The dorsal attention network (DAN) is a bilateral frontoparietal brain system that mediates the top-down, goal-directed control of visual attention, enabling the voluntary selection of specific locations, objects, or features in the environment based on internal goals and expectations. The DAN was first delineated in the early 2000s through neuroimaging studies, notably by Corbetta and Shulman (2002), distinguishing it from the ventral attention network. This network plays a crucial role in preparing sensory cortices for expected stimuli, maintaining spatial priority maps, and coordinating overt and covert shifts of attention, such as during saccade planning or visual search tasks.Anatomically, the DAN encompasses key cortical regions including the frontal eye fields (FEF)—located at the intersection of the precentral and superior frontal sulci—and the intraparietal sulcus (IPS), which includes subregions like the anterior, posterior, and ventral IPS, along with the superior parietal lobule. These areas are organized in a retinotopic manner, reflecting their sensitivity to visual space, and exhibit sustained activation during attentional tasks.[1] Subcortically, the DAN incorporates structures such as the pulvinar nucleus of the thalamus, superior colliculi, caudate head, and brainstem nuclei (e.g., pedunculopontine and cuneiform), which support preparatory signaling and sensory modulation.[2] Neurochemically, its activity correlates with acetylcholine (α4β2 nicotinic receptors), dopamine transporters, and serotonin systems, facilitating efficient top-down processing.[2]Functionally, the DAN sustains attentional sets for stimuli and responses, exerting modulatory influences on early visual areas to enhance processing of task-relevant information while suppressing distractors.[3] It is activated during endogenous cueing paradigms, such as Posner's spatial attention task, where bilateral FEF and IPS show prolonged responses to directional cues, contrasting with transient sensory activations.[1] The network also overlaps with visuospatial working memory, maintaining delay-period activity for up to 10 seconds to rehearse locations or features. Unlike stimulus-driven systems, the DAN does not respond strongly to unattended or unexpected events, focusing instead on proactive, voluntary orienting.[1]The DAN interacts dynamically with the ventral attention network (VAN)—a right-lateralized system involving the temporoparietal junction and ventral frontal cortex—to achieve flexible attentional control. While the DAN handles goal-directed selection, the VAN acts as a "circuit breaker" for reorienting to salient, behaviorally relevant stimuli, with the two networks connected via the superior longitudinal fasciculus for coordinated signaling.[1] Disruptions in DAN connectivity, as seen in conditions like spatial neglect, impair top-down attention and spatial working memory.[1][4]
Overview
Definition and characteristics
The dorsal attention network (DAN) is a frontoparietal brain network primarily responsible for endogenous, goal-directed control of attention, enabling the voluntary selection and prioritization of sensory information based on internal intentions or task demands.[5] This network facilitates top-down modulation of perceptual processing, allowing individuals to orient attention toward specific locations or features in the environment without reliance on salient external cues.[5]Key characteristics of the DAN include its bilateral organization, which supports attention deployment across both visual fields symmetrically, and its central role in spatial orienting and selection mechanisms.[5] Unlike bottom-up systems that respond reflexively to unexpected or salient stimuli, the DAN operates through sustained preparatory activity to maintain focus on relevant targets, ensuring efficient allocation of cognitive resources.[5] Functional connectivity analyses using resting-state fMRI have consistently identified the DAN as one of the major intrinsic brain networks.[6]In cognitive neuroscience, the DAN represents a foundational system for understanding voluntary attention, with its intrinsic connectivity patterns observable even in the absence of tasks, highlighting its stable architectural role in the human connectome.[6]
Historical background
The foundations of the dorsal attention network (DAN) concept trace back to early experimental paradigms in cognitive psychology that distinguished between voluntary and involuntary forms of attention. In the 1980s, Michael Posner's cueing paradigm provided critical evidence for separating endogenous attention, which involves goal-directed, top-down control using central symbolic cues, from exogenous attention, driven by salient peripheral stimuli. This distinction laid the groundwork for understanding attention as comprising multiple systems, with endogenous processes later associated with dorsal frontoparietal mechanisms.Building on these behavioral insights, computational models further shaped the theoretical framework for the DAN. The biased competition theory, proposed by Robert Desimone and John Duncan in 1995, posited that neural representations of multiple stimuli compete for processing resources, with top-down signals from higher cortical areas biasing selection toward behaviorally relevant information. This model emphasized the role of prefrontal and parietal regions in resolving competition, influencing subsequent conceptualizations of the DAN as a network facilitating such voluntary attentional control.A pivotal milestone came in 2002, when Maurizio Corbetta and Gordon L. Shulman integrated neuroimaging data from positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) to propose segregated dorsal and ventral attention systems. Their analysis of task-evoked activations revealed the dorsal system, encompassing frontoparietal regions, as primarily responsible for endogenous, goal-directed orienting, contrasting with the ventral system's role in stimulus-driven reorienting to salient events.The DAN's status as a distinct intrinsic network was solidified through resting-state fMRI studies in the mid-2000s. In 2006, Michael D. Fox and colleagues demonstrated that spontaneous low-frequency fluctuations in the BOLD signal exhibited strong anticorrelations between dorsal and ventral attention systems during rest, confirming their functional independence and extending the task-based findings to intrinsic brain organization. This work marked a shift toward viewing the DAN as a core component of the brain's large-scale networks, paving the way for broader applications in cognitive neuroscience.
Neuroanatomy
Core brain regions
The dorsal attention network (DAN) primarily consists of bilateral cortical regions in the frontal and parietal lobes that support voluntary spatial orienting and attention. These core areas are organized in a frontoparietal architecture, with frontal components focused on executive control of eye movements and parietal regions involved in mapping attentional priorities across space.The bilateral frontal eye fields (FEF), located in the superior frontal gyrus near the intersection of the precentral sulcus and superior frontal sulcus (Brodmann area 8), serve as key nodes for planning and executing saccades, integrating top-down signals to direct gaze toward behaviorally relevant locations. Adjacent supporting structures include the supplementary eye fields (SEF), situated on the medial surface of the superior frontal gyrus within the paracentral sulcus, which contribute to the coordination of complex oculomotor sequences and attentional shifts.[3] Additionally, regions in the middle frontal gyrus act as auxiliary nodes, facilitating the integration of attentional demands with prefrontal executive functions.[7]In the parietal cortex, the intraparietal sulcus (IPS), particularly its posterior segments, and the superior parietal lobule (SPL) form the dorsal posterior hubs, enabling the representation and selection of spatial attentional maps to guide voluntary orienting. These parietal areas are bilaterally symmetric and positioned along the lateral and medial aspects of the parietal lobe, respectively, providing a topographic organization for attentional allocation.[1]Subcortical modulation of the DAN arises from structures including the thalamus, basal ganglia, superior colliculi, and brainstem nuclei (e.g., pedunculopontine and cuneiform), which gate attentional signals and enhance network efficiency. The pulvinar nucleus of the thalamus plays a critical role in attentional gating, filtering sensory inputs to prioritize relevant spatial information for cortical processing. The superior colliculi contribute to the integration of sensory and motor signals for orienting responses. Contributions from the basal ganglia, including the head of the caudate nucleus, support the modulation of goal-directed attention through dopaminergic pathways that influence frontal and parietal activity. The pedunculopontine and cuneiform nuclei provide neuromodulatory inputs via cholinergic, glutamatergic, and GABAergic pathways.[2]
Connectivity patterns
The dorsal attention network (DAN) exhibits robust intra-network connectivity, primarily mediated by white matter tracts that link its core frontal and parietal regions. The superior longitudinal fasciculus (SLF), a major association fiber bundle, provides direct structural connections between the frontal eye fields (FEF) and the intraparietal sulcus (IPS), facilitating rapid signal transmission for spatial attention orienting.[8] Diffusion tensor imaging studies have delineated the SLF's role in linking these areas bilaterally, with the SLF III segment specifically connecting the FEF to the superior parietal lobule and IPS, supporting the network's coordinated activation during goal-directed tasks.[8] Additionally, the supplementary eye field (SEF) connects to the superior parietal lobule (SPL) via parallel pathways, contributing to the representation of attentional priorities in both egocentric and object-centered reference frames.[3]Functionally, the DAN demonstrates strong within-network coherence during attention-demanding tasks, with hubs in the FEF and IPS showing synchronized BOLD signal fluctuations. Resting-state fMRI reveals that these task-positive regions are intrinsically anticorrelated with the default mode network (DMN), exhibiting negative correlations (e.g., r ≈ -0.3 to -0.5 across seeds) that reflect a push-pull dynamic between external focus and internal mentation. This anti-correlation persists across resting conditions (eyes open, closed, or fixation) and is modulated during tasks, where DAN activation suppresses DMN activity to enhance attentional control. Such patterns underscore the DAN's role as a functional hub for top-down modulation, with connectivity strength predicting behavioral performance in spatial attention paradigms.[9]Afferent inputs to the DAN originate from early visual processing areas, integrating sensory information to guide attentional selection. Projections from visual cortical regions, including area V4, convey feature-specific signals to the IPS, enabling the network to prioritize relevant stimuli based on color, shape, and motion attributes. These bottom-up afferents support the DAN's ability to maintain a dynamic salience map, combining sensory inputs with goal-directed biases. Efferent outputs from the DAN, particularly from the FEF, project to motor areas such as the superior colliculus and brainstem oculomotor nuclei, executing orienting responses like saccades and shifts in gaze. These descending pathways ensure that attentional priorities translate into overt behavioral actions, closing the loop from perception to response.[10]The DAN's connectivity reflects a hierarchical organization, with the FEF serving as a primary source of top-down control that influences downstream parietal regions. Effective connectivity analyses indicate unidirectional influences from FEF to IPS during spatial attention tasks, where frontal signals bias parietal processing to align with behavioral goals. This frontoparietal hierarchy allows the FEF to generate preparatory attentional sets that propagate to the IPS for sensory enhancement and to motor effectors for execution, establishing a graded flow of information within the network.[10]
Functions
Role in top-down attention
The dorsal attention network (DAN) plays a central role in top-down attention by generating predictive signals that bias sensory processing toward task-relevant stimuli, enhancing the representation of goal-directed information while modulating irrelevant inputs. This voluntary control mechanism allows individuals to prioritize endogenous goals, such as searching for a specific object in a cluttered scene, by amplifying neural responses in sensory cortices aligned with current intentions. For instance, preparatory activity in the DAN facilitates the selection of spatial locations or features, drawing from internal representations to guide behavior without reliance on external salience.[10]In spatial attention shifts, the DAN supports endogenous cueing, as demonstrated in Posner's paradigm, where symbolic cues (e.g., arrows) direct attention to anticipated locations, resulting in faster reaction times to validly cued targets compared to invalid ones. This process involves preparatory orienting, with the DAN activating to reallocate attentional resources voluntarily, leading to improved detection accuracy and speed for task-relevant stimuli. Experimental evidence from event-related fMRI shows that valid cues elicit sustained activation in DAN nodes, particularly during the delay period before target onset, confirming its involvement in goal-directed spatial selection. The DAN, comprising key frontoparietal areas like the frontal eye fields and intraparietal sulcus, underlies this endogenous control.[10][11]For sustained attention, the DAN maintains focus over extended periods, such as in working memory tasks requiring the holding of spatial information against interference. It achieves this by continuously suppressing distractors through top-down inhibition, preventing shifts to irrelevant stimuli and preserving the attentional set. In feature-based attention paradigms, like variants of the sustained attention to response task, increased DAN activity correlates with effective distractor suppression, particularly when high-priority irrelevant items demand greater inhibitory control. This sustained engagement ensures stable performance in prolonged goal-directed activities.[10][12]Neural dynamics of the DAN during top-down attention are characterized by increased BOLD signals in event-related fMRI studies of voluntary orienting tasks. Preparatory cues trigger bilateral activation in the intraparietal sulcus and frontal eye fields, with signal magnitude scaling with attentional demands and persisting through maintenance phases. These patterns reflect the network's role in generating top-down biases that enhance target processing while dampening distractor responses, as observed in tasks involving spatial or feature-based selection.[10][13]
Integration with sensory processing
The dorsal attention network (DAN) interfaces with early sensory cortices through top-down feedback mechanisms that amplify relevant perceptual signals, enhancing the processing of attended stimuli while suppressing irrelevant ones. Specifically, regions such as the frontal eye fields (FEF) and intraparietal sulcus (IPS) within the DAN exert modulatory influences on primary visual cortex (V1), increasing neural activity for stimuli at attended locations. This feedback is evidenced by directional connectivity analyses, including Granger causality and dynamic causal modeling, which demonstrate that FEF and IPS activity precedes and predicts V1 responses during spatial attention tasks. Similar amplification occurs for auditory signals, where DAN projections to early auditory areas sharpen representations of attended sounds, prioritizing them amid competing inputs.In feature-based attention, the DAN, particularly via IPS involvement, selects and enhances specific object properties across sensory modalities, refining perceptual representations without reliance on spatial cues alone. For instance, attending to a particular color or orientation activates IPS regions that bias processing in visual areas like V4, facilitating the integration of features from multiple objects or locations. This process allows for selection of visual features, where IPS signals propagate to visual cortices to prioritize matching attributes, such as a specific motion direction. Functional imaging studies confirm IPS activation during feature-cueing paradigms, underscoring its role in modulating sensory cortices to improve featurediscrimination.[14]The DAN addresses attentional bottlenecks by implementing resource allocation models that prioritize competing sensory inputs, preventing overload in limited-capacity perceptual systems. Drawing from biased competition frameworks, DAN nodes like FEF generate top-down biases that resolve conflicts among multiple stimuli, favoring goal-relevant ones through enhanced gain in sensory areas. This prioritization is crucial during high-load scenarios, where DAN-mediated suppression of unattended inputs reduces interference, as seen in models of neural competition in frontoparietal circuits. Such mechanisms ensure efficient sensory throughput, with DAN activity scaling to task demands to maintain perceptual acuity.[15]Evidence from cueing studies further illustrates DAN's role in sensory integration, showing that valid spatial or temporal cues lead to improved detection thresholds for attended stimuli. In Posner's classic paradigm, predictive cues activate the DAN, resulting in faster response times and higher accuracy for cued locations, attributed to enhanced sensory gain in contralateral visual fields. Neuroimaging during these tasks reveals DAN connectivity strengthening to sensory regions, correlating with behavioral benefits like reduced perceptual uncertainty. These findings highlight how DAN-driven orienting refines sensory processing, amplifying signal-to-noise ratios for task-relevant inputs.[16][3]
Network interactions
Relation to ventral attention network
The dorsal attention network (DAN) and ventral attention network (VAN) form a complementary dichotomy in attentional control, with the DAN supporting sustained, voluntary, goal-directed attention to expected stimuli, while the VAN facilitates transient, reflexive reorienting to unexpected or salient events.[1] This division enables efficient allocation of cognitive resources, where the DAN maintains focus on task-relevant information and the VAN detects deviations that require rapid shifts in attention.[6]At the circuit level, the right-lateralized VAN, encompassing the temporoparietal junction (TPJ) and ventral frontal cortex, interacts with the DAN by interrupting its ongoing activity during behaviorally salient events, such as novel or unexpected stimuli, to facilitate attentional reorienting. This interaction is evident in neuroimaging studies showing VAN activation suppresses DAN regions, particularly in the intraparietal sulcus and frontal eye fields, allowing for bottom-up capture of attention.[1]The circuit-switching model describes this dynamic interplay as a mutual suppression and enhancement mechanism, particularly in tasks involving invalid cueing where the VAN disengages the DAN to redirect attention toward uncued locations.[11] In such paradigms, the VAN's role in signaling discrepancies leads to enhanced DAN engagement post-reorienting, ensuring adaptive attentional control without persistent interference.[1]Behavioral evidence underscores the VAN's critical role in supporting DAN function; lesions to the right VAN, as seen in hemispatial neglect syndrome, impair the ability to reengage DAN-mediated attention toward contralesional stimuli, resulting in profound deficits in detecting and responding to left-sided events. This asymmetry highlights the VAN's dominance in initiating reorienting, without which the DAN's top-down mechanisms fail to compensate effectively.[1]
Links to default mode and salience networks
The dorsal attention network (DAN) displays a robust anti-correlation with the default mode network (DMN), a pattern where increased DAN activity during goal-directed tasks suppresses DMN engagement, thereby minimizing mind-wandering and internal mentation. This intrinsic opposition ensures efficient allocation of cognitive resources toward external stimuli, as observed in functional magnetic resonance imaging (fMRI) studies of working memory and attentional tasks.[17] Disruptions in this anti-correlation have been linked to attentional inefficiencies, highlighting its role in maintaining focused states.[18]The salience network (SN), anchored in the anterior insula and dorsal anterior cingulate cortex, serves as a mediator in modulating interactions between the DAN and DMN by detecting behaviorally relevant stimuli and directing network switching. Upon identifying salience, the SN facilitates DAN activation for top-down control while inhibiting DMN activity, enabling rapid transitions from introspective to externally oriented processing.[19] This mechanism underscores the SN's pivotal function in integrating sensory inputs with attentional priorities.[20]These dynamics are formalized in the triple-network model, which posits the SN as a central hub for orchestrating switches between the central executive network (encompassing the DAN) and the DMN during cognitive state changes. Empirical evidence from Granger causality analyses supports this framework, demonstrating directional influences from SN regions that bias toward DAN dominance in task-relevant contexts.[21]At rest, fluctuations in DAN-DMN connectivity further illustrate these links, with reduced anti-correlations predicting increased attentional lapses and variability in sustained performance.[22] Such resting-state patterns reflect the networks' ongoing antagonism, where lapses occur when DMN incursions overpower DAN suppression, as evidenced in studies of vigilance tasks.[23] This connectivity metric thus serves as a biomarker for attentional stability across individuals.[24]
Clinical and research implications
Disorders and dysfunctions
Hemispatial neglect, a common consequence of right-hemisphere strokes, arises from disruptions to the dorsal attention network (DAN), particularly in the intraparietal sulcus and frontal eye fields, leading to profound inattention to stimuli on the contralesional (typically left) side of space.[25] This impairment manifests as a failure to orient toward behaviorally relevant targets in the neglected hemifield, despite intact primary sensory processing, and is exacerbated by damage that severs the network's ability to mediate voluntary spatial attention.[26] Patients often exhibit anosognosia for their deficit, complicating rehabilitation, with severity correlating to the extent of DAN lesion overlap.[27]In attention-deficit/hyperactivity disorder (ADHD), the DAN shows hypoactivation and reduced functional connectivity during tasks requiring sustained attention, such as continuous performance tests, contributing to core symptoms of inattention and executive dysfunction.[28] This network inefficiency is evident in both children and adults with ADHD, where lower intra-network coherence in frontal and parietal regions impairs goal-directed attentional control and sensory selection.[29] Structural alterations, including decreased white matter integrity in DAN pathways, further underlie these deficits, linking them to broader cognitive impairments like working memory lapses.[30]Schizophrenia is associated with aberrant DAN connectivity and atypical activation patterns, which disrupt attentional fragmentation and contribute to disorganized cognition and perceptual disturbances.[31] During tasks involving perceptual organization, individuals with schizophrenia exhibit distinct DAN hypoengagement in right-hemisphere regions, predictive of symptom severity such as cognitive disorganization.[32] These alterations often involve imbalanced interactions with the ventral attention network, leading to inefficient reorienting and heightened distractibility in everyday functioning.[33]Aging leads to declining DAN efficiency, characterized by reduced within-network functional connectivity and weakened anticorrelations with the default mode network, resulting in difficulties with divided attention and selective focus.[34] Older adults display slower reorienting responses and diminished parietal alpha/beta oscillations during attentional shifts, correlating with everyday lapses like forgetting task goals.[35] These changes, progressive from middle age onward, reflect broader network dedifferentiation that impairs top-down control, though interventions like social engagement can partially restore DAN integrity.[36]
Neuroimaging evidence and recent studies
Functional magnetic resonance imaging (fMRI) studies using task-based paradigms, such as the Posner cueing task, have demonstrated robust activation in the dorsal attention network (DAN) during endogenous orienting of attention, with key nodes including the frontal eye fields (FEF) and intraparietal sulcus (IPS) showing increased BOLD signals when participants voluntarily shift spatial attention to cued locations.[1] These paradigms reveal that the DAN facilitates top-down control by enhancing sensory processing in expected visual fields, as evidenced by effective connectivity from FEF and IPS to early visual areas during valid cues.[37]Resting-state fMRI combined with independent component analysis (ICA) has been instrumental in parcellating the DAN, identifying its core components through intrinsic functional connectivity patterns that persist in the absence of tasks, such as anticorrelations with the default mode network.[38] This approach has refined DAN boundaries, distinguishing it from adjacent frontoparietal networks and highlighting developmental asymmetries in connectivity between children and adults.[39]Diffusion tensor imaging (DTI) and tractography techniques have mapped the structural integrity of frontoparietal tracts underpinning the DAN, revealing white matter pathways like the superior longitudinal fasciculus that connect FEF and IPS, with fractional anisotropy values indicating their role in efficient attentional orienting.[40] These methods demonstrate that disruptions in tract integrity correlate with impaired attentional performance, providing a structural basis for functional observations.[8]Recent 2024 research has explored DAN-ventral attention network (VAN) interactions in pain modulation, showing altered functional connectivity in chronic pain patients where enhanced DAN activity contributes to endogenous pain amplification via top-down mechanisms.[41]A 2025 study on functional connectivity between the dorsal and ventral attention networks found that VAN regions, particularly in the right hemisphere (e.g., inferior frontal gyrus and supramarginal gyrus), show stronger associations with attentional disengagement effects compared to DAN regions.[42]Integrations of transcranial magnetic stimulation (TMS) with electroencephalography (EEG) have established causal roles of the FEF in attentional orienting within the DAN, where inhibitory TMS over FEF disrupts alpha-band oscillations in contralateral visual cortex, impairing spatial attention shifts.[43] Repetitive TMS protocols reveal lasting effects, with post-stimulation reductions in FEF excitability persisting for minutes and correlating with prolonged deficits in visuospatial orienting tasks.[44]