Brain Networks and Attention

Learning Objectives

After completing this section, you will be able to:

A Neurofeedback Paradigm Shift

With neurofeedback, we want to know where to place active electrodes and what brain activity to train. It is natural to assume that the sensor should sit over the part of the brain corresponding to a Brodmann area that needs to be up-trained or down-trained. That approach can work for some conditions that respond to single-channel training. However, one site connects to another, and so on, within networks.

Networks, which can be functional or structural, mediate connectivity. Connectivity usually serves a purpose, such as mediating a cognitive or emotional process like thinking or feeling. In functional networks, regions show correlated activity over time. Structural networks consist of axonal projections and pathways. Functional and structural networks can overlap, but functional connectivity does not automatically imply an underlying anatomical connection.

Fornito et al. (2016) emphasized the role of time scale in their convergence. As functional connectivity is averaged over longer periods, it may converge onto structural connectivity, although the two remain different measures that can yield connectomes with different topological values (p. 25).

Interventions as different as immobilizing a limb and neurofeedback can change functional connectivity. Newbold and Dosenbach (2021) placed a cast on the intact dominant hand of participants for two weeks and ran repeated fMRI scans to observe the effect on motor cortex connectivity. Within a day or two, functional connectivity between the left and right motor cortices had nearly disappeared, a faster timescale than many had predicted. Recovery was also rapid in two of the three participants.

The premise of connectivity training is that neurofeedback, using EEG and functional MRI, can increase or decrease functional connectivity to improve performance. Single-channel neurofeedback training may also alter connectivity within brain networks that are functionally or structurally linked to the trained cortical site.

Commonsense About Functional Brain Areas

Petersen and Fiez (1993) observed that a functional area of the brain is not a task area. There is no "tennis forehand area" waiting to be discovered, and no single area is devoted to a complex function, so attention and language are not localized to one Brodmann area or lobe. Any task or function draws on a complex, distributed set of brain areas. These networks, or coordinated sets of brain regions, likely activate and deactivate in intricate patterns that remain poorly understood.

The areas that perform a task are distributed across different locations, yet the processing is not diffusely spread among them. Each area makes a specific contribution that is determined by where it sits within its richly connected, parallel, distributed hierarchy.

Key Anatomical Terms

Long brain region names can be intimidating, and six descriptions can serve as a decoder ring. Anterior or rostral refers to toward the head end, whereas posterior or caudal means toward the tail. Dorsal means toward the top of the brain, while ventral means toward the bottom. A gyrus is a ridge of convoluted brain tissue, whereas a sulcus is a furrow (Breedlove & Watson, 2023).

Networks carry both functional and anatomical names. For example, the dorsal attention network (DAN) corresponds to the dorsal frontoparietal network (D-FPN).

Brodmann Areas

The original Brodmann areas consisted of 47 numbered cytoarchitectural zones of the cerebral cortex based on Nissl staining. They require subdividing certain areas, and they vary in size and shape across individuals.

The numbered Brodmann areas of the cerebral cortex.

Gordon et al. (2016) compared the original Brodmann areas with several other brain atlases using fMRI. They concluded that Brodmann's parcellation captures real structure in the data but is too coarse to represent true cortical areas. Modern attempts to parcellate the cortex frequently reveal finer architectonic divisions than Brodmann reported. The more than 100-year-old Brodmann areas therefore lack some of the specificity needed for in vivo and functional studies.

Revised Brodmann maps reveal 180 regions per hemisphere, 97 of which were not previously identified (Glasser et al., 2016). Cognitive neuroscience tends to use modern brain atlases of defined regions (e.g., Yeo et al., 2011) and exact coordinates such as Montreal Neurological Institute (MNI) space and Talairach space (Chau & McIntosh, 2005). Researchers sometimes convert these coordinates to corresponding Brodmann areas because the areas are well known.

Brodmann areas participate in networks. They matter for neurofeedback because they help us target functional and structural networks rather than discrete, disconnected scalp sites. Neuroscience reveals the connections between Brodmann areas and how different networks become active or quiet together across diverse conditions.

For example, attention to an object's location engages the bilateral dorsal attention network (dorsal frontoparietal network), comprising the intraparietal sulcus and the frontal eye fields, while the largely right-lateralized ventral attention network (ventral frontoparietal network), comprising the temporoparietal junction and ventral frontal cortex, automatically processes unanticipated events. Flexible attention control involves the dynamic interaction of these top-down and bottom-up systems (Vossel et al., 2014).

Psychological disorders can be associated with shifts from normal activity in particular brain areas and their connections. After traumatic brain injury (TBI), neuronal connections change across the entire brain, and surviving long axonal projections no longer target inhibitory neurons (Frankowski et al., 2022).

Current neurofeedback protocols allow us to train networks involved in cognitive functions like attention or psychological disorders like depression using multiple electrodes simultaneously. Connectivity training enables us to increase or decrease communication between brain locations to treat symptoms and improve performance, and both activations and deactivations matter. Quantitative EEG (qEEG) normative databases can reveal the key parts of a network that need training and the required direction.

Brain Organization and Dynamics

This section introduces the brain's network architecture, from large-scale systems to specific cortical regions. Understanding these systems is essential for neurofeedback practitioners, since many protocols target the networks described here.

The brain is organized into interactive, functional, distributed networks with spatial, temporal, and content-based relationships. These networks interact through feedback loops and transiently organized aggregates of neurons, all mediated by rhythmic, oscillatory electrical discharges that ultimately produce the EEG. This process is further shaped by selective attention to specific categories of interest.

Each type of local cognitive, sensory processing, or emotional network produces oscillatory activity and contains internal stabilizing characteristics. These local networks exist within a global dynamic network system that links and provides interactive capacity to the smaller networks, also operating within an oscillatory framework.

A densely connected lateral prefrontal and posterior parietal cortical network orchestrates responses to novel cognitive tasks using flexible hubs. This frontoparietal network assigns tasks to the most appropriate brain regions and shares information among them to master new skills (Cole et al., 2013).

The central nervous system processes incoming content through separate regions that handle specialized input (e.g., auditory, kinesthetic, tactile, visual). Content is shared, integrated, compared to previous content, and analyzed, and decisions are made about memory and responses. All of this activity occurs within interacting networks linked by electrical and chemical signals, and the electrical discharges from this activity are recorded from the scalp surface as the EEG.

Network Overview

The networks most relevant to attention include the oculomotor, motor, affective, central autonomic, social, and executive circuits. Each plays a distinct role in how we focus, respond, and regulate behavior.

Oculomotor Network

The frontal eye field (FEF), in concert with the dorsolateral prefrontal cortex, posterior parietal cortex, basal ganglia, and thalamus, programs and initiates voluntary eye movements, inhibits eye movements toward distracting stimuli, and allows us to return our focus to locations we have previously attended (Thompson & Thompson, 2015).

Motor Network

The supplementary motor area (SMA), in concert with the premotor cortex, primary motor cortex, sensorimotor cortex, and cerebellum, plans, initiates, and inhibits voluntary movements and muscle contractions (Breedlove & Watson, 2023; Thompson & Thompson, 2015). Understanding the motor network helps clinicians appreciate why surface EMG (SEMG) biofeedback and sensorimotor rhythm (SMR) neurofeedback can influence broader neural circuits.

Somato-Cognitive Action Network (SCAN)

Gordon and colleagues (2023) used precision fMRI from seven participants together with datasets from the Adolescent Brain Cognitive Development Study, the Human Connectome Project, and the UK Biobank, drawing on roughly 50,000 individuals. They identified three interconnected primary motor cortex (M1) regions that participate in the integrated movement of multiple body parts.

The somato-cognitive action network (SCAN) consists of three inter-effector regions of M1, the SMA, the dorsal anterior cingulate cortex, the centromedian and ventral intermediate nuclei of the thalamus, the posterior putamen, and the vermis and flocculonodular lobe of the cerebellum, which mediate posture and balance. Connectivity analysis showed that the SCAN communicates with the cingulo-opercular network, or salience network, which supports cognitive control and sustains task focus over extended periods.

The somato-cognitive action network (SCAN).

M1's two interlacing systems establish a pattern of integration and isolation. Effector-specific regions for the foot, hand, and mouth support the isolation of fine motor control, while the SCAN integrates goals, body movement, and physiology.

The relative expansion of SCAN regions in humans may support uniquely human integrated actions, such as coordinating breathing for speech and combining hand, body, and eye movement for tool use. By enabling anticipatory postural, respiratory, cardiovascular, and arousal adjustments before action, such as shoulder tension, increased heart rate, or butterflies in the stomach, the SCAN provides a substrate that may help explain why mental and bodily states so often interact.

Mesa (2023) placed these findings in context. The dominant paradigm has treated the motor cortex as a simple relay that passes movement commands to muscles, with planning, cognition, and conscious initiation of movement occurring elsewhere in the brain.

The SCAN, in concert with the salience network, supports complex adaptations such as allostasis. These findings agree with primate studies showing that more M1 neurons govern movements independent of the specific muscles used than govern the contraction of particular muscles (Griffin et al., 2015; Kaufman et al., 2014). Together they challenge the cortical homunculus that Penfield and colleagues mapped in the 1930s, the distorted human figure whose body parts are sized by the amount of cortical area dedicated to them.

Affective Network

The pre- and subgenual (beneath the curved "knee") areas of the anterior cingulate cortex (ACC) participate in affective circuits triggered when we make mistakes (Arnsten, 2009). The dorsal rostral cingulate zone monitors cognitive activity to predict when errors are likely and greater executive control may be needed (Thompson & Thompson, 2015). The ventromedial prefrontal cortex projects to the amygdala, basal ganglia, hypothalamus, and brainstem arousal and reward pathways. This network is relevant to neurofeedback for anxiety and mood disorders.

Central Autonomic Network

The central autonomic network (CAN) consists of forebrain, limbic, and brainstem regions (Benarroch, 1993). Neuroimaging studies reveal a cortico-limbic network for autonomic control that includes the ventromedial prefrontal cortex, cingulate cortex, insula, mediodorsal thalamus, hypothalamus, and amygdala (Beissner et al., 2013; Schumann et al., 2021; Shoemaker et al., 2015).

Thayer and Lane (2000) proposed a neurovisceral integration model that places the prefrontal cortex at the top of a hierarchy, with direct functional links to the insula and cingulate. The limbic system extends these connections through the amygdala to downstream subcortical regions, such as the hypothalamus and brainstem nuclei, that drive parasympathetic and sympathetic heart rate modulation. The model highlights how prefrontal control over subcortical structures links sympathovagal balance with cognitive and emotional processes. This framework is central to HRV biofeedback, which trains the prefrontal and autonomic circuitry that governs heart rate regulation.

Magnetic resonance imaging (MRI) and resting-state functional connectivity (RSFC) studies have reinforced the role of medial prefrontal and limbic interplay in heart rate control (Kumral et al., 2019; Sakaki et al., 2016). Individuals with slower heart rates show heightened RSFC within a network of central autonomic and sensorimotor regions compared with those who have faster heart rates (de la Cruz et al., 2019). Slower heart rates were associated with elevated RSFC between the ventromedial prefrontal cortex and the anterior insula.

Social Network

The orbitofrontal cortex (OFC), along with the basal ganglia and thalamus, orchestrates the highest level of emotional processing in the nervous system. The social network is responsible for socially responsible behavior, empathy, behavioral inhibition, emotional regulation, and sound judgment (Thompson & Thompson, 2015).

A mentalizing network that includes the ventromedial, orbitofrontal, dorsolateral, and dorsomedial prefrontal cortices lets us reason about our own and others' mental states (Hoskinson et al., 2019). A mirror network centered on the superior temporal sulcus is active during our own actions and our observation of others' actions, supporting observational learning and social cognition (Sadeghi et al., 2022). The amygdala detects salient stimuli, while the entorhinal cortex and anterior insular cortex contribute to this social circuitry.

Executive Network

The dorsolateral prefrontal cortex plays a critical role in executive functions, which Kropotov (2009) described as the coordination and control of motor and cognitive actions to attain specific goals. Executive functions include allocation of attention, cognitive inhibition, behavioral inhibition, working memory, and cognitive flexibility. The executive network focuses and maintains continuous attention (Faraone et al., 2006) and shows reduced activation and connectivity in ADHD, making it a primary target of neurofeedback protocols for ADHD and other attention-related conditions.

Attentional Processes

Attention is the selection of sensory information or cognition for enhanced processing. We can overtly or covertly attend to stimuli. In overt attention, our attentional focus and sensory orientation coincide. For example, you parse this sentence as you focus your gaze on it.

In covert attention, we shift our attentional focus from our sensory orientation. For example, you attend to a reminder on the corner of your screen while gazing at this sentence. While the midbrain superior colliculus is mainly implicated in overt attention, it may also regulate covert attention (Breedlove & Watson, 2023).

Attention is more selective than arousal, which is our overall level of alertness (Breedlove & Watson, 2023).

Research using divided attention tasks, where subjects simultaneously process multiple stimuli, shows that attentional resources are finite. The challenge of attending to more than one target increases when targets occupy different spatial locations. When we attend to a stimulus, we shift our attentional spotlight (focus) to select that stimulus for enhanced analysis (Bee & Micheyl, 2008).

Perceptual load (stimulus processing demands) determines the level at which an attentional bottleneck (stimulus selection) occurs. Complex stimuli involving a high perceptual load monopolize processing resources, resulting in early selection, where we filter out lower-priority competing stimuli before preliminary perceptual and semantic analysis.

Conversely, simple stimuli involving a low perceptual load leave free processing resources, resulting in late selection, where we filter out competing stimuli after performing extensive analysis (Lavie et al., 2004). This distinction has practical implications for neurofeedback, since training in high-demand environments may engage different attentional mechanisms than training in quiet clinical settings.

Cortical Regions That Guide Attention

The dorsal frontoparietal system, comprising the intraparietal sulcus and frontal eye field, is responsible for the top-down direction of attention (Breedlove & Watson, 2023). The intraparietal sulcus (IPS), located in the parietal lobe, provides voluntary top-down steering of attention (Corbetta & Shulman, 1998).

The frontal eye field (FEF), found in the premotor region of the frontal lobes, directs gaze toward targets selected by the IPS (Paus et al., 1991). Target selection is guided by cognitive goals (top-down processing) rather than stimulus characteristics (bottom-up processing).

In contrast, the temporoparietal junction (TPJ), where the superior temporal gyrus and inferior parietal lobe intersect, mediates bottom-up shifts in attention in response to stimulus attributes (Corbetta & Shulman, 2002). The TPJ functions like a circuit breaker, overruling immediate attentional priorities and reallocating attentional resources to a new target (Breedlove & Watson, 2023).

Two Cortical Networks Regulate Attention

Two cortical networks cooperatively regulate subcortical and cortical systems to produce a coherent perceptual experience (Breedlove & Watson, 2023). A dorsal frontoparietal system provides top-down control of endogenous attention (voluntary attention), directing the attentional spotlight to support cognitive system priorities.

A right temporoparietal system provides bottom-up control of exogenous attention (involuntary reflexive attention), redirecting attention based on the novelty or importance of incoming stimuli.

Extensive interconnections between the two networks allow us to fluidly redirect attention from stimuli that are forebrain priorities (IPS) to those that are unexpected. This balance between top-down and bottom-up attention is a key consideration in neurofeedback training for attention disorders.

Salience Network

The salience network comprises structures that monitor our external and internal environments to determine which inputs are essential and require further processing and attention. The insula, primarily the anterior insula, is a crucial component because it facilitates bottom-up access to the brain's attentional and working memory resources (Menon & Uddin, 2010). The cingulate gyrus, particularly the right dorsal anterior cingulate cortex, is another crucial component (Thompson & Thompson, 2015).

The salience network. Graphic shared by Nekovarova, Fajnerova, Horacek, and Spaniel under the Creative Commons Attribution 3.0 license.

The clinical literature on ADHD, depression, and schizophrenia has explored the role of the anterior cingulate cortex in these disorders. Deep brain stimulation of this region has successfully improved treatment-resistant depression (Mayberg et al., 2005).

The insula, a cortical region located within the lateral sulcus, functions as an integrative hub for the salience network. The insula integrates interoceptive awareness (perception of internal body signals), emotional experience, and external perception to facilitate an individual's global perception of the world. It directs specific networks in processing salient stimuli and generating appropriate responses (Wiebking & Northoff, 2014). For biofeedback practitioners, the insula's role in interoceptive awareness is particularly relevant, since biofeedback training fundamentally depends on clients learning to detect and interpret internal signals.

The insula appears to provide an interface between the brain's cognitive, homeostatic, and affective systems, linking the areas involved in monitoring internal signals with those engaged in processing incoming external sensory streams. The insula detects salient events via afferent pathways and switches between other large-scale networks when these events are recognized, thereby guiding attention and working memory.

The anterior and posterior insula interact to regulate autonomic responses to salient stimuli. Interactive communication between the insula and anterior cingulate cortex facilitates access to the motor system (Menon & Uddin, 2010).

This network appears to help us switch between the task-oriented (executive) network and the internally focused default mode network (Seeley et al., 2007; Shirer et al., 2012).

Default Mode Network (DMN)

Brain regions are selectively active when we are conscious (Breedlove & Watson, 2023). The default mode network (DMN) consists of frontal, temporal, and parietal lobe circuits active during spontaneous cognition like introspection, daydreaming, and streams of consciousness. The DMN appears to contribute flexible memory retrieval and idea generation, critical elements of creativity, and is relatively inactive when pursuing external goals (Andrews-Hanna et al., 2010). In neurofeedback practice, understanding the DMN helps explain why some clients need to learn to suppress mind-wandering, as in ADHD, while others may benefit from cultivating it, as in creative performance training.

ADHD has been linked to irregular connectivity among brain regions, including within the DMN (Cao et al., 2014). The strength of specific brain connections can forecast variations in a person's capacity to sustain attention, and this holds true even at rest when the individual is not engaged in a specific task (Rosenberg et al., 2016, 2017).

There has been increasing discussion about whether some DMN regions belong to a distinct Parietal Memory Network that includes the precuneus, the mid-cingulate cortex, and the posterior inferior parietal lobule and dorsal angular gyrus (Gilmore et al., 2015; Hu et al., 2016).

Deactivations can be as important as activations. The degree of DMN deactivation appears critical for attentional control, since people who suppress it more readily can learn new material more easily (Nelson et al., 2016; Zerr et al., 2018). The DMN also helps synthesize details into single coherent events and supports envisioning the future (Gilmore et al., 2018).

The DMN may contribute to creative fluency, the ability to generate innovative ideas like alternative uses for everyday objects. A study of neurosurgical patients showed that left DMN stimulation reduced the number of uses generated but not their originality (Shofty et al., 2022).

Understanding Ourselves

The posterior cingulate cortex (PCC) and precuneus combine bottom-up attention with information from memory and perception. The ventral (lower) part of the PCC activates in all tasks involving the DMN, including those related to the self or others, remembering the past, thinking about the future, processing concepts, and spatial navigation. The dorsal (upper) part of PCC mediates involuntary awareness and arousal. The precuneus is concerned with visual, sensorimotor, and attentional information.

The medial prefrontal cortex (mPFC) participates in decisions about the self, such as personal information, autobiographical memories, future goals and events, and decision-making regarding those close to us like family members. The ventral (lower) part is involved in positive emotional information and reward.

The angular gyrus connects perception, attention, spatial cognition, and action and helps us recall episodic memories.

Understanding Others

The major functional hubs include the PCC, mPFC, and angular gyrus. The dorsal medial prefrontal cortex (dmPFC) participates in analyzing others' objectives. The temporoparietal junction (TPJ) contributes to theory of mind, models of others' cognitive processes, emotions, knowledge, and motivation. The lateral temporal cortex is concerned with short-term verbal memory, naming, and reading. Finally, the anterior temporal pole is part of a bilateral semantic system representing object concepts and a left hemisphere-dominant network concerned with naming and understanding object names.

Autobiography and Future Simulations

The major functional hubs include the PCC, mPFC, and angular gyrus. The hippocampus forms new declarative memories. The parahippocampal cortex (PHC) mediates spatial memory, navigation, and high-level visual processing like facial recognition. The retrosplenial cortex (RSC) is involved in episodic memory, navigation, predicting future events, and analyzing visual scenes. Finally, the posterior inferior parietal lobe (pIPL) integrates sensory information and participates in top-down attentional orienting.

The Pulvinar Mediates Attentional Shifts

The pulvinar nucleus, comprising the posterior quarter of the human thalamus, processes visual information and directs attention. The pulvinar shares widespread connections with the cingulate, parietal cortex, and superior colliculus, and is crucial for orienting, shifting attention, and filtering out irrelevant stimuli. Tasks that present subjects with more distracting stimuli increase pulvinar activation, as shown by functional MRI (fMRI) (Buchsbaum et al., 2006).

Overall, the pulvinar guides the processing of relevant information across wide-ranging cortical networks based on dynamically changing attentional priorities (Breedlove & Watson, 2023; Saalmann et al., 2012).

Training Networks Summary

Neurofeedback training increasingly monitors and trains network activity using qEEG normative databases. To employ these protocols effectively, neurofeedback professionals must thoroughly understand Brodmann areas and the functional and structural networks in which they participate. The dorsal frontoparietal system provides top-down attentional control while the temporoparietal system enables bottom-up redirection, with the salience network serving as the switch between them. This network architecture provides the rationale for targeting specific networks and frequencies to address attention disorders, optimize performance, and restore healthy brain function.

Assignment

Choose one client presentation you see often, such as inattention, anxiety, or performance under pressure. Identify which of the networks in this chapter you would expect to be involved, and explain how you would decide between a single-channel protocol and a connectivity protocol. What would a qEEG normative database need to show you before you committed to the connectivity approach?

Glossary

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