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Two-streams hypothesis

The two-streams hypothesis, a foundational model in visual neuroscience, proposes that the primate brain processes visual information through two parallel cortical pathways originating from primary visual cortex: the ventral stream, which extends into the temporal lobe and supports object identification and perceptual representation (often termed the "what" pathway), and the dorsal stream, which projects to the parietal lobe and facilitates spatial awareness and visually guided actions (known as the "where" or "how" pathway).^[1]^[2] This dichotomy highlights functional specialization in the visual system, where perception and action are dissociated to enable efficient interaction with the environment.^[2] The hypothesis originated from lesion studies in monkeys conducted by Leslie G. Ungerleider and Mortimer Mishkin in the early 1980s, who observed that damage to the temporal lobe impaired object recognition but spared spatial abilities, while parietal lesions disrupted localization without affecting identification.^[1] Their work built on earlier neuroanatomical evidence of segregated projections from striate cortex (V1) to inferotemporal and posterior parietal regions, establishing the streams as distinct multisynaptic pathways.^[1] In 1992, Melvyn A. Goodale and A. David Milner refined the model based on human patient data, particularly from D.F., who suffered ventral stream damage yet could perform accurate actions on objects despite profound perceptual deficits, shifting emphasis from "where" to "how" for the dorsal pathway.^[2] Subsequent neuroimaging and electrophysiological studies in both monkeys and humans have corroborated the hypothesis, revealing hierarchical processing within each stream: the ventral pathway involves areas like V4 and inferotemporal cortex for feature integration and category-specific recognition, while the dorsal stream includes regions such as the lateral intraparietal area (LIP) and medial superior temporal area (MST) for motion and depth processing critical to motor control.^[3] Evidence from functional MRI shows stream-specific activation patterns during tasks requiring perception versus grasping, underscoring their independence yet interconnected nature through feedback loops.^[4] The two-streams model has profoundly influenced cognitive science, informing theories of visuomotor coordination, agnosia syndromes, and even computational vision algorithms that mimic biological separation of recognition and navigation.^[5] Challenges include debates over stream interactions and whether the dorsal pathway includes substreams for different action types, but it remains a cornerstone for understanding visual cortex organization.^[6]

Overview

Core Concepts

The two-streams hypothesis proposes that the brain processes sensory information, particularly visual input, through parallel cortical pathways that specialize in distinct functions. In its core formulation for vision, one pathway—the ventral stream—handles the identification and recognition of objects ("what" processing), while the other—the dorsal stream—manages spatial awareness and guidance of actions ("where" or "how" processing). This division allows for efficient, segregated handling of perceptual and motor demands, emerging from the initial convergence of visual signals in primary visual cortex before diverging into these streams.^[1] The hypothesis was originally articulated by Ungerleider and Mishkin in 1982, based on anatomical and behavioral evidence from rhesus monkeys, which demonstrated dissociable pathways projecting from occipital cortex to temporal and parietal regions, respectively.^[1] This model emphasized functional specialization, with the temporal pathway supporting object discrimination and the parietal pathway aiding visuospatial tasks. Subsequent refinement by Goodale and Milner in 1992 shifted the emphasis from purely spatial versus object processing to a sharper distinction between conscious visual perception (ventral stream) and unconscious, real-time visuomotor control (dorsal stream), drawing on neuropsychological cases of dissociated visual deficits.^[7] This functional dissociation underscores the hypothesis's explanatory power: the ventral stream enables deliberate, recognition-based perception that integrates with memory and language, whereas the dorsal stream operates rapidly and automatically to support immediate actions like grasping or navigation, often bypassing conscious awareness.^[8] Analogous parallel streams have been hypothesized for auditory processing, where ventral pathways focus on sound identification and dorsal ones on spatial localization and action integration, extending the model's principles beyond vision.

Anatomical Foundations

The two-streams hypothesis posits a segregation of visual processing into parallel cortical pathways originating from the primary visual cortex (V1), with distinct anatomical trajectories supported by neuroanatomical tracing studies in primates.^[7] This segregation begins early in the visual system, influenced by inputs from the lateral geniculate nucleus (LGN) of the thalamus, where magnocellular layers predominantly project to regions that feed the dorsal stream, emphasizing motion and spatial sensitivity, while parvocellular layers primarily innervate areas contributing to the ventral stream, focusing on color and form details. Although both streams receive mixed inputs, this differential emphasis from LGN layers 1-2 (magnocellular) and 3-6 (parvocellular) establishes the foundational parallel processing architecture. The ventral stream pathway follows an occipitotemporal route, progressing from V1 through the ventral portions of V2 and V4 to the inferotemporal cortex (IT), particularly areas TE and TEO. These connections form a hierarchical series of corticocortical projections, with V4 serving as a key intermediate station that receives color- and shape-selective inputs before relaying to IT, where neurons exhibit invariance to object position and size. Anatomical studies using anterograde tracers in monkeys confirm dense, reciprocal linkages along this path, underscoring its role as a segregated conduit for detailed visual analysis. In contrast, the dorsal stream pathway takes an occipitoparietal course, extending from V1 via the dorsal aspects of V2 and V3 to the motion-sensitive middle temporal area (MT/V5), and thence to the posterior parietal cortex (PPC), including regions such as the lateral intraparietal area (LIP) and ventral intraparietal area (VIP). This route is characterized by broader, more divergent projections, with MT receiving strong inputs from V1's thick stripes in V2 and projecting forward to parietal areas via area V3A. Tracer studies reveal that these connections maintain spatial and motion-biased selectivity, forming a parallel hierarchy distinct from the ventral pathway. While the streams operate in parallel from their early divergence at V1, evidence from connectivity mapping indicates limited cross-stream interactions at higher cortical levels, such as bidirectional projections between IT area TEO and PPC's LIP, which may allow for integration without fully merging the pathways.^[9] This anatomical arrangement preserves the hypothesis's core principle of functional specialization through segregated processing routes.^[7]

Historical Development

Origins in Visual Neuroscience

The foundations of the two-streams hypothesis emerged from mid-20th-century studies elucidating the functional organization of the primate visual system. In the 1960s, David Hubel and Torsten Wiesel conducted seminal electrophysiological recordings in the primary visual cortex (V1) of cats and monkeys, identifying specialized "feature detectors" such as simple cells responsive to oriented edges and complex cells integrating motion and directionality. Their work demonstrated a hierarchical progression from basic edge detection in V1 to more complex feature integration in higher areas, suggesting segregated pathways for processing different aspects of visual information. This laid critical groundwork for understanding how visual inputs diverge into specialized streams beyond V1. Preceding the formal hypothesis, lesion studies in the late 1960s provided early evidence for dual visual processing mechanisms. George Schneider's 1969 experiments on hamsters revealed that tectal lesions (targeting the superior colliculus) selectively impaired visuomotor orienting and localization, while lesions to the visual cortex disrupted pattern discrimination and object recognition without affecting spatial guidance. These dissociations indicated two anatomically and functionally distinct systems: one for reflexive localization via subcortical structures and another for perceptual analysis in cortical areas, influencing subsequent models of visual pathway segregation in mammals.^[10] The two-streams hypothesis was explicitly formulated in 1982 by Leslie G. Ungerleider and Mortimer Mishkin, building on these influences through targeted lesion experiments in rhesus monkeys. They proposed a ventral stream projecting from occipital cortex to the inferior temporal lobe for "what" processing—enabling object identification and form perception—and a dorsal stream to the posterior parietal cortex for "where" processing—supporting spatial localization and visually guided actions.^[1] Monkeys with temporal lesions exhibited deficits in recognizing objects across views or delays, whereas parietal lesions caused impairments in reaching toward or discriminating locations of stimuli, confirming the streams' dissociable roles. This primate-based framework began transitioning to human applications in the late 1980s with the emergence of positron emission tomography (PET) imaging, which enabled noninvasive mapping of visual cortex activation. Early PET studies, such as those demonstrating retinotopic organization in human V1 during visual stimulation, provided initial correlates for segregated processing pathways analogous to the monkey model. These findings marked the onset of functional neuroimaging's role in validating the hypothesis across species, highlighting occipital-parietal and occipital-temporal activations during spatial versus object tasks.^[11]

Key Experimental Milestones

In the early 1990s, a significant refinement to the two-streams hypothesis emerged from studies of patient D.F., who suffered from visual form agnosia due to damage in the ventral stream following carbon monoxide poisoning. D.F. exhibited intact visuomotor abilities, such as accurately grasping objects despite being unable to consciously perceive their shape or orientation, demonstrating a dissociation between perception and action.^[12] This evidence prompted A. David Milner and Melvyn A. Goodale to propose in their 1992 paper that the dorsal stream is specialized for "how" to interact with objects in real-time visuomotor control, rather than merely "where" they are located, building on earlier lesion studies while emphasizing functional independence between the streams; they elaborated on this in their 1995 book The Visual Brain in Action.^[2] Concurrent electrophysiological recordings in awake monkeys during the 1990s provided cellular-level validation of stream-specific processing. In the ventral stream, single-unit recordings from the inferotemporal cortex (IT) revealed neurons highly selective for complex object features, such as shapes and textures, supporting its role in object recognition independent of spatial location or action demands. Complementing this, recordings in the dorsal stream's middle temporal area (MT) confirmed sensitivity to motion direction and speed, while posterior parietal cortex (PPC) neurons, particularly in areas like the anterior intraparietal area (AIP), responded to visual cues for grasping and reaching, such as object size and orientation for preshaping the hand.^[13] Early functional magnetic resonance imaging (fMRI) studies in humans during the late 1990s and early 2000s further corroborated stream segregation by examining brain activation during perception versus action tasks. For instance, when participants viewed objects for identification (engaging the ventral stream), activation was prominent in lateral occipital complex regions, whereas grasping tasks elicited stronger responses in dorsal areas like the superior parietal lobule and intraparietal sulcus, with minimal ventral involvement. These patterns highlighted how the streams process visual information differently based on task demands, aligning with primate data and extending the hypothesis to human cognition.^[14]

Visual Streams

Ventral Stream: Perception and Object Recognition

The ventral stream, often referred to as the "what" pathway, plays a central role in visual perception by enabling the identification and categorization of objects through a hierarchical progression of neural processing. This pathway begins in the primary visual cortex (V1) and extends through secondary areas like V2, integrating basic features such as edges and orientations, before advancing to area V4, where more complex attributes like color, form, and texture are combined to form intermediate representations of object surfaces and shapes.^[15] In V4, neurons exhibit selectivity for contour fragments and surface properties, facilitating the parsing of object boundaries from backgrounds and contributing to the synthesis of coherent visual forms essential for recognition.^[16] This integration culminates in the inferior temporal (IT) cortex, where neurons achieve viewpoint- and size-invariant representations of entire objects, allowing for robust identification regardless of retinal position or transformations. Behavioral manifestations of ventral stream processing are evident in specialized tasks requiring object recognition. For instance, face recognition relies heavily on the fusiform face area (FFA) within the ventral occipitotemporal cortex, where neurons respond selectively to facial configurations, supporting the rapid discrimination of individual identities.^[17] Similarly, word reading engages the visual word form area (VWFA) in the left fusiform gyrus, a region tuned to orthographic strings and invariant to font or case variations, enabling efficient decoding of written language as a perceptual category.^[18] Scene categorization, in turn, activates the parahippocampal place area (PPA), which processes holistic spatial layouts and environmental contexts to distinguish indoor from outdoor settings or natural from urban scenes.^[19] The ventral stream functions as a slower, more deliberative pathway compared to its dorsal counterpart, with neural latencies typically ranging from 200 to 300 ms post-stimulus onset in human magnetoencephalography (MEG) recordings, reflecting the time required for hierarchical feature integration and recurrent processing that supports conscious awareness, semantic interpretation, and integration with long-term memory.^[20] This temporal profile allows the stream to contribute to enduring perceptual representations, such as linking object identities to stored knowledge, rather than immediate visuomotor responses.^[21]

Dorsal Stream: Action and Spatial Processing

The dorsal stream, extending from the primary visual cortex to the posterior parietal cortex (PPC), specializes in visuomotor transformations that enable egocentric spatial processing and the guidance of actions, such as localizing objects relative to the observer and coordinating movements in real time.^[22] This pathway supports the "how" aspect of vision by integrating dynamic visual cues to direct behaviors without relying on conscious object identification.^[22] Key inputs to the dorsal stream include motion signals, which facilitate rapid adjustments to changing environments. The dorsal stream comprises two main subdivisions: the dorsomedial stream and the dorsolateral stream, each contributing to distinct aspects of spatial and action processing.^[23] The dorsomedial stream, involving areas like V6A in the superior parietal lobule and the medial intraparietal area (MIP), focuses on the online control of reaching and grasping movements, integrating visual form and motion for arm trajectories.^[23] In contrast, the dorsolateral stream encompasses the anterior intraparietal area (AIP) for precise hand shaping during manipulation and the lateral intraparietal area (LIP) for broader spatial navigation and attention shifts.^[23] These subdivisions ensure segregated yet interconnected processing for complex visuomotor tasks. Central to dorsal stream function is real-time motion analysis in area MT/V5, which detects object trajectories and feeds into parietal regions for immediate action planning.^[24] The PPC then executes coordinate transformations, remapping retinotopic visual inputs into somatotopic frames for hand-eye coordination, allowing seamless alignment of gaze and limb movements.^[25] This process underpins behaviors like visually guided saccades, where LIP neurons encode target locations to direct rapid eye shifts toward salient stimuli.^[26] Behavioral evidence highlights the dorsal stream's role in practical actions, such as obstacle avoidance during reaching, where automatic trajectory corrections prevent collisions based on spatial layout rather than object details.^[27] Patients with dorsal stream lesions, like those exhibiting optic ataxia, fail to adapt hand paths around barriers, underscoring the pathway's necessity for implicit visuomotor guidance.^[27] Similarly, tool use demonstrates AIP's involvement in grip preshaping for functional manipulation—such as wielding a hammer—without explicit awareness of the tool's identity, as observed in individuals with ventral stream damage who perform actions accurately despite perceptual deficits.^[28]

Extensions to Auditory Processing

Auditory Ventral Stream

The auditory ventral stream represents the auditory counterpart to the visual ventral stream, emphasizing the identification and semantic interpretation of sounds rather than their spatial attributes. This pathway processes auditory "what" information, enabling the recognition of sound objects and their associated meanings, much like object recognition in the visual domain. Proposed as an extension of the two-streams hypothesis to audition, it highlights modality-specific adaptations for perceiving complex acoustic environments. The pathway originates in the primary auditory cortex (A1) and proceeds anteriorly through belt regions such as the middle lateral (ML) and anterolateral (AL) areas, then along the superior temporal gyrus (STG) toward the anterior temporal lobe and ventrolateral prefrontal cortex (vlPFC).^[29] This hierarchical progression allows for increasingly abstract representations, with early stages encoding basic acoustic features and later regions integrating contextual and semantic information. In humans, the stream projects ventrolaterally from anterior and middle STG to middle and inferior temporal cortices, facilitating interfaces between sound processing and higher cognitive networks.^[30]^[31] Core functions of the auditory ventral stream include speech recognition, environmental sound categorization, and phonological processing. In speech, the STG exhibits a gradient: middle regions handle phonemes, anterior-superior areas process words, and the most anterior parts manage phrases, supporting comprehension through invariant representations of temporally complex sounds.^[29] For environmental sounds, AL regions categorize stimuli like vocalizations despite acoustic variability, contributing to scene analysis.^[29] Phonological processing occurs early, with AL neurons showing categorical responses to speech contrasts, such as distinguishing consonants in ambiguous tokens.^[29] Overall, the stream maps acoustic input to lexical-semantic meanings, essential for language and auditory object identification.^[31] Evidence from a neuroimaging meta-analysis supports the ventral stream's role in "what" tasks, showing segregated activation patterns that align with processing specialization, with nonspatial (identity) tasks primarily activating temporal lobe regions and spatial (location) tasks activating parietal regions, indicating functional independence of the auditory ventral pathway in humans.^[32]

Auditory Dorsal Stream

The auditory dorsal stream, analogous to its visual counterpart, processes spatial aspects of sound, originating in the core auditory cortex including primary auditory cortex (A1) within Heschl's gyrus and extending through the lateral belt and parabelt regions of the superior temporal gyrus (STG).^[33] From these posterior superior temporal areas, the pathway projects dorsally to the inferior parietal lobule (IPL), particularly the intraparietal sulcus and planum temporale extensions, and further to frontal regions such as the premotor cortex and posterior inferior frontal gyrus.^[34] This fronto-parietal trajectory facilitates the integration of auditory spatial information with motor systems, distinguishing it from the ventral stream's focus on sound identification.^[35] Key functions of the auditory dorsal stream include sound localization, which relies on extracting binaural cues to determine a sound source's position in three-dimensional space. Neurons along this pathway, particularly in the posterior STG and IPL, are tuned to interaural time differences (ITDs) and interaural level differences (ILDs), enabling precise encoding of azimuth (horizontal position) relative to the listener's head.^[33] For elevation (vertical position), spectral cues from pinna filtering are processed in parietal regions, with human neuroimaging showing activation in the IPL during tasks involving vertical sound motion.^[36] Beyond localization, the dorsal stream supports auditory attention shifts and sensorimotor integration, guiding orienting responses such as head turning or eye movements toward sound sources. Functional MRI studies demonstrate IPL and premotor activation during tasks requiring spatial attention to sounds, linking auditory input to motor planning for actions like vocalization or gesture in response to auditory events.^[35] This sensorimotor role extends to coordinating auditory-motor mappings, as evidenced by primate electrophysiology where dorsal stream neurons respond to both sound location and associated movements.^[33]

Supporting Evidence

Neuroimaging and Functional Studies

Functional magnetic resonance imaging (fMRI) studies have provided robust evidence for the segregation of the ventral and dorsal visual streams in healthy human brains by demonstrating selective activations during tasks emphasizing perception versus action. For instance, when participants viewed objects for recognition purposes, ventral stream regions such as the lateral occipital complex (LOC) showed heightened activation, whereas dorsal stream areas like the anterior intraparietal sulcus (AIP) exhibited minimal response.^[37] In contrast, during visually guided grasping tasks with real 3D objects, AIP and other dorsal regions were preferentially activated, while LOC activity was suppressed, highlighting a functional dissociation between streams for object perception and manipulation.^[38] These findings from early 2000s research, including seminal work by Culham and colleagues, underscore how task demands can isolate stream-specific processing in non-invasive group studies.^[37] Electroencephalography (EEG) and magnetoencephalography (MEG) have further elucidated the temporal dynamics of the two streams through event-related potentials (ERPs) and fields, revealing differences in processing speed for motion and form features. Dorsal stream responses to dynamic visual stimuli, such as motion onset, emerge rapidly around 100 ms post-stimulus, as indexed by early positive components over posterior parietal electrodes, reflecting quick spatial and action-oriented computations.^[39] Ventral stream processing of static form and object identity, however, manifests later, typically 150-200 ms after stimulus presentation, with components like the N170 over occipitotemporal sites supporting perceptual categorization.^[39] These temporal distinctions, observed in healthy subjects during simple visual discrimination tasks, support the hypothesis of parallel but temporally offset pathways, with dorsal responses preceding ventral ones to facilitate rapid integration for behavior.^[40] Recent advancements in the 2020s, particularly resting-state fMRI, have revealed intrinsic connectivity patterns that delineate stream-specific networks while also indicating cross-modal influences. Analyses of spontaneous BOLD fluctuations show stronger within-network coherence in ventral regions (e.g., fusiform gyrus) linked to object semantics and in dorsal areas (e.g., superior parietal lobule) associated with spatial attention, confirming segregated functional architectures even without external stimuli.^[41] Moreover, these studies highlight inter-stream and cross-modal connections, such as between visual dorsal areas and auditory motion-sensitive regions, suggesting dynamic interactions that extend the two-streams model beyond isolated visual processing.^[42] Such connectivity profiles, derived from large-scale datasets, provide evidence for the hypothesis's applicability in understanding integrated sensory-motor functions in typical brains.^[43]

Lesion and Behavioral Evidence

Lesions to the ventral stream, which is associated with object recognition and perception, often result in visual agnosia, where patients exhibit profound deficits in identifying and recognizing visual forms despite intact basic visual abilities such as acuity and color vision. A seminal case is that of patient D.F., who suffered bilateral damage to the lateral occipital complex following carbon monoxide poisoning in 1988; she demonstrated visual form agnosia, performing at chance levels (around 50% accuracy) on tasks requiring explicit judgments of object orientation, width, or shape, such as matching the slant of a slot or the width of Efron blocks. However, D.F. showed preserved visuomotor abilities, accurately preshaping her hand grip to match the width or orientation of objects during grasping tasks, with maximum grip aperture scaling appropriately to object size (e.g., errors less than 5 mm deviation from controls). This dissociation provided key evidence for the functional independence of the ventral stream in conscious perception from the dorsal stream's role in action guidance.^[44] In contrast, lesions to the dorsal stream, typically from strokes or trauma affecting the superior parietal lobule and intraparietal sulcus, lead to optic ataxia, characterized by impaired visually guided reaching and pointing while object perception remains relatively intact. Patients with unilateral parietal lesions, such as those studied in cases of posterior parietal stroke, can accurately describe object features like size, shape, and location verbally or through perceptual matching tasks (e.g., achieving over 90% accuracy in object identification), but exhibit significant errors in manual actions, with reaching deviations up to 10-15 cm from target locations under visual guidance. For instance, in a series of patients with right parietal damage, visuomotor coordination was disrupted in the contralesional field, leading to misreaching and grasping inaccuracies, yet discrimination of the same stimuli in non-action tasks was unimpaired. These findings underscore the dorsal stream's specialized role in transforming visual information into spatial-motor coordinates for action. Behavioral paradigms further illustrate dorsal stream deficits through targeted experiments revealing dissociations in spatial and temporal processing. In Efron's delayed matching task, adapted for visual stimuli, patients with parietal lesions show impairments in judging the simultaneity of events occurring at different spatial locations, with thresholds for detecting asynchrony elevated by 50-100 ms compared to controls, while judgments of temporal order for colocalized stimuli remain intact. This deficit, observed in individuals with right posterior parietal damage, highlights the dorsal stream's involvement in integrating spatial attention with temporal resolution, as the task requires binding location-specific visual inputs over time without relying on object recognition. Such paradigms complement lesion studies by isolating dorsal functions in healthy and impaired populations alike.^[45]

Criticisms and Modern Perspectives

Challenges to Stream Independence

While the two-streams hypothesis posits a largely independent segregation of the ventral and dorsal visual pathways, neuroanatomical and functional studies have revealed substantial bidirectional interactions that challenge this modularity. Evidence from diffusion tensor imaging indicates the existence of white matter pathways, such as the vertical occipital fasciculus, that facilitate cross-talk between the streams, supporting the integration of object recognition (ventral) with spatial action guidance (dorsal). For instance, ventral stream feedback to dorsal areas enables object-specific grasping adjustments, as demonstrated in models accounting for neural dynamics during perception-action tasks where bidirectional connections best explain observed behaviors. Neuroimaging further supports this, showing ventral contributions to dorsal processing in tasks requiring object-guided actions, such as reaching, where perceptual details from the ventral stream modulate motor planning in the dorsal stream. Recent psychophysical reexaminations have intensified critiques of stream independence, particularly through scrutiny of classic experiments underpinning the hypothesis. A 2024 analysis revisited Milner and Goodale's foundational studies, arguing that visual illusions affect action to a greater degree than previously reported, with confounds like attention and task timing potentially inflating perceived dissociations between perception and action. These critiques highlight shared representational processes across streams in attention-demanding tasks, where a single underlying mechanism—rather than fully distinct systems—can account for both perceptual reports and motor responses, as evidenced by non-replications of key illusion effects and Bayesian modeling favoring integrated processing. Ongoing debates between proponents like Milner and Goodale and critics emphasize that while distinct representations may exist (two-reps view), the evidence for entirely separate processing systems (two-systems) remains moderate at best. The overlap of impairments in neurodegenerative disorders like Alzheimer's disease further undermines the notion of strict stream independence, revealing concurrent dysfunctions that suggest intertwined dependencies. In Alzheimer's, both ventral stream deficits—such as impaired object and face recognition—and dorsal stream impairments—like reduced motion perception and spatial navigation—are evident early, linked to amyloid deposition and atrophy in shared visual cortices (e.g., Brodmann area 19). Retinal thinning and optic nerve changes correlate with declines in both pathways, as shown in tasks engaging visuospatial organization that recruit elements from each stream, indicating that pathological processes do not respect modular boundaries. This widespread involvement challenges the hypothesis's modularity by demonstrating how unified neurodegenerative mechanisms disrupt the purportedly segregated functions of perception and action. A 2022 computational study used recurrent neural networks to examine the two-streams hypothesis, finding that ventral stream-like models require longer memory for viewpoint-invariant object classification compared to dorsal stream models for orientation and size estimation tasks.^[5] A 2025 study using intracranial EEG during memory-guided actions found increased alpha power (8-13 Hz) in both dorsal (inferior parietal lobule) and ventral (ventral temporal cortex) streams during working memory delays, with theta oscillations (2-7 Hz) in the ventral stream. Cross-frequency interactions between these regions indicated coupled stream activity, suggesting the streams operate in an integrated rather than strictly parallel manner during cognition.^[46] Proposals for refinements include subdividing the dorsal stream into multiple sub-streams and incorporating cross-sensory integrations to address limitations in unimodal processing. Neuroanatomical studies propose a dorso-dorsal sub-stream for online visuomotor control and a ventro-dorsal sub-stream for semantic action understanding, supported by connectivity patterns in primate parietal cortex that differentiate reaching (dorodorsal) from object manipulation (ventrodorsal) tasks. Additionally, visuo-auditory integrations in the posterior parietal cortex (PPC) enable multisensory binding, as evidenced by synchronized neural responses to congruent audiovisual stimuli, enhancing spatial localization accuracy by 40% in behavioral paradigms and extending the two-streams framework to multimodal cognition.^[47]^[48]

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A double dissociation between semantic and spatial cognition - eLife
May 17, 2024 · Analysis of intrinsic functional connectivity using resting-state fMRI. The results of Study 1 suggested separate visual-DMN pathways ...
[40]
The 'when' pathway of the right parietal lobe - PMC - PubMed Central
Much of the work we review investigates judgments of whether two events are simultaneous or whether two events are seen as independent or integrated into the ...
[41]
Neural Dynamics of Visual Stream Interactions During Memory ...
Mar 17, 2025 · Our study aimed to investigate the neural dynamics and functional connectivity between the dorsal and ventral streams, as well as the ...
[42]
Two what, two where, visual cortical streams in humans
A ventromedial 'Where' visual stream builds feature combinations for scenes, and provides 'Where' inputs via the parahippocampal scene area to the hippocampal ...Missing: dorsomedial | Show results with:dorsomedial
[43]
A modality-independent proto-organization of human multisensory ...
Jan 16, 2023 · Altogether, these results indicated that congruent auditory and visual streams elicited a functional synchronization in the superior temporal ...