Ventral Stream Research Articles

A concentration of visual cortex-like neurons in prefrontal cortex

Visual recognition is largely realized through neurons in the ventral stream, though recently, studies have suggested that ventrolateral prefrontal cortex (vlPFC) is also important for visual processing. While it is hypothesized that sensory and cognitive processes are integrated in vlPFC neurons, it is not clear how this mechanism benefits vision, or even if vlPFC neurons have properties essential for computations in visual cortex implemented via recurrence. Here, we investigated if vlPFC neurons in two male monkeys had functions comparable to visual cortex, including receptive fields, image selectivity, and the capacity to synthesize highly activating stimuli using generative networks. We found a subset of vlPFC sites show all properties, suggesting subpopulations of vlPFC neurons encode statistics about the world. Further, these vlPFC sites may be anatomically clustered, consistent with fMRI-identified functional organization. Our findings suggest that stable visual encoding in vlPFC may be a necessary condition for local and brain-wide computations.

Exploring perceptual grouping by proximity principle in multistable dot lattices: Dissociation between vision-for-perception and vision-for-action.

Perceptual grouping, a fundamental mechanism in our visual system, significantly influences our interpretation of and interaction with the surrounding world. This study explores the impact of the proximity principle from the perspective of the Two Visual Systems (TVS) model. The TVS model argues that the visual system comprises two distinct streams: the ventral stream, which forms the neural basis for "vision-for-perception," and the dorsal stream, which underlies "vision-for-action." We designed a perceptual grouping task using dot lattices as well as a line-orientation discrimination task. Data were collected using vocal and mouse methods for the vision-for-perception mode, and joystick and pen-paper methods for the vision-for-action mode. Each method, except for vocal, included separate blocks for right and left hands. The proximity data were fitted using exponential and power models. Linear mixed-effects models were used for the statistical analyses. The results revealed similar line-orientation discrimination accuracy across all conditions. The exponential model emerged as the best fit, demonstrating adherence to the Pure Distance Law in both perceptual modes. Sensitivity to the proximity principle was higher in the vision-for-action mode compared to the vision-for-perception. In terms of orientation biases, a strong preference for vertical orientation was observed in the vision-for-perception mode, whereas a noticeable preference toward either of the oblique orientations was detected in the vision-for-action mode. Analysis of free-drawn lines demonstrated an affordance bias in the vision-for-action mode. This suggests a remarkable tendency to perceive organizations within specific orientations that offer more affordances due to the interaction between the body postures and tools.

Visual system structural and functional connections during face viewing in body dysmorphic disorder.

Individuals with body dysmorphic disorder (BDD) perceive distortions in their appearance, which could be due to imbalances in global and local visual processing. The vertical occipital fasciculus connects dorsal and ventral visual stream regions, integrating global and local information, yet the role of this structural connection in BDD has not been explored. Here, we investigated the vertical occipital fasciculus's white matter microstructure in those with BDD and healthy controls and tested associations with psychometric measures and effective connectivity while viewing their face during fMRI. We analyzed diffusion MRI and fMRI data in 17 unmedicated adults with BDD and 21 healthy controls. For diffusion MRI, bundle-specific analysis was performed, enabling quantitative estimation of neurite density and orientation dispersion of the vertical occipital fasciculus. For task fMRI, participants naturalistically viewed photos of their own face, from which we computed effective connectivity from dorsal to ventral visual regions. In BDD, neurite density was negatively correlated with appearance dissatisfaction and negatively correlated with effective connectivity. Further, those with weaker effective connectivity while viewing their face had worse BDD symptoms and worse insight. In controls, no significant relationships were found between any of the measures. There were no significant group differences in neurite density or orientation dispersion. Those with BDD with worse appearance dissatisfaction have a lower fraction of tissue having axons or dendrites along the vertical occipital fasciculus bundle, possibly reflecting impacting the degree of integration of global and local visual information between the dorsal and ventral visual streams. These results provide early insights into how the vertical occipital fasciculus's microstructure relates to the subjective experience of one's appearance, as well as the possibility of distinct functional-structural relationships in BDD.

Interaction between the prefrontal and visual cortices supports subjective fear.

It has been reported that threatening and non-threatening visual stimuli can be distinguished based on the multi-voxel patterns of haemodynamic activity in the human ventral visual stream. Do these findings mean that there may be evolutionarily hardwired mechanisms within early perception, for the fast and automatic detection of threat, and maybe even for the generation of the subjective experience of fear? In this human neuroimaging study, we presented participants ('fear' group: N = 30; 'no fear' group: N = 30) with 2700 images of animals that could trigger subjective fear or not as a function of theindividual's idiosyncratic 'fear profiles' (i.e. fear ratings of animals reported by a given participant). We provide evidence that the ventral visual stream may represent affectively neutral visual features that are statistically associated with fear ratings of participants, without representing the subjective experience of fear itself. More specifically, we show that patterns of haemodynamic activity predictive of a specific 'fear profile' can be observed in the ventral visual stream whether a participant reports being afraid of the stimuli or not. Further, we found that the multivariate information synchronization between ventral visual areas and prefrontal regions distinguished participants who reported being subjectively afraid of the stimuli from those who did not. Together, these findings support the view that the subjective experience of fear may depend on the relevant visual information triggering implicit metacognitive mechanisms in the prefrontal cortex. This article is part of the theme issue 'Sensing and feeling: an integrative approach to sensory processing and emotional experience'.

Dissonant music engages early visual processing

The neuroscientific examination of music processing in audio-visual contexts offers a valuable framework to assess how auditory information influences the emotional encoding of visual information. Using fMRI during naturalistic film viewing, we investigated the neural mechanisms underlying the effect of music on valence inferences during mental state attribution. Thirty-eight participants watched the same short-film accompanied by systematically controlled consonant or dissonant music. Subjects were instructed to think about the main character's intentions. The results revealed that increasing levels of dissonance led to more negatively valenced inferences, displaying the profound emotional impact of musical dissonance. Crucially, at the neuroscientific level and despite music being the sole manipulation, dissonance evoked the response of the primary visual cortex (V1). Functional/effective connectivity analysis showed a stronger coupling between the auditory ventral stream (AVS) and V1 in response to tonal dissonance and demonstrated the modulation of early visual processing via top-down feedback inputs from the AVS to V1. These V1 signal changes indicate the influence of high-level contextual representations associated with tonal dissonance on early visual cortices, serving to facilitate the emotional interpretation of visual information. Our results highlight the significance of employing systematically controlled music, which can isolate emotional valence from the arousal dimension, to elucidate the brain's sound-to-meaning interface and its distributive crossmodal effects on early visual encoding during naturalistic film viewing.

Factorized visual representations in the primate visual system and deep neural networks.

Object classification has been proposed as a principal objective of the primate ventral visual stream and has been used as an optimization target for deep neural network models (DNNs) of the visual system. However, visual brain areas represent many different types of information, and optimizing for classification of object identity alone does not constrain how other information may be encoded in visual representations. Information about different scene parameters may be discarded altogether ('invariance'), represented in non-interfering subspaces of population activity ('factorization') or encoded in an entangled fashion. In this work, we provide evidence that factorization is a normative principle of biological visual representations. In the monkey ventral visual hierarchy, we found that factorization of object pose and background information from object identity increased in higher-level regions and strongly contributed to improving object identity decoding performance. We then conducted a large-scale analysis of factorization of individual scene parameters - lighting, background, camera viewpoint, and object pose - in a diverse library of DNN models of the visual system. Models which best matched neural, fMRI, and behavioral data from both monkeys and humans across 12 datasets tended to be those which factorized scene parameters most strongly. Notably, invariance to these parameters was not as consistently associated with matches to neural and behavioral data, suggesting that maintaining non-class information in factorized activity subspaces is often preferred to dropping it altogether. Thus, we propose that factorization of visual scene information is a widely used strategy in brains and DNN models thereof.

Four dimensions of naturalistic language production in aphasia after stroke.

There is a rich tradition of research on the neuroanatomical correlates of spoken language production in aphasia using constrained tasks (e.g., picture naming), which offer controlled insights into the distinct processes that govern speech and language (i.e., lexical-semantic access, morphosyntactic construction, phonological encoding, speech motor programming/execution). Yet these tasks do not necessarily reflect everyday language use. In contrast, naturalistic language production (also referred to as connected speech or discourse) more closely approximates typical processing demands, requiring the dynamic integration of all aspects of speech and language. The brain bases of naturalistic language production remain relatively unknown, however, in part because of the difficulty in deriving features that are salient, quantifiable, and interpretable relative to both speech-language processes and the extant literature. The present cross-sectional observational study seeks to address these challenges by leveraging a validated and comprehensive auditory-perceptual measurement system that yields four explanatory dimensions of performance-Paraphasia (misselection of words and sounds), Logopenia (paucity of words), Agrammatism (grammatical omissions), and Motor speech (impaired speech motor programming/execution). We used this system to characterize naturalistic language production in a large and representative sample of individuals with acute post-stroke aphasia (n = 118). Scores on each of the four dimensions were correlated with lesion metrics, and multivariate associations among the dimensions and brain regions were then explored. Our findings revealed distinct yet overlapping neuroanatomical correlates throughout the left-hemisphere language network. Paraphasia and Logopenia were associated primarily with posterior regions, spanning both dorsal and ventral streams, which are critical for lexical-semantic access and phonological encoding. In contrast, Agrammatism and Motor speech were associated primarily with anterior regions of the dorsal stream that are involved in morphosyntactic construction and speech motor planning/execution respectively. Collectively, we view these results as constituting a brain-behavior model of naturalistic language production in aphasia, aligning with both historical and contemporary accounts of the neurobiology of spoken language production.

Neural correlates of musical timbre: an ALE meta-analysis of neuroimaging data.

Timbre is a central aspect of music that allows listeners to identify musical sounds and conveys musical emotion, but also allows for the recognition of actions and is an important structuring property of music. The former functions are known to be implemented in a ventral auditory stream in processing musical timbre. While the latter functions are commonly attributed to areas in a dorsal auditory processing stream in other musical domains, its involvement in musical timbre processing is so far unknown. To investigate if musical timbre processing involves both dorsal and ventral auditory pathways, we carried out an activation likelihood estimation (ALE) meta-analysis of 18 experiments from 17 published neuroimaging studies on musical timbre perception. We identified consistent activations in Brodmann areas (BA) 41, 42, and 22 in the bilateral transverse temporal gyri, the posterior superior temporal gyri and planum temporale, in BA 40 of the bilateral inferior parietal lobe, in BA 13 in the bilateral posterior Insula, and in BA 13 and 22 in the right anterior insula and superior temporal gyrus. The vast majority of the identified regions are associated with the dorsal and ventral auditory processing streams. We therefore propose to frame the processing of musical timbre in a dual-stream model. Moreover, the regions activated in processing timbre show similarities to the brain regions involved in processing several other fundamental aspects of music, indicating possible shared neural bases of musical timbre and other musical domains.

Relationship Between Resting State Functional Connectivity and Reading-Related Behavioural Measures in 69 Adults

Abstract In computational models of reading, written words can be read using print-to-sound and/or print-to-meaning pathways. Neuroimaging data associate dorsal stream regions (left posterior occipitotemporal cortex, intraparietal cortex, dorsal inferior frontal gyrus [dIFG]) with the print-to-sound pathway and ventral stream regions (left anterior fusiform gyrus, middle temporal gyrus) with the print-to-meaning pathway. In 69 typical adults, we investigated whether resting state functional connectivity (RSFC) between the visual word form area (VWFA) and dorsal and ventral regions correlated with phonological (nonword reading, nonword repetition, spoonerisms), lexical-semantic (vocabulary, sensitivity to morpheme units in reading), and general literacy (word reading, spelling) skills. VWFA activity was temporally correlated with activity in both dorsal and ventral reading regions. In pre-registered whole-brain analyses, spoonerisms performance was positively correlated with RSFC between the VWFA and left dorsal regions (dIFG, superior parietal and intraparietal cortex). In exploratory region-of-interest analyses, VWFA-dIFG connectivity was also positively correlated with nonword repetition, spelling, and vocabulary. Connectivity between the VWFA and ventral stream regions was not associated with performance on any behavioural measure, either in whole-brain or region-of-interest analyses. Our results suggest that tasks such as spoonerisms and spellings, which are both complex (i.e., involve multiple subprocesses) and have high between-subject variability, provide greater opportunity for observing resting-state brain-behaviour associations. However, the complexity of these tasks limits the conclusions we can draw about the specific mechanisms that drive these associations. Future research would benefit from constructing latent variables from multiple tasks tapping the same reading subprocess.

Cholinergic deficiency in Parkinson's disease patients with visual hallucinations.

Visual hallucinations can increase the burden of disease for both patients with Parkinson's disease and their caregivers. Multiple neurotransmitters have been implicated in the neuropathology of visual hallucinations, which provide targets for treatment and prevention. In this study, we assessed the association between cholinergic denervation and visual hallucinations in Parkinson's disease in vivo, using PET imaging of the cholinergic system. A total of 38 patients with Parkinson's disease participated in this study. A group of 10 healthy subjects, matched for age, sex and education, was included for comparison. None of the participants used cholinergic drugs. Thirteen patients who had experienced visual hallucinations in the past month (VH+) were compared with 20 patients who had never experienced visual hallucinations in their lives (VH-). Cholinergic system integrity was assessed with PET imaging using 18F-fluoroethoxybenzovesamicol as the tracer. We assessed the differences in tracer uptake between groups by cluster-based analysis and by analysis of predefined regions of interest consisting of the ventral visual stream, the dorsal attentional network, the ventral attentional network and the lateral geniculate nucleus and mediodorsal nucleus of the thalamus. The Parkinson's disease group (n = 38) showed an extensive pattern of decreased tracer uptake throughout the brain compared with the controls (n = 10). Within the Parkinson's disease group, the VH+ group (n = 13) showed a cluster of decreased tracer uptake compared with the VH- group (n = 20), which covered most of the left ventral visual stream and extended towards superior temporal areas. These results were mirrored in the regions of interest-based analysis, in which the VH+ group showed the strongest deficits in the left inferior temporal gyrus and the left superior temporal gyrus compared with the VH- group. Visual hallucinations in Parkinson's disease are associated with a marked cholinergic deficiency in the left ventral visual stream and the left superior temporal lobe, in addition to an extensive global cholinergic denervation in the general Parkinson's disease population.

Functional and Effective Connectivity Underlying Semantic Verbal Fluency.

Semantic verbal fluency (SVF) impairment is present in several neurological disorders. Although activation in SVF-related areas has been reported, how these regions are connected and their functional roles in the network remain divergent. We assessed SVF static and dynamic functional connectivity (FC) and effective connectivity in healthy participants using functional magnetic resonance imaging. We observed activation in the inferior frontal (IFG), middle temporal (pMTG) and angular gyri (AG), anterior cingulate (AC), insular cortex, and regions of the superior, middle, and medial frontal gyri (SFG, MFG, MidFG). Our static FC analysis showed a highly interconnected task and resting state network. Increased connectivity of AC with the pMTG and AG was observed for the task. The dynamic FC analysis provided circuits with connections similarly modulated across time and regions related to category identification, language comprehension, word selection and recovery, word generation, inhibition of speaking, speech planning, and articulatory planning of orofacial movements. Finally, the effective connectivity analysis provided a network that best explained our data, starting at the AG and going to the pMTG, from which there was a division between the ventral and dorsal streams. The SFG and MFG regions were connected and modulated by the MidFG, while the inferior regions formed the ventral stream. Therefore, we successfully assessed the SVF network, exploring regions associated with the entire processing, from category identification to word generation. The methodological approach can be helpful for further investigation of the SVF network in neurological disorders.

Selective activations and functional connectivities to the sight of faces, scenes, body parts and tools in visual and non-visual cortical regions leading to the human hippocampus

Connectivity maps are now available for the 360 cortical regions in the Human Connectome Project Multimodal Parcellation atlas. Here we add function to these maps by measuring selective fMRI activations and functional connectivity increases to stationary visual stimuli of faces, scenes, body parts and tools from 956 HCP participants. Faces activate regions in the ventrolateral visual cortical stream (FFC), in the superior temporal sulcus (STS) visual stream for face and head motion; and inferior parietal visual (PGi) and somatosensory (PF) regions. Scenes activate ventromedial visual stream VMV and PHA regions in the parahippocampal scene area; medial (7m) and lateral parietal (PGp) regions; and the reward-related medial orbitofrontal cortex. Body parts activate the inferior temporal cortex object regions (TE1p, TE2p); but also visual motion regions (MT, MST, FST); and the inferior parietal visual (PGi, PGs) and somatosensory (PF) regions; and the unpleasant-related lateral orbitofrontal cortex. Tools activate an intermediate ventral stream area (VMV3, VVC, PHA3); visual motion regions (FST); somatosensory (1, 2); and auditory (A4, A5) cortical regions. The findings add function to cortical connectivity maps; and show how stationary visual stimuli activate other cortical regions related to their associations, including visual motion, somatosensory, auditory, semantic, and orbitofrontal cortex value-related, regions.

Selective Neural Entrainment Reveals Hierarchical Tuning to Linguistic Regularities in Reading.

Reading is both a visual and a linguistic task, and as such it relies on both general-purpose, visual mechanisms and more abstract, meaning-oriented processes. Disentangling the roles of these resources is of paramount importance in reading research. The present study capitalizes on the coupling of fast periodic visual stimulation and MEG recordings to address this issue and investigate the role of different kinds of visual and linguistic units in the visual word identification system. We compared strings of pseudo-characters; strings of consonants (e.g., sfcl); readable, but unattested strings (e.g., amsi); frequent, but non-meaningful chunks (e.g., idge); suffixes (e.g., ment); and words (e.g., vibe); and looked for discrimination responses with a particular focus on the ventral, occipito-temporal regions. The results revealed sensitivity to alphabetic, readable, familiar, and lexical stimuli. Interestingly, there was no discrimination between suffixes and equally frequent, but meaningless endings, thus highlighting a lack of sensitivity to semantics. Taken together, the data suggest that the visual word identification system, at least in its early processing stages, is particularly tuned to form-based regularities, most likely reflecting its reliance on general-purpose, statistical learning mechanisms that are a core feature of the visual system as implemented in the ventral stream.

Beyond ventral and dorsal streams: thalamo-cortical connections for subcortical language integration.

Beyond ventral and dorsal streams: thalamo-cortical connections for subcortical language integration.

The attentive reconstruction of objects facilitates robust object recognition.

Humans are extremely robust in our ability to perceive and recognize objects-we see faces in tea stains and can recognize friends on dark streets. Yet, neurocomputational models of primate object recognition have focused on the initial feed-forward pass of processing through the ventral stream and less on the top-down feedback that likely underlies robust object perception and recognition. Aligned with the generative approach, we propose that the visual system actively facilitates recognition by reconstructing the object hypothesized to be in the image. Top-down attention then uses this reconstruction as a template to bias feedforward processing to align with the most plausible object hypothesis. Building on auto-encoder neural networks, our model makes detailed hypotheses about the appearance and location of the candidate objects in the image by reconstructing a complete object representation from potentially incomplete visual input due to noise and occlusion. The model then leverages the best object reconstruction, measured by reconstruction error, to direct the bottom-up process of selectively routing low-level features, a top-down biasing that captures a core function of attention. We evaluated our model using the MNIST-C (handwritten digits under corruptions) and ImageNet-C (real-world objects under corruptions) datasets. Not only did our model achieve superior performance on these challenging tasks designed to approximate real-world noise and occlusion viewing conditions, but also better accounted for human behavioral reaction times and error patterns than a standard feedforward Convolutional Neural Network. Our model suggests that a complete understanding of object perception and recognition requires integrating top-down and attention feedback, which we propose is an object reconstruction.

Neuronal tuning and population representations of shape and category in human visual cortex

Object recognition and categorization are essential cognitive processes which engage considerable neural resources in the human ventral visual stream. However, the tuning properties of human ventral stream neurons for object shape and category are virtually unknown. We performed large-scale recordings of spiking activity in human Lateral Occipital Complex in response to stimuli in which the shape dimension was dissociated from the category dimension. Consistent with studies in nonhuman primates, the neuronal representations were primarily shape-based, although we also observed category-like encoding for images of animals. Surprisingly, linear decoders could reliably classify stimulus category even in data sets that were entirely shape-based. In addition, many recording sites showed an interaction between shape and category tuning. These results represent a detailed study on shape and category coding at the neuronal level in the human ventral visual stream, furnishing essential evidence that reconciles human imaging and macaque single-cell studies.

Predicting high-level visual areas in the absence of task fMRI.

The ventral visual stream is organized into units, or functional regions of interest (fROIs), specialized for processing high-level visual categories. Task-based fMRI scans ("localizers") are typically used to identify each individual's nuanced set of fROIs. The unique landscape of an individual's functional activation may rely in large part on their specialized connectivity patterns; recent studies corroborate this by showing that connectivity can predict individual differences in neural responses. We focus on the ventral visual stream and ask: how well can an individual's resting state functional connectivity localize their fROIs for face, body, scene, and object perception? And are the neural processors for any particular visual category better predicted by connectivity than others, suggesting a tighter mechanistic relationship between connectivity and function? We found, among 18 fROIs predicted from connectivity for each subject, all but one were selective for their preferred visual category. Defining an individual's fROIs based on their connectivity patterns yielded regions that were more selective than regions identified from previous studies or atlases in nearly all cases. Overall, we found that in the absence of a domain-specific localizer task, a 10-min resting state scan can be reliably used for defining these fROIs.

Visuospatial processing in early brain-based visual impairment is associated with differential recruitment of dorsal and ventral visual streams.

Visuospatial processing impairments are prevalent in individuals with cerebral visual impairment (CVI) and are typically ascribed to "dorsal stream dysfunction" (DSD). However, the contribution of other cortical regions, including early visual cortex (EVC), frontal cortex, or the ventral visual stream, to such impairments remains unknown. Thus, here, we examined fMRI activity in these regions, while individuals with CVI (and neurotypicals) performed a visual search task within a dynamic naturalistic scene. First, behavioral performance was measured with eye tracking. Participants were instructed to search and follow a walking human target. CVI participants took significantly longer to find the target, and their eye gaze patterns were less accurate and less precise. Second, we used the same task in the MRI scanner. Along the dorsal stream, activation was reduced in CVI participants, consistent with the proposed DSD in CVI. Intriguingly, however, visual areas along the ventral stream showed the complete opposite pattern, with greater activation in CVI participants. In contrast, we found no differences in either EVC or frontal cortex between groups. These results suggest that the impaired visuospatial processing abilities in CVI are associated with differential recruitment of the dorsal and ventral visual streams, likely resulting from impaired selective attention.

Neural computations in prosopagnosia.

We report an investigation of the neural processes involved in the processing of faces and objects of brain-lesioned patient PS, a well-documented case of pure acquired prosopagnosia. We gathered a substantial dataset of high-density electrophysiological recordings from both PS and neurotypicals. Using representational similarity analysis, we produced time-resolved brain representations in a format that facilitates direct comparisons across time points, different individuals, and computational models. To understand how the lesions in PS's ventral stream affect the temporal evolution of her brain representations, we computed the temporal generalization of her brain representations. We uncovered that PS's early brain representations exhibit an unusual similarity to later representations, implying an excessive generalization of early visual patterns. To reveal the underlying computational deficits, we correlated PS' brain representations with those of deep neural networks (DNN). We found that the computations underlying PS' brain activity bore a closer resemblance to early layers of a visual DNN than those of controls. However, the brain representations in neurotypicals became more akin to those of the later layers of the model compared to PS. We confirmed PS's deficits in high-level brain representations by demonstrating that her brain representations exhibited less similarity with those of a DNN of semantics.

Modulation of the neuronal response in human primary visual cortex by re-entrant projections during retinal input processing as manifest in the visual evoked potential

Initial deflections in the visual evoked potential (VEP) reflect the neuronal process of extracting features from the retinal input; a process not modulated by re-entrant projections. Later deflections in the VEP reflect the neuronal process of combining features into an object, a process referred to as ‘object closure’ and modulated by re-entrant projections. Our earlier work indicated that the VEP reflects independent neuronal responses processing temporal – and spatial luminance contrast and that these responses arise from an interaction between forward and re-entrant input. In this earlier work, changing the temporal luminance contrast property of a stimulus altered its spatial luminance contrast property. We recorded the VEP in 12 volunteers viewing image pairs of a windmill, regular dartboard or an RMS dartboard rotated by either Π/4, Π/2, 3Π/4 or Π radians with respect to each other. The windmill and regular dartboard had identical white to black ratio, while the two dartboards identical contrast edges per unit area. Rotation varied temporal luminance contrast of a stimulus without affecting its spatial luminance contrast. N75, P100, N135 and P240 amplitude and latency were compared and a source localisation and temporal frequency analysis performed. P100 amplitude signals a neuronal response processing temporal luminance contrast that is modulated by re-entrant projections with fast axonal conduction velocities. N135 and P240 signal the neuronal response processing spatial luminance contrast and is modulated by re-entrant projections with slow axonal conduction velocities. The dorsal stream is interconnected by fast axonal conduction velocities, the ventral stream by slow axonal conduction velocities.