Evaluation

Selye's model excluded psychological factors like appraisal. His assumption that stress involves nonspecific physiological changes is incorrect. All stressors do not produce a uniform endocrine stress response. Instead, stressors can change multiple body systems in response to stressor intensity, biological predisposition, cognitive appraisal, perceived support, and emotional response (Mason, 1971, 1975; McEwen, 2005; Taylor, 2021). Finally, whereas Selye proposed that our stress responses attempt to maintain physiological processes within a narrow optimal range, current research focuses on adaptation (e.g., allostatic load model) instead of set points (Brannon et al., 2022).

The Stress Response Is Multidimensional

The human stress response is multidimensional and involves diverse systems, ranging from the central nervous system to the immune system. Each person uniquely responds to stressors. This individualized pattern is called response stereotypy. Individuals differ in the systems impacted, their activation or suppression, and how these changes affect health. While stressors can produce system-wide macroscopic changes like increased blood pressure, they can also cause epigenetic changes that alter DNA expression. Graphic © Sergey Nivens/Shutterstock.com.

Sympathetic and Parasympathetic Responders

Two modal patterns of autonomic response to stressors have been observed. A sympathetic responder may increase blood pressure, heart rate, and sweat gland activity and decrease heart rate variability and peripheral blood flow. These changes may result from increased sympathetic activation, decreased parasympathetic activation called vagal withdrawal, or a combination of both.




Listen to a mini-lecture on Sympathetic and Parasympathetic Responders © BioSource Software LLC.

In contrast, a parasympathetic responder may increase digestive activity, constrict the lungs' alveoli, and faint from low blood pressure.

Individuals can react to stressors with elements of both autonomic patterns. For example, they can increase heart rate and blood pressure (sympathetic) while experiencing the gastrointestinal symptoms of diarrhea and gas (parasympathetic).

Stephen Porges proposes that vagal withdrawal is our modal response to everyday stressors. Real Genius is drawn by Dani S@unclebelang.

Biopsychosocial Model

Engel's (1977, 1980) biopsychosocial model proposes that the complex interplay of psychological, biological, and sociological factors results in health or illness.



In this model, stress is a psychological risk factor that affects and is influenced by an individual's biology and sociology. The biopsychosocial model challenges the biomedical model that illness is primarily due to biological abnormalities, which has influenced medical practice since the 1700s (Taylor, 2012). A clinician who adopts the biopsychosocial model assumes that health depends on all three factors. When treating illness, these factors must be assessed and addressed using an interdisciplinary approach (Schwartz, 1982).

The illustration below was adapted from Brannon et al. (2022).



Evidence supporting the biopsychosocial model includes increased psychological and medical disorders in divorced and bereaved persons (Schneider, 1984).

Allostatic Load Model

Allostasis, which means achieving stability through change, involves matching type (sympathetic or parasympathetic) and intensity of physiological activation to situational demands (Brannon et al., 2022).

McEwen and Seeman's (1999) allostatic load model proposes that biological responses to stress (e.g., cortisol secretion or elevated glutamate transmission) can harm the body when stressors are chronic or repeatedly occur. Over time, the stress response itself may overwhelm the body's adaptive capacity.

Caption: Bruce McEwen

Stress-Diathesis Model

The stress-diathesis model explains that stressors interact with our inherited or acquired biological vulnerabilities, diatheses, to produce medical and psychological symptoms. From this perspective, disease results when an individual is predisposed to disease and experiences stress. Life event scales like the SRRS and USQ may achieve low predictive validity because they only and incompletely assess stress; they do not evaluate illness vulnerability (Marsland et al., 2001).

The interrelationship among stress, insomnia, and depression illustrates the stress-diathesis model. Sleep reactivity, the tendency for sleep to be disrupted by stress, appears to be a shared vulnerability in insomnia and depression. Greater sleep reactivity produces insomnia, and insomnia contributes to depression. Chronic insomnia can also increase blood pressure, blood sugar levels, and weight (Drake et al., 2014).

System-Wide Effects of Stress

Stress and the Microbiome

The human body primarily comprises single-celled organisms, which outnumber our human cells 1.3-to-1 (Sender et al., 2016). Viruses vastly outnumber bacteria and could sharply increase microbe-to-human cell ratio (Saey, 2016). Most human bacteria are located in our gut. The human microbiome is the collection of microorganisms that reside in our bodies. Graphic © Kateryna Kon/Shutterstock.com.

Stressful Life Events

Cataclysmic Events

Lazarus and Cohen (1977) described cataclysmic events as "sudden, unique, and powerful single life-events requiring major adaptive responses from population groups sharing the experience” (p. 91).

Intentional and unintentional, these events can impact local communities (e.g., mass shootings), geographic regions (e.g., earthquakes, fires, hurricanes, and tsunamis), and the entire planet (e.g., the COVID-19 pandemic. These catastrophes can produce death, dislocation, fear, grief, trauma, and Post-Traumatic Stress Disorder. Graphic © Syda Productions/Shutterstock.com.




Many factors influence survivor response to these powerful stressful events, including perceived discrimination, resources, support, vulnerability to future harm, distance from the devastation, and media coverage. The stressfulness of an event is influenced by geographic proximity, its recency, and whether it was intended. Intentional events are more traumatic than natural disasters because the perpetrators targeted the victims and could do so again (Brannon et al., 2022).

Life Events

Cataclysmic events like a pandemic are so disruptive become they change our lives in various ways: education, employment, exercise, personal, family member, and friend illness, routines, sleep, social interaction, and working conditions.

Life events differ from cataclysmic events in three ways. They affect fewer individuals. They require adjustment, whether positive (e.g., the birth of a child) or negative (e.g., the death of a loved one). Last, they can develop more slowly (e.g., divorce) or suddenly (e.g., injury in a car crash).

Holmes and Rahe (1967) measured major positive and negative life changes using their Social Readjustment Rating Scale (SRRS). The scale lists 43 events, each assigned a different Life Change Unit (LCU) value. They arranged these events in descending order from the death of a spouse (100 LCUs) to minor law violations (11 LCUs). Individuals select the events they have experienced within the last 6 to 24 months. Researchers calculate a stress score by summing the LCU value of the checked events.

Studies that combine prospective (subjects report current events) and retrospective methods (researchers examine subsequent health records) have reported increased illness and accidents following increased stressful events (Johnson, 1986; Rahe & Arthur, 1978). However, the correlation between SRRS scores and disease is around + 0.30 (Dohrenwend & Dohrenwend, 1984), which means that the SRRS accounts for only 9% of the variance in disease.




Listen to a mini-lecture on the Effects of Major Life Changes and Hassles © BioSource Software LLC.

The SRRS has received severe criticism, and its popularity has declined. Critics have argued that its positive events can reduce the risk of illness (Ray, Jefferies, & Weir, 1995). Many individuals who exceed 300 points in a year remain healthy. Scales like the SRRS underestimate African-American life stress (Turner & Avison, 2003). The SRRS assumes that an event impacts all people equally. The wording of some items is vague (e.g., "change in responsibilities at work"). Pessimism can distort recollections of life events (Brett et al., 1990). Finally, the scale does not consider whether an event has been resolved (Turner & Avison, 1992) or an event's controllability or probability (Gump & Matthews, 2000).

The Undergraduate Stress Questionnaire (USQ) developed by Crandall and colleagues (1992) instructs students to select events—mostly hassles—they have experienced during the past two weeks. Higher USQ scores are associated with increased use of health services.

The Perceived Stress Scale (PSS) developed by Cohen and colleagues (1983) measures perceived hassles, major life changes, and shifts in coping resources during the previous month using a 14-item scale. PSS items assess the degree to which respondents rate their lives as unpredictable, uncontrollable, and overloaded (p. 387). The PSS achieves good reliability and validity (Brannon et al., 2022). PSS scores predict cortisol levels (Harrell et al., 1996), fatigue, headache, sore throat (Lacey et al., 2000), and immune changes (Maes et al., 1997).

Hassles and Uplifts

A hassle is a minor stressful event like waiting in a checkout line or experiencing a traffic jam. Hassles can produce illness via several pathways. First, hassles can cause accumulating allostatic load. Second, hassles can amplify the effects of adverse life events and chronic stress (Marin et al., 2007; Serido et al., 2004; Taylor, 2021). Graphic © Dmitry Kalinvosky/Shutterstock.com.




 


An uplift is a minor positive event like receiving an unexpected call from a friend or playing with new puppies. Graphic © Orientgold/Shutterstock.com.





The interaction between hassles and chronic stress is complex. Hassles may increase the psychological distress produced by chronic stress (Serido et al., 2004). Conversely, chronic stress may reduce the effects of hassles by placing them in perspective (McGonagle & Kessler, 1990).

Hardiness

Some individuals do not experience illness or psychological distress when exposed to adverse life events and hassles. Although they may experience brief distress, they generally recover (Lehrer, 2021). Researchers use the concept of hardiness to explain these outliers (Maddi, 2017; Maddi et al., 2017; Pitts et al., 2016; Stoppelbein et al., 2017). Graphic ©lassedesignen/Shutterstock.com.

Psychological Factors in Stress

Traumatic Stress and Post-Traumatic Stress Disorder (PTSD)

Traumatic stress is produced by a highly intense stressor that disrupts coping and endangers ourselves or others. Post-Traumatic Stress Disorder (PTSD) is a severe and long-lasting trauma and stressor-related disorder that often develops within three months of a traumatic event. DSM-5 (APA, 2013) divides its symptoms into four clusters: intrusion, avoidance, negative alterations in cognitions and mood, and alterations in arousal and reactivity. Graphic © John Gomez/Shutterstock.com.




The exposure can also be second-hand, such as witnessing domestic violence or learning about a family tragedy (Crider, 2004; Lamprecht & Sack, 2002). The lifetime prevalence of adult PTSD in the United States is about 6.8% (Kessler et al., 2005).

Negative Affect States and Affectivity

From a mindfulness perspective, we should not consider emotional responses like depression as negative but as difficult emotions. A clinician would encourage clients to accept depressed feelings as part of themselves without judgment and to focus on specific behaviors that can be changed (Khazan, 2013).

Stressors can trigger complex adjustments, including challenging affective states (anxiety) and corresponding psychophysiological changes (decreased HRV). Barrett and Russell's (1998) structural model represents each affective state within a circumplex based on its degrees of affective valence (unpleasant to pleasant) and affective intensity (activation to deactivation). Affective states fall inside or along the surface of this circular structure. Diagram adapted from Russell and Barrett (1999).

Anxiety

A Framingham study report by Markovitz et al. (1993) showed that men with elevated anxiety had twice the risk of middle-age hypertension as men with lower anxiety. This increased risk was not found for women. A prospective study by Kawachi et al. (1994) revealed that men diagnosed with phobic anxiety had a three times greater risk of sudden cardiac death. Albert et al. (2005) found that women diagnosed with phobic anxiety had a 59% greater risk of sudden cardiac death and a 31% greater risk of fatal coronary heart disease than women who scored low. These increased risks were associated with risk factors such as diabetes, hypertension, and high cholesterol. Graphic © Malochka Mikalai/ Shutterstock.com.

Depression

Pratt et al. (1996) reported that depressed individuals had a four times greater risk of a heart attack in the next 14 years than non-depressed individuals. Frasure-Smith et al. (1995) found that depressed heart attack patients had a four times greater risk of another heart attack in the next 18 months than non-depressed heart attack patients. Carney et al. (2005) discovered that depressed heart attack patients were almost three times more likely to die over 30 months than non-depressed heart patients. Decreased heart rate variability accounted for a significant share of the increased risk of death.

Acute and Chronic Stress Responses

Acute Stress

Cannon's fight-or-flight response focuses on sympathetic nervous system responses to an acute stressor and describes the sympathetic-adrenomedullary (SAM) pathway that releases the hormones epinephrine and norepinephrine. Selye's General Adaptation Syndrome (GAS) describes our prolonged response to a chronic stressor across three stages. The GAS summarizes changes in the hypothalamic-pituitary-adrenal (HPA) axis, which releases the hormones CRH, ACTH, and cortisol, and explains how chronic stress can produce disease and death.

The Fight-or-Flight Response

Cannon (1932) described the fight-or-flight response, in which an individual confronts or flees a stressor. During an acute stress response, which corresponds to the end of Selye's alarm stage, we activate the sympathetic nervous system (SNS), increasing respiration, cardiac output, blood flow to skeletal muscles, and metabolism while decreasing digestion and the reproductive system activity. The SNS, in turn, activates the hard-wired sympathetic-adrenomedullary (SAM) pathway, resulting in the release of the hormones epinephrine and norepinephrine by the adrenal medulla (inner adrenal gland).




Listen to a mini-lecture on the SAM Pathway © BioSource Software LLC. The acute stress response is illustrated © Designua/Shutterstock.com.








The adrenal medulla releases epinephrine and norepinephrine in a 4:1 ratio (Fox, 2019). The adrenal medulla is the inner region of the adrenal glands located at the top of each kidney. Graphic © Designua/Shutterstock.com.

The Hypothalamic-Pituitary-Adrenal (HPA) Axis

The hypothalamic-pituitary-adrenal (HPA) axis is the second stage of the body's defense. The HPA axis is the foundation of allostasis and its failure, allostatic overload (McEwen, 2002). The HPA axis releases the hormones CRH, ACTH (corticotropin), and cortisol. This cascade starts with signals from the amygdala to the hypothalamus and ultimately targets the adrenal glands located at the top of each kidney.




Listen to a mini-lecture on the HPA Axis © BioSource Software.

The adrenal cortex, which is the adrenal gland's outer region, produces the hormones aldosterone and cortisol. Cortisol is the most important glucocorticoid. Cortisol levels peak from 20-40 minutes following a stressor (Brannon et al., 2022).

Chronic Stress

General Adaptation Syndrome (GAS)

The General Adaptation Syndrome (GAS) was Selye’s (1956) three-stage model of chronic autonomic and endocrine system responses to stressors.




Listen to a mini-lecture on the General Adaptation Syndrome © BioSource Software LLC.


Selye argued that diverse stressors produce a three-stage response (alarm, resistance, and exhaustion) in all subjects. In this model, a cold stressor is interchangeable with a shock stressor because they produce the same autonomic and endocrine responses. Whereas Cannon showed that acute stress could change the functions of our internal organs, Selye mainly demonstrated using animal models that chronic stress can change their structure (Crider, 2004).

Alarm is the first stage of Selye’s GAS and consists of shock and countershock phases. The shock phase includes reduced body stress resistance and increased autonomic arousal and hormone release (ACTH, cortisol, epinephrine, and norepinephrine) that comprise the fight-or-flight response. In the countershock phase, resistance increases due to local defenses.

Resistance is the second stage of Selye’s General Adaptation Syndrome. Local defenses have made the generalized stress response unnecessary. Both cortisol output and stress symptoms, like adrenal gland enlargement, decline. While the person appears to be normal, adaptation to the stressor places mounting demands on the body, leading to diseases of adaptation like hypertension as adaptation energy is depleted. Local defenses will break down if stressors persist. McEwen calls these adjustments allostatic load.

Recovery or Exhaustion is the third stage of Selye’s General Adaptation Syndrome. We recover when a stressor has ended, and we can restore homeostasis. In exhaustion, increased endocrine activity depletes body resources and raises cortisol levels, resulting in suppressed immunity and stress syndrome symptoms. Selye believed that these changes could cripple immunity and cause bronchial asthma, cardiovascular disease, depression, hypertension, hyperthyroidism, peptic ulcer, ulcerative colitis, and possibly death (Brannon et al., 2022).

While Selye conceptualized stress as the outcome of the three-stage GAS, stress may occur at any time. Individuals may experience stress-related changes in anticipation of an event or after it has ended. Chronic resistance appears to produce more significant harm than exhaustion (Taylor, 2021).

Graphic adapted from Biostrap.

Brain Structures Involved in Stress

The amygdala, hypothalamus, hippocampus, and prefrontal cortex are the four brain structures most important to the stress response.

Psychoneuroimmunology (PNI)

Nonspecific and Specific Immune Mechanisms

The human body utilizes nonspecific and specific immune mechanisms to protect itself against invading organisms, damaged cells, and cancer.

The main nonspecific, or innate, mechanisms are relatively rapid in response and include anatomical barriers (skin and mucous membranes) and phagocytosis (ingestion of microorganisms) by macrophages and neutrophils. Natural killer cells and neutrophils destroy infectious agents. Nonspecific mechanisms release antimicrobial agents (hydrochloric acid, interferons, and lysozymes) and signal to other immune responders. Local inflammatory responses confine microbes, allowing white blood cells and other immune cells to attack them.

In contrast, specific or adaptive immune mechanisms are slower and aim to target invaders and diseased cells selectively. The specific immune response involves the production and proliferation of antibodies by B cells, targeted destruction of foreign material by T cells, and preparation for future infiltrations of the same antigen (Shaffer & Mannion, 2016). In the illustration below, macrophages arrive to protect endangered cells. Graphic © UGREEN 3S/Shutterstock.com.

Cognitive Appraisal of Stressors and Coping

Lazarus and Folkman's (1984) Transactional Model of Stress has more strongly influenced psychologists than Selye's General Adaptation Syndrome.

Caption: Richard Lazarus

Whereas Selye's stimulus model theorized that events determine stress, Lazarus' cognitive model proposed that stress is determined by our perception of the situation and emphasizes person-environment fit (Taylor, 2012).

Coping is central to the Transactional Model of Stress. Lazarus and Folkman defined it as "constantly changing cognitive and behavioral efforts to manage specific external and/or internal demands that are appraised as taxing or exceeding the resources of the person” (1984, p. 141). Coping is an effortful learned process whose goal is to manage a situation (Brannon et al., 2022).




Listen to a mini-lecture on the Transactional Model of Stress © BioSource Software LLC.

Perception of Stressors Impacts Health

Cannon (1942) studied voodoo deaths in shamanistic cultures and published speculation about how beliefs can produce lethal cascades of system failure in the American Anthropologist. Graphic © Fer Gregory/Shutterstock.com.



Keller et al. (2012) conducted a regression analysis of the 1998 National Health Interview Survey and prospective National Death Index data from 28,753 adults. Both elevated levels of reported stress and the perception that stress negatively impacted health independently and jointly predicted poor physical and psychological health outcomes. While neither variable independently predicted premature death, subjects who reported high levels of stress and the perception that stress impacts health experienced a 43% greater risk of early death.




Listen to a mini-lecture on the Effect of the Perception That Stress Harms Health © BioSource Software LLC.

Check out Kelly McGonigal's TED Talk How to Make Stress Your Friend. McGonigal proposes that we train clients to reframe sympathetic activation as evidence of our courage to rise to a challenge instead of the body injuring itself.





Real Genius was drawn by Dani S@unclebelang.

Personality Dimensions

The personality dimensions of mastery, impulsiveness, hopelessness, optimism, and pessimism, and constructs like alexithymia and reactivity are important to understanding stress responses.

Resources to Buffer Stress

Social Support

Social support consists of received support (support provided) and perceived support (expected support) from individuals and organizations. Both forms of social support include informational, material, and psychological assistance from others. The value of each kind of social support depends on an individual's specific needs. Graphic © Yuri/iStockphoto.com.

Prosocial Behavior

A prospective study of 846 adults by Poulin et al. (2013) showed that providing tangible assistance to friends or family members in the previous year buffered the effects of stress on mortality over 5 years. In contrast, stress predicted mortality for participants who did not help others. Real Genius was drawn by Dani S@unclebelang.




Follow-up data from the longitudinal National Survey of Midlife Development in the U.S. (MIDUS II) of 1054 middle-aged adults found that women who perceived that they supported others in positive social relationships had lower IL-6 levels, a marker for inflammation (Jiang et al., 2021). Elevated IL-6 levels are linked to an increased risk for serious diseases.




Listen to a mini-lecture on the importance of prosocial behavior © BioSource Software LLC.

Graphic © Monkey Business img/Shutterstock.com.

Aerobic Exercise

A sedentary lifestyle marked by prolonged sitting and TV-viewing time was linked to chronic health conditions and all-cause mortality in a meta-analysis of data from over 1 million middle-aged men and women. Individuals who engaged in moderate-intensity exercise 60-75 minutes per day eliminated the increased risk of death due to sitting but only attenuated the risk associated with TV-viewing time (Ekelund et al., 2016). Graphic © racornShutterstock.com.






Mildly depressed college women who participated in an aerobic exercise program showed markedly reduced depression compared with those who did relaxation exercises or received no treatment (McCann & Holmes, 1984). Graphic © Maksim Toome/Shutterstock.com.

Religious and Spiritual Activity

In a national health survey financed by the U.S. Centers for Disease Control and Prevention, religiously active people had longer life expectancies (Hummer et al., 1999). McCullough and colleagues (2000) performed a meta-analysis that assigned greater weights to studies that controlled confounding variables like age, gender, health, and social support. They found that religious involvement was linked to a slightly lower mortality rate and that this was not due to social support. Graphic © LincolnRogers/ iStockphoto.com.

BASIC PHYSIOLOGY OF ATTENTION

Brain Organization and Dynamics

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 controlled/informed by selective attention to specific categories of interest.

Graphic uploaded to ResearchGate by Maurizio Corbetta from the The dynamical balance of the brain at rest.



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 an interactive capacity to the smaller networks, also operating within an oscillatory framework.

The central nervous system processes incoming content. Separate regions process specialized content (e.g., auditory, kinesthetic, tactile, visual). Content is shared, integrated, compared to previous content, and analyzed. Decisions are made regarding memory and responses. All of this central nervous system activity occurs within interacting networks linked by electrical/chemical signals. Electrical discharges from network activity are recorded from the scalp surface as the EEG.

Network Overview

The networks most relevant to attention include the executive, affective, motor, oculomotor, and social circuits.

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. The diagrams below were adapted from Alexander et al. (1986) and Thompson and Thompson (2016).




DLPC = dorsolateral prefrontal cortex, PPC = posterior parietal cortex, APA = arcuate premotor area, GPi = globus pallidus internal segment, SNr = substantia nigra, pars reticulata, VA = ventral anterior nucleus of the thalamus, MD = medial dorsal nucleus of the thalamus.

Affective Network

The pre- and subgenual areas of the anterior cingulate cortex (ACC) participate in affective circuits triggered when we make mistakes. The dorsal rostral cingulate zone monitors cognitive activity to predict when errors are likely, and greater executive control may be needed (Thompson & Thompson, 2016).


HC = hippocampal cortex, EC = entorhinal cortex, STG = superior temporal gyrus, ITG = inferior temporal gyrus, posterior parietal cortex, GPi = globus pallidus internal segment, SNr = substantia nigra, pars reticulata, MD = medial dorsal nucleus of the thalamus.

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, 2020; Thompson & Thompson, 2016).




SMA = supplementary motor area, APA = arcuate premotor area, PMC = primary motor cortex, PSC = primary somatosensory cortex, GPi = globus pallidus internal segment, SNr = substantia nigra, pars reticulata, VL = ventral lateral nucleus of the thalamus.

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've experienced in the past (Thompson & Thompson, 2016).



FEF = frontal eye field, DLPFC = dorsolateral prefrontal cortex, PPC = posterior parietal cortex, GPi = globus pallidus internal segment, SNr = substantia nigra, pars reticulata, VA = ventral anterior nucleus of the thalamus, MD = medial dorsal nucleus of the thalamus.

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, 2016).




OFC = orbitofrontal cortex, STG = superior temporal gyrus, ITG = inferior temporal gyrus, ACC = anterior cingulate cortex, GPi = globus pallidus internal segment, SNr = substantia nigra, pars reticulata, VA = ventral anterior nucleus of the thalamus, MD = medial dorsal nucleus of the thalamus.

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 you gaze at this sentence. While the midbrain superior colliculus is mainly implicated in overt attention, it may also regulate covert attention (Breedlove & Watson, 2023).

The Pulvinar Mediates Attentional Shifts

The pulvinar nucleus, which comprises the posterior quarter of the human thalamus, processes visual information and directs attention. The pulvinar plays a pivotal role in processing visual information and shares widespread connections with the cingulate, parietal cortex, and superior colliculus. The pulvinar 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). Graphic © Songkram Chotik-anuchit/Shutterstock.com.




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

Cortical Regions That Guide Attention

The dorsal frontoparietal system, which is comprised of the intraparietal sulcus and frontal eye field, is responsible for 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 towards 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). The dorsal frontoparietal system was created by Simone Vossel, Joy J. Geng, Gereon R. Fink, CC BY 3.0.






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 by 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), which redirects 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.

Salience Network

The salience network comprises structures that seek to monitor our external and internal environments to determine which of these inputs are essential and require further processing and attention. The insula, primarily the anterior insula, is a crucial component of this network 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, 2016). Graphic shared by Nekovarova, Fajnerova, Horacek, and Spaniel under the Creative Commons Attribution 3.0.




The insula, which is a cortical region located within the lateral sulcus, functions as an integrative and organization hub for the salience network. The insula integrates interoceptive awareness, emotional experience, and external perception to facilitate an individual's global perception of the world and its relationship. The insula directs specific networks in processing salient stimuli and the generation of appropriate responses to stimuli (Wiebking & Northoff, 2014).

The insula graphic provided by Fathy, Y.Y., Hoogers, S.E., Berendse, H.W. et al. Brain Imaging and Behavior (2019). https://doi.org/10.1007/s11682-019-00099-3. This file is licensed under the Creative Commons Attribution 4.0 International license.





The insula appears to provide an interface between the human brain's cognitive, homeostatic, and affective systems. It, therefore, provides a link between the areas of the brain involved in monitoring internal signals and those engaged in watching incoming external sensory streams. The insula detects salient events via afferent pathways and switches between other large-scale networks when these events are recognized to guide 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 task-oriented (executive) and default mode (attention) networks (Seeley et al., 2007; Shirer et al., 2012).

Default Mode Network (DMN)

Brain regions are selectively active when we are conscious (Breedlove & Watson, 2020). 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, which are critical elements of creativity. The DMN is relatively inactive when pursuing external goals (Andrews-Hanna et al., 2010).

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

The DMN graphic shows default mode and task-related maps for healthy subjects. On a green background, the default mode network is highlighted in warm colors (red and yellow) and the task-related network is highlighted in cold colors (blue and light blue) depending on the p-value of one sample t-test. Authors are Shim, G., Oh, J. S., Jung, W. H., et al., CC BY 3.0.

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 which involve 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) analyzes others' objectives. The temporoparietal junction (TPJ) constructs theories of mind, which are 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.

Glossary

acute stress response: autonomic changes that occur at the end of Selye’s alarm stage, for example, increased heart rate.

adaptation energy: in Selye’s General Adaptation Syndrome, the capacity of local organs to respond to the demands created by stressors. Depletion of adaptation energy theoretically results in local adaptation syndromes like muscle fatigue and tissue inflammation.

adrenal cortex: the outer region of the adrenal gland that produces the hormone cortisol.

adrenal medulla: the inner region of the adrenal gland that produces the hormones epinephrine and norepinephrine.

affective intensity: in Barrett and Russell’s structural model, a dimension that ranges from activated to deactivated states.

affective network: a network that is triggered when we make mistakes and that monitors cognitive activity to predict when errors are likely, and greater executive control may be needed. The affective network includes the anterior cingulate cortex, hippocampal cortex, entorhinal cortex, superior temporal gyrus, inferior temporal gyrus, posterior parietal cortex, globus pallidus internal segment, substantia nigra, pars reticulata, and medial dorsal nucleus of the thalamus.

affective valence: in Barrett and Russell’s structural model, a dimension that ranges from unpleasant to pleasant affective states.

alarm: the first stage of Selye’s General Adaptation Syndrome that consists of shock and countershock phases.

allostasis: the maintenance of stability through change by mechanisms that anticipate challenge and adapt through behavior and physiological change.

allostatic load model: McEwen and Seeman’s hypothesis that when stressors are acute or repeatedly occur, biological responses to stress can harm the body.

amygdala: limbic system structure that participates in evaluating whether stimuli are threatening, establishing unconscious emotional memories, learning conditioned emotional responses, and producing anxiety and fear responses.

arousal: overall level of alertness.

attention: the selection of sensory information or cognition for enhanced processing.

attentional bottleneck: a filter that limits enhanced processing to only the highest priority stimuli.

attentional spotlight: a shift of selective attention to choose stimuli for enhanced processing.

biofeedback: (1) learning process that teaches an individual to control her physiological activity, (2) biofeedback training aims to improve health and performance, (3) instruments rapidly monitor an individual's performance and display it back to her, (4) the individual uses this feedback to produce physiological changes, (5) changes in thinking, emotions, and behavior often accompany and reinforce physiological changes, and (6) these changes become independent of external feedback from instruments. Information about psychophysiological performance is obtained by noninvasive monitoring and used to help individuals achieve self-regulation through a learning process that resembles motor skill learning.

biopsychosocial model: Engel’s perspective that the complex interplay of psychological, biological, and sociological factors results in health or illness.

catecholamines: chemical compounds containing catechol and amine groups, like dopamine, epinephrine, and norepinephrine, derived from the amino acid tyrosine.

central executive network: structures including the dorsolateral prefrontal cortex, anterior cingulate cortex, and orbitofrontal cortex responsible for cognitive regulation of behavior.

central nucleus of the amygdala: the region of the amygdala that orchestrates the amygdala’s response to stressors. For example, it activates the paraventricular nucleus (PVN) of the hypothalamus, resulting in increased CRH release to the pituitary gland.

challenge: in Lazarus and Folkman’s Transactional Model of stress, an individual’s assessment during primary appraisal of her potential to cope with an event and gain from this opportunity.

cingulate cortex: cortex which lies above the corpus callosum that is responsible for the motivation dimension of attention, like pain due to physical injury and social rejection.

conditioned response (CR): in classical conditioning, a response (blood pressure rise) elicited by a conditioned stimulus (criticism).

conditioned stimulus: in classical conditioning, a stimulus (dentist's office), that in association with an unconditioned stimulus (pain), elicits an unconditioned response (anxiety) like the original unconditioned response.

corticotropin (ACTH): the hormone released by the pituitary gland following CRH binding that triggers cortisol release by the adrenal cortex.

corticotropin releasing hormone (CRH): the hormone released by the hypothalamus that triggers ACTH release by the pituitary gland.

cortisol: a glucocorticoid produced by the adrenal cortex that helps convert fat and protein to glucose and reduces inflammation.

countershock phase: the last part of Selye’s alarm stage of the General Adaptation Syndrome during which resistance increases due to increased activity by local defenses.

covert attention: an attentional focus independent of sensory orientation.

creative fluency: generating creative ideas like alternative uses for everyday objects.

default mode network: frontal, temporal, and parietal lobe circuits that are active during introspection and daydreaming and relatively inactive when we pursue external goals.

diathesis: inherited or acquired biological vulnerability. For example, obesity is a diathesis for diabetes.

diseases of adaptation: in Selye’s General Adaptation Syndrome, depletion of adaptation energy theoretically results in adaptation syndromes like atrial tachycardia.

distress: Selye’s term for stress due to negative stimuli.

divided attention tasks: situations where subjects must simultaneously process two or more stimuli.

dorsal frontoparietal system: the network comprised of the intraparietal sulcus and frontal eye field responsible for the top-down direction of attention.

dorsolateral prefrontal cortex: the left dorsolateral prefrontal cortex is concerned with approach behavior and positive affect. It helps us select positive goals and organizes and implements behavior to achieve these goals. The right dorsolateral prefrontal cortex organizes withdrawal-related behavior and negative affect and mediates threat-related vigilance. It plays a role in working memory for object location.

dove strategy: passive, reactive, nonaggressive, and cautious response to stressors.

early selection: filtering out lower-priority competing stimuli before preliminary perceptual and semantic analysis.

endogenous attention: voluntary attention that directs the attentional spotlight to support cognitive system priorities.

epinephrine: adrenal medulla hormone that increases muscle blood flow, converts stored nutrients into glucose for use by skeletal muscles, and initiates cardiac muscle contraction when it binds to β1 receptors.

eustress: Selye’s term for stress due to positive stimuli.

executive network: a network responsible for allocating attention, cognitive inhibition, behavioral inhibition, working memory, and cognitive flexibility. The executive network includes the dorsolateral prefrontal cortex, posterior parietal cortex, arcuate premotor area, globus pallidus internal segment, substantia nigra, pars reticulata, ventral anterior nucleus of the thalamus, and medial dorsal nucleus of the thalamus.

exhaustion: the third stage of Selye’s General Adaptation Syndrome during which increased endocrine activity depletes body resources and raises cortisol levels resulting in suppressed immunity, stress syndrome symptoms, and possible hospitalization and death.

exogenous attention: involuntary reflexive attention that redirects attention based on the novelty or importance of incoming stimuli.

fight-or-flight response: Cannon’s response of confronting or fleeing a threat that occurs at the end of Selye’s alarm stage.

frontal eye field (FEF): region of the premotor cortex that directs gaze towards targets selected by the IPS.

General Adaptation Syndrome (GAS): Selye proposed that diverse stressors produce a three-stage (alarm, resistance, and exhaustion) autonomic and endocrine response in all subjects.

harm: in Lazarus and Folkman's Transactional Model of Stress, damage that has already occurred. For example, a heart attack survivor may perceive harm as damage to the heart muscle.

hassle: minor stressful event, for example, waiting in a checkout line.

Hassles and Uplifts Scale: DeLongis, Folkman, and Lazarus’s revised 53-item scale measured hassle frequency and intensity. This scale better predicted headache frequency and intensity and inflammatory bowel disease frequency than the Social Readjustment Rating Scale (SRRS).

Hassles Scale: Kanner and colleagues’ 117-item scale that measures negative daily experiences.

hawk strategy: a proactive and bold response to stressors.

heart rate variability (HRV): beat-to-beat changes in heart rate, including changes in the RR intervals between consecutive heartbeats.

hippocampal formation: the limbic structure involved in controlling the endocrine system’s response to stressors, formation of explicit memories, and navigation. Cortisol binding to this structure disrupts these functions, interferes with creating new neurons, and harms and kills hippocampal neurons.

hypothalamic-pituitary-adrenal (HPA) axis: the hormonal cascade that starts with signals from the amygdala to the hypothalamus and ultimately targets the adrenal glands, releasing the hormones CRH, ACTH (corticotropin), and cortisol.

hypothalamus: a forebrain structure located below the thalamus that dynamically maintains homeostasis by controlling the autonomic nervous system, endocrine system, survival behaviors, and interconnections with the immune system.

insula: the cortical region located within the lateral sulcus of the frontal, parietal, and temporal lobes that functions as an integrative and organization hub for the salience network.

intraparietal sulcus (IPS): the region of the parietal lobe that provides voluntary top-down steering of attention.

late selection: filtering out competing stimuli after performing extensive analysis.

locus of control: Rotter’s concept of a continuum of control of outcomes where internals attribute outcomes to their efforts and externals attribute them to external events. This concept overlaps with the concepts of mastery, perceived control, and self-efficacy.

mastery: the relatively stable expectancy that we can control our personal outcomes. This concept overlaps with the concepts of locus of control, perceived control, and self-efficacy.

microbiome: the collection of the microorganisms that reside in the human body.

motor network: the network that plans, initiates and inhibits voluntary movements and muscle contractions. The motor network includes the supplementary motor area, premotor cortex, primary motor cortex, primary somatosensory cortex, cerebellum, arcuate premotor area, globus pallidus internal segment, substantia nigra, pars reticulata, and ventral lateral nucleus of the thalamus.

myelinated vagus: in Porges' polyvagal theory, the phylogenetically newer ventral vagal complex that rapidly adjusts cardiac output and promotes social engagement.

negative affective states: in Barrett and Russell’s structural model, unpleasant states like sadness are located in the left hemisphere.

negative affectivity: a predisposition toward distress and dissatisfaction. Individuals who are rated high on this trait negatively perceive themselves, others, and the environment and have a pessimistic perspective.

neurofeedback: information about EEG activity obtained by noninvasive monitoring and used to help individuals achieve self-regulation through a learning process that resembles motor skill learning.

neuroplasticity: the ability of neurons and their networks to remodel themselves in response to experience.

norepinephrine: an adrenal medullary hormone that increases muscle blood flow and converts stored nutrients into glucose for skeletal muscles.

nucleus (PVN) of the hypothalamus: when activated by the central nucleus of the amygdala, this nucleus releases CRH to the pituitary gland.

oculomotor network: a network that programs and initiates voluntary eye movements, inhibits eye movements toward distracting stimuli, and allows us to return our focus to locations we've experienced in the past. The oculomotor network includes the frontal eye field, dorsolateral prefrontal cortex, posterior parietal cortex, caudate, globus pallidus internal segment, substantia nigra, pars reticulata, ventral anterior nucleus of the thalamus, and medial dorsal nucleus of the thalamus.

overt attention: the agreement between attentional focus and sensory orientation.

oxytocin: a hormone and neurotransmitter that may contribute to social bonding, anxiety following exposure to stressors, and the milk letdown reflex.

palliation: in Lazarus and Folkman’s Transactional Model of Stress, secondary appraisal can lead to efforts to reduce our stress response rather than attack the stressor.

parasympathetic responder: a response stereotypy in which an individual may increase digestive activity, constrict the lungs' alveoli, and faint from low blood pressure when challenged by stressors.

perceived control: an individual’s expectancy that she can influence her outcomes. This concept overlaps with the concepts of locus of control, mastery, and self-efficacy.

Perceived Stress Scale (PSS): Cohen and colleagues’ scale that measures perceived hassles, major life changes, and shifts in coping resources during the previous month using a 14-item scale. PSS items assess the degree to which respondents rate their lives as unpredictable, uncontrollable, and overloaded.

perceptual load: stimulus processing demands.

pituitary gland: the endocrine gland found at the base of the skull that is divided into the anterior pituitary, which secretes the tropic hormones adrenocorticotropic hormone (ACTH), thyroid-stimulating hormone (TSH), follicle-stimulating hormone (FSH), luteinizing hormone (LH), prolactin, and growth hormone (GH), and the posterior pituitary that releases oxytocin or vasopressin produced by the hypothalamus.

polyvagal theory: Porges' theory that the unmyelinated vagus (dorsal vagus complex) and newer myelinated vagus (ventral vagal complex) mediate competing adaptive responses.

positive states: in Barrett and Russell’s structural model, pleasant states like contentment are located in the right hemisphere.

posttraumatic stress disorder (PTSD): severe and long-lasting trauma and stressor-related disorder that often develops within three months of a traumatic event and may include re-experiencing a traumatic event, avoidance of stimuli associated with the trauma, numbing of responsiveness, and hyperarousal.

prefrontal cortex: the most anterior region of the frontal lobes divided into orbitofrontal and ventromedial, dorsolateral prefrontal cortex, and anterior and ventral cingulate cortex subdivisions, and is responsible for the brain’s executive functions.

primary appraisal: in Lazarus and Folkman’s Transactional Model of Stress, the first stage of our response to an event during which we categorize the consequences of events as positive, neutral, or negative and determine whether an event is relevant and negative, or potentially harmful. We evaluate these events for their possible harm, threat, or challenge.

psychoneuroimmunology: Solomon and Moos’ term for a multidisciplinary field that studies the interactions between behavior and the nervous system, endocrine system, and immune system.

pulvinar nucleus: the posterior region of the thalamus that processes visual information and directs attention.

raphe system: a network of serotonergic neurons located in the brainstem that includes anxiety-producing and anxiety-reducing pathways that terminate on the hippocampus.

reappraisal: in Lazarus and Folkman’s Transactional Model of Stress, secondary appraisal can modify our perception of a threat when direct action is impractical or unsuccessful.

resistance: the second stage of Selye’s General Adaptation Syndrome where local defenses have made the generalized stress response unnecessary. Both cortisol output and stress symptoms, like adrenal gland enlargement, decline.

response stereotypy: a person’s unique response pattern to stressors of identical intensity.

salience network: structures including the insula and anterior cingulate cortex that seek to monitor our external and internal environments to determine which of these inputs are salient and require further processing and attention.

secondary appraisal: in Lazarus and Folkman’s Transaction Model of Stress, the second stage of our response to an event during which we evaluate whether our coping abilities and resources can surmount an event's harm, threat, or challenge.

self-efficacy: an individual’s expectancy that they can influence her outcomes. This concept overlaps with the concepts of locus of control, mastery, and perceived control.

shock phase: the first part of Selye’s alarm stage of the General Adaptation Syndrome that includes the reduced body stress resistance and increased autonomic arousal and hormone release (ACTH, cortisol, epinephrine, and norepinephrine) that comprise the “fight-or-flight” response.

social network: the network that mediates socially responsible behavior, empathy, behavioral inhibition, emotional regulation, and sound judgment. The social network includes the orbitofrontal cortex, superior temporal gyrus, inferior temporal gyrus, anterior cingulate cortex, caudate, globus pallidus internal segment, substantia nigra, pars reticulata, ventral anterior nucleus of the thalamus, and medial dorsal nucleus of the thalamus.

Social Readjustment Rating Scale (SRRS): Holmes and Rahe measured major positive and negative life changes using their Social Readjustment Rating Scale (SRRS), which lists 43 events, each assigned a different Life Change Unit (LCU) value.

stress: Selye’s term for a nonspecific response to stimuli called stressors.

stress-diathesis model: the view that stressors interact with our inherited or acquired biological vulnerabilities, diatheses, to produce medical and psychological symptoms.

stressors: Selye’s term for stimuli that elicit the stress response.

structural model: Barrett and Russell's representation of affective states within a circumplex (circular structure) based on its degrees of affective valence (unpleasant to pleasant) and affective intensity (activation to deactivation).

superior colliculus: the dorsal midbrain structure composed of gray matter that processes visual information, directs visual gaze and visual attention to selected stimuli, and participates in overt and covert attention.

sympathetic nervous system (SNS): autonomic nervous system branch that regulates activities that expend stored energy, such as when we are excited.

sympathetic responder: a response stereotypy in which an individual may increase blood pressure, heart rate, and sweat gland activity and decrease heart rate variability and peripheral blood flow pressure when challenged by stressors.

sympathetic-adrenomedullary path (SAM) pathway: during an acute stress response, the sympathetic nervous system directs the release of the hormones epinephrine and norepinephrine by the adrenal medulla.

telomerase: an enzyme that adds DNA to telomeres. Telomerase levels decline with chronic stress and cellular aging.

telomere: DNA and protein that cover the ends of chromosomes. Telomeres shorten with chronic stress and cellular aging.

temporoparietal junction (TPJ): the intersection of the superior temporal gyrus and inferior parietal lobe that mediates bottom-up shifts in attention in response to stimulus attributes.

tend-and-befriend response: Taylor and colleagues’ proposition that women may tend (nurture others) and befriend (seek and provide social support) in response to stressors.

threat: in Lazarus and Folkman’s Transactional Model of Stress, we evaluate events for the damage they could inflict in the future during primary appraisal.

Transactional Model of Stress: Lazarus and Folkman’s cognitive model proposed that stress is determined by our perception of the situation.

traumatic stress: stress produced by a highly intense stressor that disrupts coping and endangers ourselves or others.

unmyelinated vagus: in Porges' polyvagal theory, the phylogenetically older dorsal vagus complex that responds to threats through immobilization, feigning death, passive avoidance, and shutdown.

uplift: a minor positive event like receiving an unexpected call from a friend.

vagal withdrawal: the inhibition of the myelinated vagus, often by daily stressors.

ventromedial prefrontal cortex: region of the prefrontal cortex may play a role in calculating risk and the emotional responses of anxiety and fear. Cortisol binding to this structure increases anxiety and fear and disrupts and kills neurons.

Yerkes-Dodson curve: an inverted U-curve that illustrates the relationship between pressure and performance.

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Assignment

Now that you have completed this module, how has it changed your understanding of stress? Explain how the top-down and bottom-up attentional systems work together to direct our focus.

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