Mark Jonathan Roberts

Theta-gamma phase-amplitude coupling (PAC) has been proposed as a mechanism for separating neural representations in time. Although item-specific PAC has been reported, its role in perception remains unclear. Notably, previous work has shown that theta phase can bias perception, raising the question whether such biases are linked to phase-specific gamma activity. Here, we investigated whether phase-dependent perceptual biases are accompanied by item-specific theta-gamma PAC. We presented ambiguous speech sounds that could be interpreted as either /da/ or /ga/ in patients with epilepsy implanted with intracranial electrocorticography. We found that pre-stimulus theta phase predicted both the neural and perceptual response, with /da/ and /ga/ tuned channels exhibiting gamma activity clustering at distinct theta phases. These findings suggest that neural and perceptual biases stem from phase-specific pre-activation of item-specific neural representations. They demonstrate the generalizability of the theta-gamma code, shaping behavior and enhancing representational capacity.

Filling-in occurs during the perceptual disappearance of a blank figure presented on a textured background. Current models of perceptual filling-in are based on a two-stage model where the figure boundary weakens after a period of adaptation, followed by the spreading of the background representation into the region representing the figure. This suggests a competition between figure boundary and background representations whereby filling-in is facilitated by a weaker boundary representation and a stronger background representation. Here, we test this interpretation, by using the oblique effect and surround-modulation suppression, which are functional properties of early visual cortex that modulate the expected strengths of the responses to the background texture and to the figure boundary. In a sample of N=58 participants, we found more filling-in with background textures of cardinal compared to oblique orientations (earlier onset time, with more and longer episodes of filling-in per trial), in line with a known, stronger neuronal response for cardinal than for oblique orientation in early visual cortex. We found more filling-in when the main axis of the rectangular figure was iso-oriented rather than cross-oriented with the background texture (more and longer episodes of filling-in per trial, but no change in onset time), in line with a lower response to oriented stimuli when surrounded by iso-oriented flankers compared to cross-oriented flankers. Overall, our results support the two-stage model and suggest the involvement of early visual cortical areas characterized by the oblique effect and orientation-tuned surround-suppression.

Visual surface perception is a fundamental aspect of vision, yet its neural implementation remains poorly understood. Troxler’s perceptual filling-in paradigm provides a tractable illusion for studying surface perception, in which a peripheral figure becomes perceptually assimilated into the surrounding background after a period of sustained fixation. Although neural correlates of this phenomenon have been reported in early visual cortex, the underlying mechanisms, particularly the contribution of feedback signaling, remain unresolved. Here we use ultra-high-field (7T) layer-fMRI to investigate perceptual filling-in in the human visual cortex. While experimentally controlling perceptual filling-in, we measured GE-BOLD responses in ten participants. Analyses across cortical depth in the independently localized figure representation in primary visual cortex (V1) revealed neural correlates of filling-in in deep cortical layers, which are associated with feedback input. These findings provide evidence that perceptual filling-in and visual surface perception in general are supported by feedback signals to early visual cortex.

Gamma synchrony is ubiquitous in visual cortex, but whether it contributes to perceptual grouping remains contentious based on observations that gamma frequency is not consistent across stimulus features and that gamma synchrony depends on distances between image elements. These stimulus dependencies have been argued to challenge the idea that the visual system groups image elements by synchronizing the neural assemblies that encode them. Here, we argue instead that these dependencies may shape synchrony in perceptually meaningful ways. Indeed, according to the theory of weakly coupled oscillators (TWCO), synchrony-based grouping mechanisms require stimulus dependence. Synchronization among coupled oscillators depends on frequency dissimilarity and coupling strength, which in early visual cortex relate to local feature dissimilarity and physical distance, respectively. We manipulated these factors in a texture segregation experiment wherein human observers identified the orientation of a figure defined by reduced contrast heterogeneity compared to the background. Human performance followed TWCO predictions both qualitatively and quantitatively, as formalized in a computational model. Moreover, we found that when enriched with a Hebbian learning rule, our model also predicted human learning effects: Increases in model gamma synchrony due to perceptual learning predicted improvements in texture segregation across sessions. Taken together, our data suggest that the stimulus-dependence of gamma synchrony captures local image statistics and is linked to the stimulus-dependence of texture segregation, and that the effect of visual experience on gamma synchrony provides a viable perceptual learning mechanism for training-induced improvements in texture segregation. Our results suggest that gamma synchrony with its inherent stimulus dependencies can provide a plausible mechanistic basis for perceptual grouping and visual scene segmentation.

Gamma synchrony is ubiquitous in visual cortex, but whether it contributes to perceptual grouping remains contentious based on observations that gamma frequency is not consistent across stimulus features and that gamma synchrony depends on distances between image elements. These stimulus dependencies have been argued to challenge the idea that the visual system groups image elements by synchronizing the neural assemblies that encode them. Here we argue instead that these dependencies may shape synchrony in perceptually meaningful ways. Indeed, according to the theory of weakly coupled oscillators (TWCO), synchrony-based grouping mechanisms require stimulus dependence. Synchronization among coupled oscillators depends on frequency dissimilarity and coupling strength, which in early visual cortex relate to local feature dissimilarity and physical distance, respectively. We manipulated these factors in a texture segregation experiment wherein human observers identified the orientation of a figure defined by reduced contrast heterogeneity compared to the background. Human performance followed TWCO predictions both qualitatively and quantitatively, as formalized in a computational model. Moreover, we found that when enriched with a Hebbian learning rule, our model also predicted human learning effects: Increases in model gamma synchrony due to perceptual learning predicted improvements in texture segregation across sessions. Taken together, our data suggest that the stimulus-dependence of gamma synchrony captures local image statistics and is linked to the stimulus-dependence of texture segregation, and that the effect of visual experience on gamma synchrony provides a viable perceptual learning mechanism for training-induced improvements in texture segregation. Our results suggest that gamma synchrony with its inherent stimulus dependencies can provide a plausible mechanistic basis for perceptual grouping and visual scene segmentation.

Prior EEG research has shown that during working memory, alpha (8 to 14 Hz) and theta (4 to 8 Hz) oscillations tend to form a 2:1 frequency ratio. According to the Binary Hierarchy Brain Body Oscillation Theory (BHBBOT), a recent model grounded in mathematical analysis, this cross-frequency configuration reflects enhanced connectivity between brain regions generating these rhythms. However, this prediction has not yet been empirically tested. In this study, we leveraged high density EEG, source localization and connectivity metrics derived from Information Theory (IT) and the Theory of Weakly Coupled Oscillators (TWCO) to examine whether the previously observed alpha-theta cross-frequency dynamics during working memory are accompanied by changes in connectivity. Our results show that a significant increase in the proportion of 2:1 ratios between regions generating frontal theta and parietal alpha rhythms was accompanied by relative decreases in connectivity, as revealed by both IT and TWCO metrics. Furthermore, phase synchrony between these two regions was significantly reduced during working memory and correlated negatively with behavioral performance. In conclusion, our results show that the increased occurrence of 2:1 alpha:theta cross-frequency ratios during working memory reflects functional segregation (rather than integration) and therefore directly challenges some of the predictions of the BHBBOT.

Previous studies using a visual texture discrimination task (TDT) have demonstrated that performance enhancements resulting from extensive daily training ( full training condition) remained intact after replacing all training, except for the first and last session, with a few daily reminder trials ( reminder condition). Omitting reminders ( control condition) yielded only limited learning, supporting their crucial contribution. We first confirmed these findings and excluded gaze position differences among conditions as a contributing factor. Next, we tested whether the reminders’ effectiveness is specific to a context of limited attention to the peripheral target caused by simultaneously performing a demanding fixation task. Removing the fixation task yielded performance levels in the first session matching those normally reached after lengthy daily training, suggesting that learning in the standard TDT involves the redeployment of attention. After changing texture parameters to increase the difficulty of the task, performing the TDT without a fixation task yielded learning in all three conditions. This indicates that in a dual-task, reminders can produce learning outcomes comparable to full training . In contrast, when the TDT is performed with full attention to the target, consolidation of the initial session alone can yield improvements equivalent to those observed in reminder and full training conditions.

Surface perception relies on interactions between boundary encoding and surface filling-in. In a perceptual filling-in paradigm, a blank figure becomes perceptually replaced by a textured background after prolonged fixation of a point away from the figure. Filling-in begins when neurons representing the figure’s boundary adapt, allowing background-related activity to spread into the figure’s retinotopic representation. Adaptation proceeds faster the better the boundary is stabilized in a neuron’s receptive field (RF). We hypothesized that moving the figure boundary beyond a neuron’s RF would reduce adaptation and hinder filling-in, with larger movements permitting progressively less filling-in. As RF size increases with eccentricity, we further hypothesized that greater eccentricities would require larger movements to interfere with filling-in. Our results confirmed both predictions. The reduction in filling-in duration with increasing motion range permitted estimating RF size at each eccentricity. The slope of a linear function relating RF size to eccentricity matched values reported in human fMRI studies of V1/V2, suggesting that boundary adaptation involves early visual areas. We also explored whether microsaccade amplitude affects filling-in, but found no supporting evidence. Thus, external figure motion and microsaccades may disrupt adaptation through different mechanisms. These findings provide new insights into neural adaptation processes preceding perceptual filling-in.

Gamma synchrony is ubiquitous in visual cortex, but whether it contributes to perceptual grouping remains contentious based on observations that gamma frequency is not consistent across stimulus features and that gamma synchrony depends on distances between image elements. These stimulus dependencies have been argued to challenge the idea that the visual system groups image elements by synchronizing the neural assemblies that encode them. Here we argue instead that these dependencies may shape synchrony in perceptually meaningful ways. Indeed, according to the theory of weakly coupled oscillators (TWCO), synchrony-based grouping mechanisms require stimulus dependence. Synchronization among coupled oscillators depends on frequency dissimilarity and coupling strength, which in early visual cortex relate to local feature dissimilarity and physical distance, respectively. We manipulated these factors in a texture segregation experiment wherein human observers identified the orientation of a figure defined by reduced contrast heterogeneity compared to the background. Human performance followed TWCO predictions both qualitatively and quantitatively, as formalized in a computational model. Moreover, we found that when enriched with a Hebbian learning rule, our model also predicted human learning effects: Increases in model gamma synchrony due to perceptual learning predicted improvements in texture segregation across sessions. Taken together, our data suggest that the stimulus-dependence of gamma synchrony captures local image statistics and is linked to the stimulus-dependence of texture segregation, and that the effect of visual experience on gamma synchrony provides a viable perceptual learning mechanism for training-induced improvements in texture segregation. Our results suggest that gamma synchrony with its inherent stimulus dependencies can provide a plausible mechanistic basis for perceptual grouping and visual scene segmentation.