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Neuroscience Breakthroughs Since Graduate School - Part 7: Emotion

Updated: Mar 2

Neuroscience research in the last decades has fundamentally changed our understanding of emotion. Bottom-up and top-down pathways generate emotions. The gut-brain axis regulates emotions and overall mental well-being. There are no "happiness" or "sadness" centers. Instead, emotions involve different activation patterns in structures comprising affective networks. The amygdala's role in emotion has been expanded from anxiety and fear to detecting salient stimuli (e.g., reinforcers and threats). Mindfulness research has challenged the popular belief that we should be able to dismiss difficult emotions. Efforts to control the uncontrollable exhaust of our finite glucose stores and reduce our ability to select adaptive responses. Finally, the intra-individual variation (IIV) perspective has challenged trait explanations of emotion, emphasizing situational factors in hourly and daily emotional variability.

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Emotion is a complex psychological state involving subjective experience, physiological response, and expressive behavior.

Emotions can be defined as organized affective and behavior-promoting reactions to significant internal and external events . . . emotions do more than mobilize behavior. They also often involve the experience of emotional feelings (affect). Emotional feelings may amplify motivation.
Motivation can be defined as the process that determines the direction (i.e. defining which goals an organism seeks to approach or to avoid) and energization (i.e. the mobilization of resources to carry out an action) of behavior (Cromwell et al., 2020, p. 205).

Plutchik's model of emotion, also known as the "Wheel of Emotions," was developed by psychologist Robert Plutchik in 1980. This model proposes that there are eight primary emotions: joy, trust, fear, surprise, sadness, disgust, anger, and anticipation. These primary emotions are grouped into four pairs of opposites: joy-sadness, trust-disgust, fear-anger, and surprise-anticipation. Plutchik suggested we can combine these emotions into more complex and subtle emotions, creating a color-wheel-like structure for understanding human emotions (Plutchik, 1980).

The model also incorporates the concept of emotional intensity, where each primary emotion can vary in intensity. For example, annoyance can intensify into anger and further into rage. The Wheel of Emotions visually represents these emotional relationships, making understanding and identifying emotions in various contexts easier.

Ekman's model of emotions, developed by psychologist Paul Ekman, is an influential theory in the field of emotion research. Although Ekman originally proposed six basic emotions (anger, disgust, fear, happiness, sadness, and surprise), his later work expanded the list to include two additional emotions—contempt and interest (Ekman & Cordaro, 2011).

The model suggests that these eight emotions are universal, meaning they are experienced and recognized across all human cultures and result from evolutionary processes. Ekman states these emotions are associated with distinct facial expressions, physiological responses, and behavioral patterns.

The support for Ekman's model comes from various cross-cultural studies demonstrating the universality of facial expressions associated with basic emotions (Ekman, 1972; Ekman & Friesen, 1971). However, the model has faced some criticism from researchers who argue that the universality of emotions is overstated (Russell, 1994) and that emotions might be more diverse and context-dependent than Ekman's model suggests (Barrett, 2017).

The Illusion of Control

Langer's (1975) illusion of control refers to the tendency of individuals to overestimate their ability to control events and outcomes, particularly in situations where chance or external factors play a significant role. The illusion of control has important implications in various aspects of human life, including decision-making, risk-taking, and stress management. It can lead to overconfidence, resulting in poor choices or increased vulnerability to negative consequences. Additionally, the illusion of control can contribute to the development of unrealistic expectations and the inability to cope with failure. Khazan (2019) cautions that we cannot control thoughts (e.g., don't think about a white bear) or emotions (e.g., anxiety), although people believe that they should. Distraction and suppression tactics do not work in the long run, can increase unwanted thoughts and emotions, and this failure can reduce perceived self-efficacy. Moreover, these futile attempts deplete the glucose needed to fuel adaptive responses.

You use up glucose on tasks that are not under your control, namely, suppressing your difficult thoughts and emotions, and then you do not have adequate resources left to exercise control over tasks that are otherwise controllable: your actions in response to these difficult thoughts and feelings (Khazan, 2019, p. 137).

Khazan (2019) likens struggling with uncontrollable thoughts and feelings to thrashing in quicksand. Instead of flailing and sinking deeper into several feet of water, you can lie flat and paddle to safety. Graphic © Cory Thoman/


Emotions Are Not Moods

Moods are longer-lasting, less intense affective states with no specific cause or object. We simultaneously experience positive and negative moods throughout our waking days.

Moods and emotions are closely related psychological phenomena but differ in duration, specificity, and cognitive appraisal.


Emotions are typically short-lived, intense responses to specific events or situations. In contrast, moods are longer-lasting, more diffuse states of feeling that may not be tied to a particular cause (Gross, 2015).

According to Ekman's (1993) research on emotions, basic emotions like happiness, sadness, anger, surprise, fear, and disgust typically last for a short duration, ranging from a few seconds to several minutes. Frijda's (1988) work suggests that emotions can last longer if continuous or recurring stimuli maintain the emotional state.

Moods, which are more diffuse and longer-lasting than emotions, can last for hours, days, or even longer. Personality traits and psychological adjustment can influence mood variability (Kuppens et al., 2007). Individual differences in personality traits, such as neuroticism and extraversion, can influence the duration and intensity of moods. People with higher levels of neuroticism may experience negative moods for longer periods, while those with higher levels of extraversion might have more positive and enduring moods (David et al., 1997).


Emotions are generally more specific and have a clear object, such as feeling angry at someone, whereas moods are more general and do not necessarily have a specific target (Gross, 2015).

Cognitive Appraisal

Cognitive appraisal refers to the process by which an individual evaluates and interprets an event or situation in relation to their well-being, goals, and values. This evaluation influences the individual's emotional response to the event or situation. Emotions involve cognitive appraisals of a situation, whereas moods do not necessarily involve such appraisals (Scherer, 2005).

How Moods Relate to Emotions

Moods can affect emotions in several ways:

Mood congruency: Research has shown that people tend to experience emotions that are congruent with their current mood state (Bower, 1981; Forgas, 1995). For example, suppose someone is in a positive mood. In that case, they are more likely to experience positive emotions like happiness or excitement. They are more likely to experience negative emotions like sadness or anger if they are in a negative mood.

Mood-dependent memory: Moods can also affect the emotions we experience by influencing the memories we retrieve. According to the mood-dependent memory hypothesis, people are more likely to recall information that is consistent with their current mood (Eich, 1995). This means that someone in a sad mood may be more likely to remember sad events, which could trigger negative emotions.

Cognitive appraisal: Moods can also affect emotions through their influence on cognitive appraisal processes. Appraisal theory posits that emotions arise from our interpretations and evaluations of events (Lazarus, 1991). Our current mood can bias these appraisals, leading us to interpret events in a way that is consistent with our mood. For example, suppose someone is in a negative mood. In that case, they may be more likely to perceive a situation as threatening or harmful, resulting in the experience of negative emotions like fear or anger.

Mood regulation: People may also use mood regulation strategies to change their emotions. According to the process model of emotion regulation, there are several different ways that individuals can regulate their emotions, including situation selection, situation modification, attentional deployment, cognitive change, and response modulation (Gross, 1998). The specific strategies an individual chooses may depend on their current mood and their motivation to maintain or change it.

How Moods Relate to Personality

We cannot disentangle moods from motivation and personality.

. . . the functions of motivation and emotion are underpinned mainly by overlapping, similar locations and networks in the brain, but may differ in specific connections and neurotransmitters that are most central, and, that these functions normally interact with one another, together moving behavior (Cromwell et al., 2020, p. 220).

According to the mood-as-input model (Clore & Huntsinger, 2007; Martin et al., 1993; Martin & Tessler, 1996), moods can serve as a source of information that influences our motivation to engage in different activities. For example, a positive mood may increase motivation to pursue rewarding experiences, while a negative mood may signal the need for self-protection or problem-solving.

The primary tenets of the model are:

Stop rules: Mood influences when individuals decide to stop processing information or engaging in a task. Two types of stop rules guide this: "Feel Good - Stop" and "Feel Bad - Continue." Positive moods signal that the task is complete or satisfactory, while negative moods indicate the need for further effort or information.

Interpretation of mood: The model suggests that individuals use their mood as a source of information, leading to different interpretations based on their current emotional state. Positive moods may result in more optimistic assessments, while negative moods can produce more pessimistic evaluations.

Mood-congruent processing: The Mood-as-Input model also proposes that mood influences memory and information processing by affecting the accessibility of mood-congruent information. People in positive moods are more likely to recall positive information, while those in negative moods are more likely to recall negative information.

Personality can influence an individual's predisposition to experience certain moods. For example, individuals with a high level of neuroticism, a personality trait associated with emotional instability, may be more prone to experiencing negative moods (Costa & McCrae, 1992).

The Stability of Positive and Negative Moods

Jeffrey Vittengl (2004) examined the stability of positive and negative moods in daily life. He found that while positive and negative moods both demonstrated significant day-to-day stability, negative mood was more stable than positive mood. Additionally, the study revealed that positive mood had a greater tendency to return to an individual's baseline level than negative mood. This suggests that negative moods may be more persistent and less influenced by daily events than positive moods.

Another study by Vittengl and colleagues (2009) investigated the effects of cognitive therapy and antidepressant medications on reducing negative mood and increasing positive mood in patients with major depressive disorder. They found that both treatments were effective at reducing negative mood, but cognitive therapy was more effective at increasing positive mood. This research highlights the importance of considering both positive and negative moods when evaluating the effectiveness of depression treatments.

Intra-Individual Variation

Intra-individual variation (IIV) refers to the differences in emotional and mood experiences within an individual over time or across situations. In his manifesto titled "Bringing the Person Back into Scientific Psychology, This Time Forever," Peter C. M. Molenaar, a retired Pennsylvania State University professor, advocated for IIV. He employed mathematical and physical computations to describe the dynamic variability in personality while criticizing conventional psychological assessment approaches.

Understanding IIV is crucial for comprehending the dynamic nature of emotions and mood and informing interventions to improve emotional well-being and mental health.

It is not only that behavioral changes happen often but that they occur from day to day and hour to hour. Someone could be open and agreeable at noon but negative and rigid at two o’clock. Such oscillations in daily feelings and behavior—designated with the bland title of intraindividual variability, or IIV—are, in fact, so great that they rival or even exceed the differences in personality traits such as extroversion or conscientiousness that can be measured between one person and another (Russo, 2023).

Some implications of intra-individual variation for emotion and mood include:

Emotional complexity: IIV highlights the multidimensional nature of emotional experiences, suggesting that individuals can simultaneously experience various, seemingly contradictory emotions. This emotional complexity has been linked to better psychological well-being (Kashdan, Barrett, & McKnight, 2015).

Emotional regulation: IIV in emotions and mood can inform the study of emotion regulation strategies. Individuals may adaptively or maladaptively regulate their emotions depending on context or personal factors. Emotional regulation significantly affects mental health and well-being (Gross, 2015).

Mood variability: High IIV in mood has been linked to an increased risk for psychological disorders such as depression, anxiety, and borderline personality disorder (Houben, Van Den Noortgate, & Kuppens, 2015).

Individual differences: IIV underscores the importance of considering individual differences in emotional experiences, as factors such as personality, genetics, and environmental influences can contribute to unique emotional patterns (Charles, Piazza, Mogle, Sliwinski, & Almeida, 2013).

A 20-year study by Penn State psychologist David Almeida (2002) revealed the importance of daily experiences in shaping long-term emotional, physical, and cognitive outcomes. By examining daily personality variability in over 3,500 adults, the researchers found that immediate circumstances significantly affect short-term psychological states.

Almeida’s team calculated how much of what we typically think of as a personality trait actually is just that or can be understood as a passing emotion. 'We see it in how grumpy people are,' he says. 'We think, Oh, this is a grumpy person. In fact, half of their grumpiness is a personality trait, and half would be within the person’s variability from day to day.' He notes that people with positive traits such as openness or agreeability show only a 30 percent variation in traits such as quickness to anger or worry (Russo, 2023).

Stanford University's Nilam Ram (2009) suggests that emotional fluctuations might reflect different aspects of an individual's personality or response to specific events.

. . . the emotional reports researchers receive from people they observe in their studies are probably a combination of some aspects of the immediate environment and elements of their personality—how reactive they are to what’s around them and how well they regulate their emotions (Russo, 2023).

Researchers have also learned to quantify hour-to-hour variability, which has led to a better understanding of personality traits and the development of new treatments. For instance, Newman (2011) from Penn State has used IIV to create specialized smartphone apps to monitor people's emotions and experiences several times daily. This has led to more nuanced insights into psychological states and has helped therapists tailor treatments to individual patients. While some studies have linked high IIV with neuroticism, others suggest that it may indicate successful adaptation to a tumultuous life. Ultimately, understanding daily emotional fluctuations can provide new insights for helping individuals become the people they want to be.

Emotional Regulation

Emotion regulation is the process by which individuals influence or manage their emotions, both in terms of the intensity and duration of the emotional experience. This can involve the conscious or unconscious modulation of emotional responses to various situations, stimuli, or events. Emotional regulation is an important skill to develop, contributing to overall well-being, social functioning, and mental health.

No Centers for Primary Emotions

There are no "happiness" or "sadness" centers. Instead, emotional regulation is a global integrative process involving the gut-brain axis, amygdala, prefrontal cortex (PFC), anterior cingulate cortex (ACC), and insula.

. . . Each emotion involves differential patterns of activation across the emotion processing network. For example, activity of the cingulate cortex is altered in sadness, happiness, and anger; the left somatosensory cortex is deactivated in both anger and fear. Feelings of regret over costly decisions apparently involves activation of the amygdala and orbitofrontal cortex. Although different emotions are associated with different patterns of activation, there is a lot of overlap among patterns for different emotions (Breedlove & Watson, 2023, p. 511).

Gut-Brain Axis

The gut-brain axis is a complex, bidirectional communication network involving the central nervous system (CNS), the enteric nervous system (ENS), and the gastrointestinal (GI) tract. Research has demonstrated that the gut-brain axis regulates emotions and overall mental well-being.

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 the 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/


The gut microbiota, consisting of trillions of microorganisms, can influence emotional processing and behavior. Dysbiosis or imbalance in the gut microbiota has been linked to anxiety, depression, and other mood disorders. Probiotic interventions have shown the potential to reduce symptoms of these disorders (Cryan & Dinan, 2012).

Stress can disrupt the gut-brain axis, leading to changes in gut microbiota composition and increased intestinal permeability. These alterations can produce or exacerbate mood disorders such as anxiety and depression (Mayer et al., 2015).

Gut peptides and neurotransmitters, such as serotonin, dopamine, and gamma-aminobutyric acid (GABA), play a critical role in mood regulation. Most of the body's serotonin is produced in the GI tract, highlighting the gut's importance in mood regulation (Rea et al., 2016).

The vagus nerve is an essential component of the gut-brain axis, transmitting information between the gut and the brain. Vagal nerve stimulation has been shown to improve mood in individuals with treatment-resistant depression (Bonaz et al., 2018). Vagus nerve graphic © Axel_Kock/

vagus nerve

In conclusion, the gut-brain axis plays a significant role in emotion regulation through its influence on gut microbiota, stress response, gut peptides, neurotransmitters, and the vagus nerve. Dysregulation of this axis can lead to the development or exacerbation of mood disorders, and interventions targeting the gut-brain axis show promise in alleviating symptoms of these disorders.


The amygdala, located deep within the brain's temporal lobes, detects salient stimuli, including threats and reinforcers.

The function of the connection between the amygdala and the PFC is typically conceptualized as a top-down (PFC to amygdala) process of emotion regulation in the service of more adaptive, planful behavior . . . Tottenham and Gabard-Durnam (2017) argue that the amygdala 'instructs the cortex' about what is salient and important (Cromwell et al., 2020, p. 216).

Amygdala graphic © Kateryna Kon/


The amygdala plays a crucial role in emotional processing, particularly in processing and generating fear, anxiety, and other emotions. It comprises two almond-shaped structures, one in each hemisphere of the brain. Research has suggested that there may be functional lateralization of the amygdala, with the left and right amygdalae serving somewhat different roles in emotional processing.

Some studies have suggested that there might be functional differences between the left and right amygdala in processing and regulating emotions. Research has shown that the left amygdala is more involved in processing positive emotions, while the right amygdala is more involved in processing negative emotions. This hypothesis is based on the broader lateralization of emotion processing in the brain, with the left hemisphere generally being associated with positive emotions and the right hemisphere with negative emotions (Davidson, 1992; Davidson & Irwin, 1999).

The right amygdala appears to be more involved in the rapid, automatic processing of emotional stimuli, while the left amygdala is more involved in the slower, more elaborative processing of emotional stimuli (Gläscher & Adolphs, 2003; Sergerie, Chochol, & Armony, 2008). This may be related to the right hemisphere's role in processing holistic, gestalt information, while the left hemisphere is more specialized for processing detailed, analytic information (Van Kleeck, 1989).

The left and right amygdala have different connectivity patterns with other brain regions. The left amygdala is more strongly connected to the prefrontal cortex, which is involved in top-down control of emotions. In contrast, the right amygdala is more strongly connected to the sensory processing areas and the autonomic nervous system, which is responsible for generating emotional responses (Baas, Aleman, & Kahn, 2004; Gainotti, 2012).

Prefrontal Cortex (PFC)

The prefrontal cortex (PFC) is a critical brain region responsible for various higher-order cognitive functions such as decision-making, planning, and emotional regulation. Graphic © Songkram Chotik-anuchit/

Prefrontal Cortex

The left and right prefrontal cortices have been found to play distinct roles in emotional regulation, with the orbitofrontal (OFC), ventrolateral (VLPFC), ventromedial (VMPFC), and dorsolateral (DLPFC) regions contributing significantly to these processes.

The OFC, VLPFC, vmPFC, and dlPFC are essential for emotional regulation. Both the left and right hemispheres contribute to these processes, albeit with different emphases.

Orbitofrontal Prefrontal Cortex

The orbitofrontal cortex (OFC) is a region in the brain's prefrontal cortex, situated just above the orbits (eye sockets). It has been implicated in various cognitive processes, including decision-making, reward and punishment, and emotional regulation. The OFC can be further divided into the left and right hemispheres, associated with different emotional regulation aspects.

The left OFC has been predominantly associated with processing positive emotions and reward-related information. It is thought to play a crucial role in evaluating and predicting the rewarding aspects of stimuli (Kringelbach, 2005). Activation of the left OFC has been linked to feelings of pleasure and the anticipation of positive outcomes (Elliott, Friston, & Dolan, 2000).

The right OFC has been associated with processing negative emotions and punishment-related information. It is thought to be involved in evaluating and predicting aversive stimuli, with activation of the right OFC linked to feelings of fear, anxiety, and punishment anticipation (O'Doherty, Kringelbach, Rolls, Hornak, & Andrews, 2001). In addition, the right OFC has been found to play a role in inhibitory control and response inhibition, which are essential for emotional regulation (Aron, Robbins, & Poldrack, 2004).

The VLPFC is a brain region that is crucial in emotional control and strategy selection. It is located in the lateral prefrontal cortex, responsible for cognitive processes such as working memory, decision-making, and control (Duncan & Owen, 2000). The VLPFC is specifically involved in controlling emotional responses and selecting appropriate strategies to adapt to changing circumstances (Goldin et al., 2008).

One of the key functions of the VLPFC is regulating emotional responses through a process called cognitive reappraisal (Ochsner & Gross, 2005). Cognitive reappraisal involves altering a stimulus's emotional impact by changing how it is perceived or interpreted. The VLPFC is believed to modulate the activity of the amygdala, a region of the brain associated with emotional processing, to regulate emotional responses (Ochsner et al., 2004).

In addition to emotional regulation, the VLPFC plays a significant role in strategy selection. It generates, evaluates, and selects goal-directed actions and strategies essential for adaptive behavior (Badre, 2008). The VLPFC integrates information from various sources, such as sensory inputs, contextual information, and long-term memory, to guide action selection (Koechlin et al., 2003).

The left and right VLPFCs are believed to have some distinct functions. Research suggests that the left VLPFC is more involved in verbal working memory and semantic processing, while the right VLPFC is more involved in visuospatial working memory and inhibitory control (Stuss et al., 2001). However, both hemispheres contribute to emotional control and strategy selection, and their specific roles may depend on task demands and individual differences.

Ventromedial Prefrontal Cortex

The left VMPFC processes and regulates positive emotions and approach-related behavior (Davidson & Irwin, 1999). It has been linked to generating positive emotional responses and integrating emotions with cognitive processes (Bechara et al., 2000). Additionally, the left vmPFC plays a role in reward processing and decision-making (Kringelbach, 2005).

The right VMPFC regulates negative emotions and withdrawal-related behavior (Davidson & Irwin, 1999). Like the left VMPFC, the right VMPFC integrates emotional and cognitive processes (Bechara et al., 2000). Moreover, it contributes to perceiving and understanding others' emotions and social cognition (Schulte-Rüther et al., 2008).

Dorsolateral Prefrontal Cortex

The left DLPFC is implicated in the cognitive aspects of emotional regulation, particularly in the conscious, effortful control of positive emotions (Ochsner & Gross, 2005). It exerts top-down control over limbic and subcortical regions that generate emotional responses, such as the amygdala (Johnstone et al., 2007).

The right DLPFC is involved in the cognitive regulation of negative emotions (Ochsner & Gross, 2005). Like the left DLPFC, it is responsible for top-down control over limbic and subcortical regions, including the amygdala, to modulate emotional responses (Johnstone et al., 2007). The right DLPFC also mediates inhibitory control and self-regulation (Aron, Robbins, & Poldrack, 2004).

In summary, the left and right VLPFC, VMPFC, and DLPFC contribute to emotional regulation. The left and right VLPFC regulate emotion and strategy selection. The left VMPFC regulates positive emotion regulation, while the right VMPFC regulates negative emotions. The left and right DLPFC are responsible for the cognitive control of emotions, with the left DLPFC associated with positive emotions and the right dlPFC with negative emotions.

Top-Down Emotional Control

The orbitofrontal cortex (OFC), ventrolateral prefrontal cortex (VLPFC), dorsolateral prefrontal cortex (DLPFC), and ventromedial prefrontal cortex (VMPFC) interact with one another to modulate emotional responses.

The OFC evaluates emotional stimuli, reward processing, and decision-making (Kringelbach, 2005). It connects with the VMPFC, which is implicated in forming emotional values and regulating affective states (Bechara et al., 2000).

The VLPFC involves cognitive control processes, such as inhibitory control and attentional selection (Aron et al., 2004). It interacts with the DLPFC, which is responsible for working memory, cognitive control, and executive functions (Miller & Cohen, 2001).

The DLPFC modulates the VMPFC during emotion regulation by exerting cognitive control over emotional responses (Ochsner et al., 2012). This interaction enables the integration of cognitive and emotional information, allowing for more adaptive emotional responses.

The VMPFC has reciprocal connections with the amygdala, a key region in generating emotional responses (Price & Drevets, 2012). The VMPFC modulates amygdala activity while regulating emotional responses, enabling the top-down control of emotion (Phelps et al., 2004).

Anterior Cingulate Cortex (ACC)

The anterior cingulate cortex (ACC) is a critical region in the brain involved in various cognitive and emotional processes.

It monitors conflicts and detects emotional salience, helping integrate cognitive control and emotional processing. It is divided into two parts, the left and right ACC, which have been implicated in different aspects of emotional regulation.

The left ACC is predominantly involved in positive affect and approach-related behaviors. At the same time, the right ACC suppresses negative affect and avoidance-related behaviors (Bush et al., 2000; Drevets & Raichle, 1998; Etkin et al., 2011).


The insula is a region of the cerebral cortex, folded deep within the lateral sulcus, which is involved in various functions, including processing and integrating interoceptive information, emotional awareness, and emotional regulation.

The insula can be divided into the left insula and the right insula. While both regions play a role in emotional regulation, they contribute distinctively to this process.

The left insula is primarily associated with positive emotions and approach-related behaviors. A study by Craig (2009) suggests that the left insula processes emotions such as happiness, joy, and love. This region has also been linked to regulating emotions by enhancing the experience of positive emotions and dampening the experience of negative emotions (Phan et al., 2002).

The right insula is more involved in processing negative emotions and withdrawal-related behaviors. Research has shown that the right insula is activated during the experience of emotions such as fear, disgust, and sadness (Paulus & Stein, 2006; Phillips et al., 2003). Furthermore, the right insula has been implicated in the regulation of emotions by amplifying the experience of negative emotions and suppressing the experience of positive emotions (Phan et al., 2002).

Bottom-Up Emotional Control

Bottom-up control of emotion refers to the process by which emotions are generated and regulated through sensory input and physiological responses rather than through cognitive processes (top-down control). In bottom-up control, emotional experiences are shaped by sensory information, such as sights, sounds, and smells, which evoke automatic physiological responses. These responses can then lead to the generation and regulation of emotions.

Several theories and empirical research support the role of bottom-up control in emotion regulation.

In a seminal paper, Zajonc (1980) posits that emotional responses can precede cognitive processes, arguing that some affective reactions can be elicited without conscious awareness or thought. This suggests that bottom-up processes are crucial in emotion generation and regulation.

LeDoux's (1996) work on the neural basis of emotions, particularly fear, has been fundamental in understanding the role of bottom-up control in emotion. His research demonstrates how sensory input can activate the amygdala and produce an emotional response even before the cortex processes the information.

Lang and colleagues (1997) highlight the importance of the automatic activation of attention and physiological arousal in response to emotionally salient stimuli. They propose that the appraisal of a stimulus's emotional significance is integral to the bottom-up control of emotion.

Pessoa's (2008) review paper explores the complex interplay between emotion and cognition, emphasizing the role of bottom-up control in emotion generation. He discusses the neural mechanisms underlying the interaction between sensory input and emotional processing and how they contribute to the overall emotional experience.

Gross' (2015) work on emotion regulation provides a comprehensive overview of individuals' strategies to regulate their emotions, including both top-down and bottom-up approaches. He acknowledges the importance of bottom-up control in the automatic generation of emotional responses and their regulation.

How Heart Rate Variability Aids Emotional Regulation

Researchers have found that heart rate variability is associated with the function of some brain structures involved in emotion regulation, particularly the PFC and the amygdala.

HRV is the organized fluctuation of time intervals between successive heartbeats defined as interbeat intervals" (Shaffer, Meehan, & Zerr, 2020).

Higher HRV has been linked to better emotion regulation and is thought to reflect greater flexibility in adapting to environmental demands. Studies have shown that increased HRV is associated with reduced amygdala reactivity and enhanced PFC function during emotion regulation tasks (Thayer et al., 2012).

HRV plays a crucial role in managing our physiological and emotional responses. Williams and colleagues (2015) discovered a connection between lower HRV and increased challenges in everyday emotion regulation, comprehending personal emotions, and impulse control.

Holzman and Bridgett (2017) studied the link between HRV and top-down self-regulation, which refers to controlling behavioral, cognitive, and emotional processes. In their meta-analysis of 123 studies exploring the relationship between HRV and self-regulation aspects, they concluded that HRV is a strong indicator of our capacity to manage emotions and behavior. HRV impacts self-regulation through two-way communication between the parasympathetic nervous system (vagus nerve) and specific areas of the prefrontal cortex, which participate in emotion regulation and behavior control.

Lane and colleagues (2017) established the connection between HRV and prefrontal cortex activity in the context of emotion regulation.

Sakaki and colleagues (2016) also found a link between higher HRV and enhanced functional connectivity between the amygdala and prefrontal cortex. This suggests that high HRV correlates with an improved ability of the prefrontal cortex to manage the activation of the fight-or-flight response in the amygdala.

Thayer and Lane's Neurovisceral Integration Model

Julian Thayer and Richard Lane's (2000) neurovisceral integration model emphasizes the connection between HRV and the prefrontal cortex. Their model investigates the neural pathways connecting the heart to cortical and subcortical brain structures through the vagus nerve.

Thayer and Lane's model is a theoretical framework emphasizing the central autonomic network's (CAN) role in regulating emotional, cognitive, and behavioral processes. This model was first introduced in a seminal paper by Thayer and Lane in 2000 and has since been refined and expanded in subsequent publications (Thayer & Lane, 2000; Thayer et al., 2009; Thayer & Lane, 2009).

The neurovisceral integration model posits that the CAN, which includes key brain regions like the prefrontal cortex, insular cortex, amygdala, hypothalamus, and brainstem nuclei, plays a crucial role in the integration of information from the body and the environment. The model suggests that the CAN modulates autonomic nervous system (ANS) activity, particularly the balance between the parasympathetic (vagal) and sympathetic nervous systems. Parasympathetic activation, often called vagal tone, is believed to be a key factor in emotional and cognitive regulation (Thayer & Lane, 2000).

According to the model, higher vagal tone is associated with better emotional regulation, cognitive flexibility, and adaptive behavioral responses. Conversely, lower vagal tone is linked to increased vulnerability to stress and negative emotions, reduced cognitive control, and maladaptive behaviors (Thayer et al., 2009).

Studies have provided empirical evidence supporting the associations between HRV and various emotional, cognitive, and behavioral outcomes, in line with the neurovisceral integration model's predictions (Thayer & Lane, 2009).

Thayer and colleagues' (2012) meta-analysis revealed a significant relationship between HRV and increased ventromedial prefrontal cortex and left amygdala blood flow.

How Mindfulness Practices Aid Emotional Regulation

Mindfulness practices, which involve cultivating non-judgmental awareness of the present moment, have been shown to have positive effects on emotional self-regulation. They are thought to influence emotional self-regulation through several mechanisms and brain structures.

Attention regulation: Mindfulness practices enhance the ability to regulate attention and maintain focus on the present moment. This improved attentional control can help individuals disengage from emotionally charged thoughts and feelings, reducing their intensity and impact (Jha, Krompinger, & Baime, 2007).

Body awareness: Mindfulness practices increase interoceptive awareness or the awareness of internal bodily sensations. This heightened body awareness can help individuals recognize and respond more adaptively to emotional states (Farb et al., 2012).

Non-reactivity: Mindfulness promotes a non-judgmental, accepting attitude toward one's thoughts and emotions. This non-reactivity can reduce emotional reactivity and facilitate more adaptive emotional responses (Chambers, Gullone, & Allen, 2009).

Several brain structures have been implicated in the effects of mindfulness on emotional self-regulation.

Prefrontal cortex (PFC): Mindfulness practices have been associated with increased activation and connectivity in the PFC, particularly the dorsolateral and ventromedial regions. These changes may reflect enhanced cognitive control and top-down regulation of emotions (Hölzel et al., 2011).

Anterior cingulate cortex (ACC): Mindfulness training has been linked to increased ACC activation and connectivity, possibly related to improved attention regulation and conflict monitoring during emotional processing (Tang et al., 2015).

Amygdala: Mindfulness practices can lead to decreased amygdala activation in response to emotional stimuli, suggesting reduced emotional reactivity and enhanced emotional self-regulation (Desbordes et al., 2012).

Insula: Mindfulness training has increased insula activation, which may be related to enhanced body awareness and more adaptive processing of emotional information (Farb et al., 2012).

Emotional Memories

Emotional memories are created and stored through a complex interplay between multiple brain regions that work together to process, encode, and retrieve these memories. Key brain structures involved in the formation and storage of emotional memories.

Amygdala: The amygdala plays a crucial role in processing and encoding emotional information, particularly for fear and negative emotions. It helps detect emotionally significant stimuli and assign emotional salience to them. The amygdala also modulates the consolidation of emotional memories by influencing other brain regions, such as the hippocampus and the prefrontal cortex.

Hippocampus: The hippocampus is essential for forming and retrieving episodic memories, including emotional memories. Graphic © Hank Grebe/

The hippocampus helps contextualize emotional experiences within relevant memories and spatial-temporal information. The hippocampus works closely with the amygdala to consolidate and store emotional memories for later retrieval.

Prefrontal cortex (PFC): The PFC, particularly the ventromedial and dorsolateral regions, is involved in the encoding and retrieval of emotional memories. It helps integrate emotional information with cognitive processes and contributes to the conscious recollecting of emotional experiences. The PFC also plays a role in emotion regulation, which can influence the formation and retrieval of emotional memories.

Insula: The insula is involved in processing bodily sensations and emotions, including awareness of one's own emotional state. It may contribute to the encoding and retrieval of emotional memories by integrating sensory information with emotional and cognitive processes.

Anterior cingulate cortex (ACC): The ACC is implicated in attention, conflict monitoring, and emotional processing. It may be involved in modulating emotional memories by assessing the emotional salience of stimuli and directing attention to emotionally significant events.

Emotional Memory Storage

Emotional memories are not stored in a single location but are rather distributed across these interconnected brain regions. The specific aspects of emotional memories, such as the emotional intensity, context, and sensory details, may be encoded and stored in different areas. Their retrieval involves the coordinated activation of multiple regions. The consolidation and retrieval of emotional memories are also influenced by factors such as stress, sleep, and emotion regulation, which can modulate the activity of the involved brain regions.


The amygdala, prefrontal cortex, and insula have been identified as crucial brain areas involved in processing emotions. Further research has revealed the significance of the connections between these areas, such as the connection between the amygdala and prefrontal cortex in emotional regulation (Etkin et al., 2016). Emotions are closely tied to physiological responses, such as changes in heart rate, blood pressure, and hormone levels. The autonomic nervous system (ANS) is critical in regulating these responses.

Recent studies have uncovered the importance of the gut-brain axis and its role in modulating emotions through communication between the enteric nervous system (ENS) and the central nervous system (CNS; Mayer et al., 2015). Cognitive and behavioral strategies are critical in regulating emotions. New research has shown that emotion regulation can be enhanced through techniques such as mindfulness and cognitive reappraisal (Goldin et al., 2013).

Studies of emotions in animals, particularly primates, have provided insights into emotions' evolutionary origins and functions. These studies suggest that emotions are adaptive responses to environmental challenges and opportunities (de Waal, 2010).


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amygdala: a group of nuclei in the brain's medial temporal lobe involved in processing emotions, particularly fear and anxiety, and forming emotional memories.

anterior cingulate cortex (ACC): a region in the brain's medial frontal cortex involved in emotional regulation, conflict monitoring, error detection, and decision-making processes.

bottom-up control of emotion: the influence of sensory and perceptual inputs on emotional experience and regulation, driven by external stimuli and early-stage processing of emotional information.

cognitive appraisal: the process through which individuals evaluate the personal significance of an event, situation, or stimulus, and its relevance to their well-being, influencing the emotional response.

central autonomic network (CAN): a network of brain regions, including the prefrontal cortex, insula, and amygdala, that work together to regulate autonomic functions and emotional experiences.

dorsolateral prefrontal cortex: a brain region within the lateral prefrontal cortex involved in cognitive control, working memory, decision-making, and emotion regulation.

Ekman's model of emotions: a theory proposing the existence of six universally recognized basic emotions: happiness, sadness, anger, fear, surprise, and disgust.

emotion: a complex psychological state involving subjective experience, physiological response, and expressive behavior reacting to a stimulus or event.

emotion regulation: the process of consciously or unconsciously modulating emotional experiences and expressions to meet personal or social goals or adapt to situational demands.

emotional memories: long-term memories associated with emotions, allowing individuals to remember the feelings associated with specific events.

gut-brain axis: the bidirectional communication system between the gastrointestinal tract and the central nervous system, mediated by the vagus nerve, immune system, and microbiome.

heart rate variability: the variation in time between successive heartbeats, a physiological measure reflecting the balance between the sympathetic and parasympathetic nervous systems, and related to emotional regulation and stress.

hippocampus: a region in the medial temporal lobe of the brain, essential for learning, memory consolidation, and spatial navigation, and involved in emotional processing.

intra-individual variation (IIV): the differences in emotional experiences and responses within an individual across different situations, contexts, and time.

insula: a region in the brain's cerebral cortex involved in interoception, self-awareness, and emotional processing, particularly for feelings of disgust and empathy.

Langer's illusion of control: a cognitive bias wherein individuals overestimate their ability to control events or outcomes, often influenced by factors such as perceived skill, choice, or familiarity.

microbiome: the collection of microorganisms, including bacteria, fungi, and viruses, living in and on the human body, significantly influencing health, behavior, and emotions.

mood congruency: the phenomenon where memory retrieval and information processing are influenced by an individual's current mood, leading to the preferential recall of mood-congruent information.

moods: prolonged, diffuse, and less intense emotional states, not necessarily linked to a specific stimulus or event, that influence an individual's perception, cognition, and behavior.

mood-as-input model: a theoretical framework suggesting that moods provide information about the value and utility of ongoing cognitive processes, influencing task persistence based on stop rules.

orbitofrontal cortex (OFC): A region in the prefrontal cortex involved in decision-making, reward evaluation, and emotional regulation, integrating sensory, affective, and cognitive information.

Plutchik's model of emotion: a psychoevolutionary theory proposing eight primary, bipolar emotions: joy-sadness, anger-fear, trust-disgust, and surprise-anticipation, organized in a wheel-like structure.

prefrontal cortex (PFC): a region of the brain located in the anterior portion of the frontal lobes, responsible for higher-order cognitive functions, such as decision-making, social behavior, working memory, and emotion regulation.

top-down control of emotion: the cognitive regulation and modulation of emotional responses, driven by higher-order brain regions such as the prefrontal cortex, which can influence and modify emotional processing in lower-order brain regions.

vagus nerve: the tenth cranial nerve, a critical component of the parasympathetic nervous system, which innervates multiple organs and is involved in various physiological processes, such as digestion, heart rate regulation, and emotional responses via the bidirectional connection between the brain and body.

ventrolateral prefrontal cortex (VLPFC): a subregion of the prefrontal cortex situated laterally and ventrally in the frontal lobes. It is involved in various cognitive functions, including language processing, response inhibition, and emotion regulation.

ventromedial prefrontal cortex (VMPFC): a subregion of the prefrontal cortex located medially and ventrally in the frontal lobes. It is crucial in processing and integrating emotional information, social cognition, decision-making, and reward evaluation.


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