair of inducing contours are of equal width, and that the strength can be increased by varying the overall size of the stimulus when the width of the inducing contour is not optimal. The results support a neural substrate that has characteristics not unlike double-opponent, color-luminance cells observed in cortical area V1.

Contributions of contour frequency, amplitude, and luminance to the watercolor effect estimated by conjoint measurement.

The watercolor effect is a long-range, assimilative, filling-in phenomenon induced by a pair of distant, wavy contours of different chromaticities. Here, we measured joint influences of the contour frequency and amplitude and the luminance of the interior contour on the strength of the effect. Contour pairs, each enclosing a circular region, were presented with two of the dimensions varying independently across trials (luminance/frequency, luminance/amplitude, frequency/amplitude) in a conjoint measurement paradigm (Luce & Tukey, 1964). In each trial, observers judged which of the stimuli evoked the strongest fill-in color. Control stimuli were identical except that the contours were intertwined and generated little filling-in. Perceptual scales were estimated by a maximum likelihood method (Ho, Landy, & Maloney, 2008). An additive model accounted for the joint contributions of any pair of dimensions. As shown previously using difference scaling (Devinck & Knoblauch, 2012), the strength increases with luminance of the interior contour. The strength of the phenomenon was nearly independent of the amplitude of modulation of the contour but increased with its frequency up to an asymptotic level. On average, the strength of the effect was similar along a given dimension regardless of the other dimension with which it was paired, demonstrating consistency of the underlying estimated perceptual scales.

A role for the parietal cortex in sensorimotor adaptation of saccades.

Sensorimotor adaptation ensures movement accuracy despite continuously changing environment and body. Adaptation of saccadic eye movements is a classical model of sensorimotor adaptation. Beside the well-established role of the brainstem-cerebellum in the adaptation of reactive saccades (RSs), the cerebral cortex has been suggested to be involved in the adaptation of voluntary saccades (VSs). Here, we provide direct evidence for a causal involvement of the parietal cortex in saccadic adaptation. First, the posterior intraparietal sulcus (pIPS) was identified in each subject using functional magnetic resonance imaging (fMRI). Then, a saccadic adaptation paradigm was used to progressively reduce the amplitude of RSs and VSs, while single-pulse transcranial magnetic stimulation (spTMS) was applied over the right pIPS. The perturbations of pIPS resulted in impairment for the adaptation of VSs, selectively when spTMS was applied 60 ms after saccade onset. In contrast, the adaptation of RSs was facilitated by spTMS applied 90 ms after saccade initiation. The differential effect of spTMS relative to saccade types suggests a direct interference with pIPS activity for the VS adaptation and a remote interference with brainstem-cerebellum activity for the RS adaptation. These results support the hypothesis that the adaptation of VSs and RSs involves different neuronal substrates.

Functional activation of the cerebral cortex related to sensorimotor adaptation of reactive and voluntary saccades.

Potentially dangerous events in the environment evoke automatic ocular responses, called reactive saccades. Adaptation processes, which maintain saccade accuracy against various events (e.g. growth, aging, neuro-muscular lesions), are to date mostly relayed to cerebellar activity. Here we demonstrate that adaptation of reactive saccades also involves cerebral cortical areas. Moreover, we provide the first identification of the neural substrates of adaptation of voluntary saccades, representing the complement to reactive saccades for the active exploration of our environment. An fMRI approach was designed to isolate adaptation from saccade production: an adaptation condition in which the visual target stepped backward 50 ms after saccade termination was compared to a control condition where the same target backstep occurred 500 ms after saccade termination. Subjects were tested for reactive and voluntary saccades in separate sessions. Multi-voxel pattern analyses of fMRI data from previously-defined regions of interests (ROIs) significantly discriminated between adaptation and control conditions for several ROIs. Some of these areas were revealed for adaptation of both saccade categories (cerebellum, frontal cortex), whereas others were specifically related to reactive saccades (temporo-parietal junction, hMT+/V5) or to voluntary saccades (medial and posterior areas of intra-parietal sulcus). These findings critically extend our knowledge on brain motor plasticity by showing that saccadic adaptation relies on a hitherto unknown contribution of the cerebral cortex.

Integration of visual information for saccade production.

To foveate a visual target, subjects usually execute a primary hypometric saccade (S1) bringing the target in perifoveal vision, followed by a corrective saccade (S2) or by more than one S2. It is still debated to what extent these S2 are pre-programmed or dependent only on post-saccadic retinal error. To answer this question, we used a visually-triggered saccade task in which target position and target visibility were manipulated. In one-third of the trials, the target was slightly displaced at S1 onset (so-called double step paradigm) and was maintained until the end of S1, until the start of the first S2 or until the end of the trial. Experiments took place in two visual environments: in the dark and in a dimly lit room with a visible random square background. The results showed that S2 were less accurate for shortest target durations. The duration of post-saccadic visual integration thus appears as the main factor responsible for corrective saccade accuracy. We also found that the visual context modulates primary saccade accuracy, especially for the most hypometric subjects. These findings suggest that the saccadic system is sensitive to the visual properties of the environment and uses different strategies to maintain final gaze accuracy.

Prior knowledge of illumination for 3D perception in the human brain.

In perceiving 3D shape from ambiguous shading patterns, humans use the prior knowledge that the light is located above their head and slightly to the left. Although this observation has fascinated scientists and artists for a long time, the neural basis of this "light from above left" preference for the interpretation of 3D shape remains largely unexplored. Combining behavioral and functional MRI measurements coupled with multivoxel pattern analysis, we show that activations in early visual areas predict best the light source direction irrespective of the perceived shape, but activations in higher occipitotemporal and parietal areas predict better the perceived 3D shape irrespective of the light direction. These findings demonstrate that illumination is processed earlier than the representation of 3D shape in the visual system. In contrast to previous suggestions, we propose that prior knowledge about illumination is processed in a bottom-up manner and influences the interpretation of 3D structure at higher stages of processing.

Effects of motion and configural complexity on color transparency perception.

We tested whether motion and configural complexity affect perceived transparency. A series of five coherent chromatic transformations in color space was applied across a figure: translation, convergence, shear, divergence and rotation. The stimuli consisted of a bipartite or a checkerboard configuration (10 x 10 degrees), with a central static or moving overlay (5 x 5 degrees). Three different luminance conditions (the plane of chromatic transformation oriented toward higher, lower, or equal luminances) were also tested for each of three modulation depths. For each stimulus, the observer judged whether the overlay appeared transparent or not. The main results indicated an interaction between the type of chromatic transformation and stimulus motion and complexity. For example, convergences are judged to appear transparent significantly more often when motion is added for bipartite configurations, or when they are generated in a checkerboard configuration. Surprisingly, shears that have been reported to appear opaque, are more frequently reported to appear transparent with short vector lengths and when combined with motion. Other transformations are also affected by motion, although the effectiveness of figural complexity on transparency seems to depend on both the type of color shifts and the presence of motion. The results indicate that adding motion and stimulus complexity are not necessarily neutral with respect to the chromatic shifts evoking transparency. Thus, studies that have used motion to enhance transparency may yield different results about the color shifts supporting transparency perception from those that did not. The same might be supposed for stimulus complexity under some conditions.