TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.     

TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.     

Selective visual attention and perceptual coherence

Conscious perception of the visual world depends on neural activity at all levels of the visual system from the retina to regions of parietal and frontal cortex. Neurons in early visual areas have small spatial receptive fields (RFs) and code basic image features; neurons in later areas have large RFs and code abstract features such as behavioral relevance. This hierarchical organization presents challenges to perception: objects compete when they are presented in a single RF, and component object features are coded by anatomically distributed neuronal activity. Recent research has shown that selective attention coordinates the activity of neurons to resolve competition and link distributed object representations. We refer to this ensemble activity as a "coherence field", and propose that voluntary shifts of attention are initiated by a transient control signal that "nudges" the visual system from one coherent state to another.     

大脑腹侧视觉通路知觉表征神经机制研究

大脑腹侧视觉通路知觉表征的神经机制是认知神经科学研究面临的基本问题。本文系统介绍了该问题研究中比较有影响力的理论模型,归纳分析了模型之间的分歧与各自的局限。文章分析指出大脑自上而下的调控机制是腹侧视觉通路神经表征机制问题研究的另一重要维度,如何有效整合知觉表征模型与大脑调控机制的相关研究是进一步深化知觉表征神经机制问题研究的关键。     

Do object-category selective regions in the ventral visual stream represent perceived distance information?

It is well established that scenes and objects elicit a highly selective response in specific brain regions in the ventral visual cortex. An inherent difference between these categories that has not been explored yet is their perceived distance from the observer (i.e. scenes are distal whereas objects are proximal). The current study aimed to test the extent to which scene and object selective areas are sensitive to perceived distance information independently from their category-selectivity and retinotopic location. We conducted two studies that used a distance illusion (i.e., the Ponzo lines) and showed that scene regions (the parahippocampal place area, PPA, and transverse occipital sulcus, TOS) are biased toward perceived distal stimuli, whereas the lateral occipital (LO) object region is biased toward perceived proximal stimuli. These results suggest that the ventral visual cortex plays a role in representing distance information, extending recent findings on the sensitivity of these regions to location information. More broadly, our findings imply that distance information is inherent to object recognition.     

The neural basis of mirror symmetry detection: a review

The human visual system is extremely sensitive to the presence of bilateral (mirror) symmetry. In this review, I summarise the results of recent work investigating the neural basis of mirror symmetry detection, focusing in particular on brain stimulation evidence. Overall, available findings converge in pointing to the lateral occipital (LO) complex, especially in the right hemisphere, as a key region causally involved in symmetry detection. Interestingly, they also suggest that another region in the right extrastriate visual cortex, the occipital face area (OFA), is causally implied in symmetry detection, posing an interesting connection at the neural level between visual cortex responses to faces and to symmetry. Finally, this review also considers evidence on haptic symmetry detection in sighted and early blind individuals that points to LO as a multi-modal symmetry-sensitive region, and suggests that symmetry is a salient perceptual feature mediated by LO even when any visual experience is missing.     

Age-related deficits in generation and manipulation of mental images: II. The role of dorsolateral prefrontal cortex.

The authors investigated neural substrates of age-related declines in mental imagery. Healthy adult participants (ages 19 to 77) performed a series of visual-spatial mental imagery tasks that varied in apparent difficulty and involved stimuli of varying graphic complexity. The volumes of the dorsolateral frontal cortex (DLPFC) and posterior visual processing areas were estimated from magnetic resonance imaging scans. The volume of the DLPFC and the fusiform cortex, working-memory capacity, and performance on the tasks involving image generation and manipulation were significantly reduced with age. Further analyses suggested that age-related deficits in performance on mental imagery tasks may stem in part from age-related shrinkage of the prefrontal cortex and age-related declines in working memory but not from age-related slowing of sensorimotor reaction time. The volume of cortical regions associated with modality-specific visual information processing did not show a consistent relationship with specific mental imagery processes.     

Cortical integration of audio–visual speech and non-speech stimuli

Using fMRI we investigated the neural basis of audio–visual processing of speech and non-speech stimuli using physically similar auditory stimuli (speech and sinusoidal tones) and visual stimuli (animated circles and ellipses). Relative to uni-modal stimuli, the different multi-modal stimuli showed increased activation in largely non-overlapping areas. Ellipse-Speech, which most resembles naturalistic audio–visual speech, showed higher activation in the right inferior frontal gyrus, fusiform gyri, left posterior superior temporal sulcus, and lateral occipital cortex. Circle-Tone, an arbitrary audio–visual pairing with no speech association, activated middle temporal gyri and lateral occipital cortex. Circle-Speech showed activation in lateral occipital cortex, and Ellipse-Tone did not show increased activation relative to uni-modal stimuli. Further analysis revealed that middle temporal regions, although identified as multi-modal only in the Circle-Tone condition, were more strongly active to Ellipse-Speech or Circle-Speech, but regions that were identified as multi-modal for Ellipse-Speech were always strongest for Ellipse-Speech. Our results suggest that combinations of auditory and visual stimuli may together be processed by different cortical networks, depending on the extent to which multi-modal speech or non-speech percepts are evoked.     

Multisensory cortical processing of object shape and its relation to mental imagery

Here, we used functional magnetic resonance imaging to investigate the multisensory processing of object shape in the human cerebral cortex and explored the role of mental imagery in such processing. Regions active bilaterally during both visual and haptic shape perception, relative to texture perception in the respective modality, included parts of the superior parietal gyrus, the anterior intraparietal sulcus, and the lateral occipital complex. Of these bimodal regions, the lateral occipital complexes preferred visual over haptic stimuli, whereas the parietal areas preferred haptic over visual stimuli. Whereas most subjects reported little haptic imagery during visual shape perception, experiences of visual imagery during haptic shape perception were common. Across subjects, ratings of the vividness of visual imagery strongly predicted the amount of haptic shape-selective activity in the right, but not in the left, lateral occipital complex. Thus, visual imagery appears to contribute to activation of some, but not all, visual cortical areas during haptic perception.     

Cognitive and neuronal processes involved in sequential generation of general and specific mental images

Mental image generation is a complex process mediated by dynamically interrelated components, e.g. image generation and image enrichment of details. This study investigated the cognitive and neural correlates of sequential image generation. An event-related fMRI experiment was carried out in which general and specific images had to be generated sequentially in two different positions. Participants had to generate either a general image first and then a specific one or a specific image first and then a general one, in response to the same word-stimulus. Generation times showed that specific images took shorter to be produced if they had been preceded by the generation of a general image. The fMRI results showed that position of generation and type of image was associated with different patterns of neurofunctional change. When an image was generated as first, areas of activation were found in the parahippocampal, fusiform and occipital regions. These are areas associated with memory retrieval and visual processing. When an image was generated as second, significant activations were found in superior temporal and precuneus areas, brain structures that are involved in the storage of visual memory for object shapes and imagery, respectively. The generation of a general image was supported by frontal areas and by the precuneus. The generation of a specific image involved frontal and thalamic areas (structures associated with visual processing of details) and the posterior cingulate cortex. When shifting from a specific image to a general one, a higher level of activity was found in the middle frontal gyrus involved in global visuo-spatial processing, suggesting that the generation of specific images required the retrieval of an object’s global shape. Altogether, these data suggest that the sequential generation of different types of image is associated with discrete processes but also shares common cognitive and neural components.     

The functions of the orbitofrontal cortex

The orbitofrontal cortex contains the secondary taste cortex, in which the reward value of taste is represented. It also contains the secondary and tertiary olfactory cortical areas, in which information about the identity and also about the reward value of odours is represented. The orbitofrontal cortex also receives information about the sight of objects from the temporal lobe cortical visual areas, and neurons in it learn and reverse the visual stimulus to which they respond when the association of the visual stimulus with a primary reinforcing stimulus (such as taste) is reversed. This is an example of stimulus-reinforcement association learning, and is a type of stimulus-stimulus association learning. More generally, the stimulus might be a visual or olfactory stimulus, and the primary (unlearned) positive or negative reinforcer a taste or touch. A somatosensory input is revealed by neurons that respond to the texture of food in the mouth, including a population that responds to the mouth feel of fat. In complementary neuroimaging studies in humans, it is being found that areas of the orbitofrontal cortex are activated by pleasant touch, by painful touch, by taste, by smell, and by more abstract reinforcers such as winning or losing money. Damage to the orbitofrontal cortex can impair the learning and reversal of stimulus-reinforcement associations, and thus the correction of behavioural responses when there are no longer appropriate because previous reinforcement contingencies change. The information which reaches the orbitofrontal cortex for these functions includes information about faces, and damage to the orbitofrontal cortex can impair face (and voice) expression identification. This evidence thus shows that the orbitofrontal cortex is involved in decoding and representing some primary reinforcers such as taste and touch; in learning and reversing associations of visual and other stimuli to these primary reinforcers; and in controlling and correcting reward-related and punishment-related behavior, and thus in emotion. The approach described here is aimed at providing a fundamental understanding of how the orbitofrontal cortex actually functions, and thus in how it is involved in motivational behavior such as feeding and drinking, in emotional behavior, and in social behavior.     

Separating figure from ground with a parallel network

The differentiation of figure from ground plays an important role in the perceptual organization of visual stimuli. The rapidity with which we can discriminate the inside from the outside of a figure suggests that at least this step in the process may be performed in visual cortex by a large number of neurons in several different areas working together in parallel. We have attempted to simulate this collective computation by designing a network of simple processing units that receives two types of information: bottom-up input from the image containing the outlines of a figure, which may be incomplete, and a top-down attentional input that biases one part of the image to be the inside of the figure. No presegmentation of the image was assumed. Two methods for performing the computation were explored: gradient descent, which seeks locally optimal states, and simulated annealing, which attempts to find globally optimal states by introducing noise into the computation. For complete outlines, gradient descent was faster, but the range of input parameters leading to successful performance was very narrow. In contrast, simulated annealing was more robust: it worked over a wider range of attention parameters and a wider range of outlines, including incomplete ones. Our network model is too simplified to serve as a model of human performance, but it does demonstrate that one global property of outlines can be computed through local interactions in a parallel network. Some features of the model, such as the role of noise in escaping from nonglobal optima, may generalize to more realistic models.     

Abnormal retinotopic representations in human visual cortex revealed by fMRI

The representation of the visual field in early visual areas is retinotopic. The point-to-point relationship on the retina is therefore maintained on the convoluted cortical surface. Functional magnetic resonance imaging (fMRI) has been able to demonstrate the retinotopic representation of the visual field in occipital cortex of normal subjects. Furthermore, visual areas that are retinotopic can be identified on computationally flattened cortical maps on the basis of positions of the vertical and horizontal meridians. Here, we investigate abnormal retinotopic representations in human visual cortex with fMRI. We present three case studies in which patients with visual disorders are investigated. We have tested a subject who only possesses operating rod photoreceptors. We find in this case that the cortex undergoes a remapping whereby regions that would normally represent central field locations now map more peripheral positions in the visual field: In a human albino we also find abnormal visual cortical activity. Monocular stimulation of each hemifield resulted in activations in the hemisphere contralateral to the stimulated eye. This is consistent with abnormal decussation at the optic chiasm in albinism. Finally, we report a case where a lesion to white matter has resulted in a lack of measurable activity in occipital cortex. The activity was absent for a small region of the visual field, which was found to correspond to the subject's field defect. The cases selected have been chosen to demonstrate the power of fMRI in identifying abnormalities in the cortical representations of the visual field in patients with visual dysfunction. Furthermore, the experiments are able to show how the cortex is capable of modifying the visual field representation in response to abnormal input.     

视觉工作记忆中注意模板的表征—来自EROS的证据

采用事件相关光信号技术,分析再认任务和搜索任务下脑区的动态激活,探讨视觉工作记忆中注意模板的表征机制。实验要求被试在记忆项目后,分别完成再认任务和搜索任务。结果发现,与再认任务相比,视觉搜索任务下记忆项诱发前额叶、枕叶更高程度的激活,且前额叶的激活增强早于枕叶。研究表明,与用于再认任务的普通记忆项相比,用于视觉搜索任务的注意模板的潜在表征机制可能是：先激活前额叶以增强注意控制,并对枕叶的活动加以调节,增强注意模板表征的复述。     

Toward a theory of visual consciousness.

The visual brain consists of several parallel, functionally specialized processing systems, each having several stages (nodes) which terminate their tasks at different times; consequently, simultaneously presented attributes are perceived at the same time if processed at the same node and at different times if processed by different nodes. Clinical evidence shows that these processing systems can act fairly autonomously. Damage restricted to one system compromises specifically the perception of the attribute that that system is specialized for; damage to a given node of a processing system that leaves earlier nodes intact results in a degraded perceptual capacity for the relevant attribute, which is directly related to the physiological capacities of the cells left intact by the damage. By contrast, a system that is spared when all others are damaged can function more or less normally. Moreover, internally created visual percepts-illusions, afterimages, imagery, and hallucinations-activate specifically the nodes specialized for the attribute perceived. Finally, anatomical evidence shows that there is no final integrator station in the brain, one which receives input from all visual areas; instead, each node has multiple outputs and no node is recipient only. Taken together, the above evidence leads us to propose that each node of a processing-perceptual system creates its own microconsciousness. We propose that, if any binding occurs to give us our integrated image of the visual world, it must be a binding between microconsciousnesses generated at different nodes. Since any two microconsciousnesses generated at any two nodes can be bound together, perceptual integration is not hierarchical, but parallel and postconscious. By contrast, the neural machinery conferring properties on those cells whose activity has a conscious correlate is hierarchical, and we refer to it as generative binding, to distinguish it from the binding that might occur between the microconsciousnesses.     

Transfer of visual information after unilateral input to the brain

Visual callosal connections are more numerous and widespread in the association areas than in the primary visual cortex and adjoining visual areas. In keeping with this, the amount of "wrong" ipsilateral visual field that is represented in the various cortical areas of primates and cats increases as one goes from primary visual cortex to extraoccipital areas. Therefore it can be argued that transfer of unilaterally presented visual stimuli occurs mainly at the temporal and parietal cortical level.     

Inter-individual variability in metacognitive ability for visuomotor performance and underlying brain structures

Metacognition refers to the ability to discriminate between one’s own correct and incorrect decisions. The neurobiological underpinnings of metacognition have mainly been studied in perceptual decision-making. Here we investigated whether differences in brain structure predict individual variability in metacognitive sensitivity for visuomotor performance. Participants had to draw straight trajectories toward visual targets, which could unpredictably deviate around detection threshold, report such deviations when detected, and rate their confidence level for such reports. Structural brain MRI analyses revealed that larger gray-matter volume (GMV) in the left middle occipital gyrus, left medial parietal cortex, and right postcentral gyrus predicted higher deviation detection sensitivity. By contrast, larger GMV in the right prefrontal cortex but also right anterior insula and right fusiform gyrus predicted higher metacognitive sensitivity. These results extend past research by linking metacognitive sensitivity for visuomotor behavior to brain areas involved in action agency (insula), executive control (prefrontal cortex) and vision (fusiform).     

Seeing in three dimensions: the neurophysiology of stereopsis

From the pair of 2-D images formed on the retinas, the brain is capable of synthesizing a rich 3-D representation of our visual surroundings. The horizontal separation of the two eyes gives rise to small positional differences, called binocular disparities, between corresponding features in the two retinal images. These disparities provide a powerful source of information about 3-D scene structure, and alone are sufficient for depth perception. How do visual cortical areas of the brain extract and process these small retinal disparities, and how is this information transformed into non-retinal coordinates useful for guiding action? Although neurons selective for binocular disparity have been found in several visual areas, the brain circuits that give rise to stereoscopic vision are not very well understood. I review recent electrophysiological studies that address four issues: the encoding of disparity at the first stages of binocular processing, the organization of disparity-selective neurons into topographic maps, the contributions of specific visual areas to different stereoscopic tasks, and the integration of binocular disparity and viewing-distance information to yield egocentric distance. Some of these studies combine traditional electrophysiology with psychophysical and computational approaches, and this convergence promises substantial future gains in our understanding of stereoscopic vision.