Accepted Manuscript

Title: Reciprocal effects between dominance and anger: A

systematic review

Author: João Carlos Centurion Cabral Patrice de Souza

Tavares Rosa Maria Martins de Almeida

PII: S0149-7634(16)30414-6
DOI: http://dx.doi.org/doi:10.1016/j.neubiorev.2016.10.021
Reference: NBR 2639

To appear in:

Received date: 5-7-2016

Revised date: 23-10-2016
Accepted date: 24-10-2016

Please cite this article as: Centurion Cabral, João Carlos, de Souza Tavares,
Patrice, de Almeida, Rosa Maria Martins, Reciprocal effects between dominance
and anger: A systematic review.Neuroscience and Biobehavioral Reviews
http://dx.doi.org/10.1016/j.neubiorev.2016.10.021

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 Reciprocal effects between dominance and anger: A systematic review

João Carlos Centurion Cabral a,*; Patrice de Souza Tavares b; and Rosa Maria
Martins de Almeida a

a Institute of Psychology, Laboratory of Experimental Psychology, Neuroscience and

Behavior, Federal University of Rio Grande do Sul (UFRGS), Porto Alegre, Brazil.
Electronic address: centurion.cabral@ufrgs.br
b Human and Information Sciences Institute, Clinical and Health Psychology
Laboratory, Federal University of Rio Grande (FURG), Rio Grande, Brazil. Electronic
address: patricetavares@hotmail.com
c Institute of Psychology, Laboratory of Experimental Psychology, Neuroscience and
Behavior, Federal University of Rio Grande do Sul (UFRGS), Porto Alegre, Brazil.
Electronic address: rosa_almeida@yahoo.com

*Corresponding author:
João Carlos Centurion Cabral
Institute of Psychology – Federal University of Rio Grande do Sul
Postal address: 2600 Ramiro Barcelos st, - Porto Alegre – Zip Code: 90035-003
Rio Grande do Sul – Brazil.– Telephone: +55 53 3201-5844 – Fax: +55 51 3308-
5470
Electronic address: centurion.cabral@ufrgs.br

1
Highlights

 There is a positive association between human dominance behavior and anger

 Social status and dominance have a direct effect on the perception of anger
 Anger expression has causal effects on the perception of dominance
 Dominance and anger have a mutual causal relationship possibly with positive feedback

ABSTRACT

Dominance and high status are directly associated with perception of angry

expressions. However, studies that have sought to empirically assess the causal

mechanisms between these construct are still relatively scarce. Moreover, several

variables can influence and be influenced by both anger and dominance, increasing

the complexity of synthesizing the findings related to the association between these

agonistic behaviors. We conducted a systematic review in five electronic databases. A

total of 207 potentially relevant publications were identified and screened. Of those, 20

articles were found eligible for detailed review, with 26 empirical studies. All reviewed

studies reported an association between dominance and anger. Social status and

dominance have a direct effect on the perception of anger. In turn, the perception of

anger has a consistent effect on attributions of dominance for those who express this

emotion. There are mutual effects between dominance and anger, which, if recurring

and positively feedback-regulated, at least in perceptual terms, can lead to the

establishment and maintenance of dominance hierarchies in social groups.

Keywords: Emotion. Anger. Dominance behavior. Social status. Submission.

2
 1. Introduction

Hostile interactions are critical for the survival of primates and other mammals (Buss,

2008; de Almeida et al., 2015; Honess and Marin, 2006). Dominance is an agonistic

behavior that has a direct impact on the organization of social groups and on

interpersonal relationships (Chiao, 2010; Johnson et al., 2012). This behavioral pattern

can be defined as a type of social relationship based on control of both the behavior of

hierarchically subordinate individuals and of valuable resources (Buss, 2008; Chiao et

al., 2009). Besides, dominance hierarchies determine the order of access to resources,

reducing energy expenditure and damage caused by intraspecific contests (Arregi et

al., 2006; Drews, 1993; Kaufmann, 1983). In humans, dominance drive and pursuit of

high social status are conceptually similar to power motivation, i.e., control of valuable

resources (Anderson and Galinsky, 2006; Ridgeway and Diekema, 1989; Salvador,

2005). However, the social hierarchy is subject to change every time that dominance is

challenged by a subordinate (Arregi et al., 2006; Campbell, 1999; Marsh et al., 2009).

Perception of threat to the status of a dominant individual induces the manifestation of

greater aggressiveness against the one who made the assault, either as a lower-

ranking individual or as an external intruder (Drews, 1993; Sewards and Sewards,

2003). Although the behavioral and physiological characteristics of power motivation

and dominance are frequently associated with the expression of some emotions in

humans, the relationship between these variables is not yet completely understood.

Dominance interaction does not only apply to physical aggression, since it also

includes the display of threats, as well as other strategies for conflict resolution

(Johnson et al., 2012; Ridgeway, 1987). Expressions of anger are often interpreted as

a threat signals (Coccaro et al., 2007; Hansen and Hansen, 1988; Hermans et al.,

2008; Hess et al., 2009; Öhman et al., 2001) and, in general terms, threat displays

exert a key role in establishing and maintaining the dominance hierarchy (Buss, 2008).
3
Anger is considered a basic emotion (Ekman, 1992; Izard, 2007) that can be broadly

defined as an emotional state that ranges from mild irritation to rage, often with a

feeling of hostility toward someone and an intent to cause damage (Berkowitz and

Harmon-Jones, 2004; Carver and Harmon-Jones, 2009; Frijda, 1986; Ramírez and

Andreu, 2006). In turn, anger expression is a behavioral response to angry feelings,

ranging from suppression of anger to its explicit expression toward someone or

something, primarily being facially characterized by frowning (Deffenbacher et al.,

1996; Hess et al., 2009; Öhman et al., 2001; Tipples et al., 2002). In healthy persons,

anger is triggered when a significant goal is frustrated by an external agent’s improper

action (Berkowitz and Harmon-Jones, 2004; Levine, 1996, 1995). Several cultural and

physiological characteristics directly influence the expression of this emotion.

Additionally, behavioral reactions of anger are commonly associated with hostility,

impulsivity and aggression (Archer and Webb, 2006; Buss and Perry, 1992; Hwang et

al., 2016; Sánchez-Martín et al., 2011). That is, angry individuals are more likely to

show aggressive behaviors, which, in turn, is also a decisive factor for dominance in

natural environment (Archer, 2006; Carré et al., 2013; Johnson et al., 2007; Mazur and

Booth, 1998).

Indeed, several anger-related characteristics are directly or indirectly associated

with dominance and power pursuit (Archer and Webb, 2006; Peterson and Harmon-

Jones, 2012; Sewards and Sewards, 2002; Shariff and Tracy, 2011). Regardless of

social status, people expressing anger are often perceived as more dominant than

when they express other emotions or are in neutral state (Archer and Webb, 2006;

Hareli et al., 2009; Knutson, 1996; Marsh et al., 2005; Tiedens, 2001; Tiedens et al.,

2000). In addition, neurobiological features typically seen in dominance drives are

equally common in situations of anger expression. Anger experience and dominance

motivation are both characterized by a reduction of top-down control and an increase

4
in bottom-up pattern systems, i.e., enhancing amygdala function and impairing areas

of prefrontal cortex (PFC) responsible for impulse control, such as orbitofrontal cortex

(OFC), ventromedial PFC (vmPFC), and anterior cingulate cortex (ACC) (Beaver et al.,

2008; Coccaro et al., 2007; de Almeida et al., 2015). Autonomically, steroid hormones

can affect the expression of anger (Carver and Harmon-Jones, 2009; Peterson and

Harmon-Jones, 2012; van Honk et al., 1999; Wirth and Schultheiss, 2007), as well as

the occurrence of dominant and competitive behaviors (Mazur and Booth, 1998; Mehta

et al., 2008). Even though the effects of dominance and anger are well known to affect

human social interaction individually, empirical studies on a possible causal

relationship between these variables are still scarce.

Aggressiveness and violence have important implications for living in social

groups, yet many of their natural and emotional determinants are not completely

known. Understanding the motivations that underlie behaviors with potential

deleterious consequences is crucial to avoiding significant social impairments. Several

variables can influence and be influenced by both anger and dominance. Thus, the

importance of reviewing the scientific literature, so that we can more deeply

understand these patterns of agonistic behaviors, becomes evident. With this in mind,

we conducted a systematic review of empirical studies on the relationship between

dominance and anger. Therefore, this review sought to answer four questions: I) Is

there a relationship between dominance and anger? II) Does dominance have a direct

effect on anger? III) Does anger have a direct effect on dominance? IV) What are the

environmental and biological factors that determine the relationship between these

variables? Given the multifactorial nature of the variables, this review focuses on data

from adult and healthy human participants. It was expected that, due to this control of

the variables, such data could enable a more concise and productive discussion on the

relationship between dominance and anger in humans.

5
 2. METHOD

The following electronic databases were searched for relevant studies in

November 2015: Scopus, Web of Science, PsycNET (APA), PubMed (Medline), and

Scielo.org (Scientific Electronic Library Online). Articles published between 1990 and

November 2014 were sampled using the following search strategy: ((anger

[Title/Abstract] OR angry [Title/Abstract]) AND ("dominance motives" [Title/Abstract]

OR "dominance motivation" [Title/Abstract] OR "Social dominance"[Title/Abstract] OR

"social hierarchy"[Title/Abstract] OR “dominance hierarchy”[Title/Abstract] OR

subordinance [Title/Abstract] OR “dominant behavior” [Title/Abstract] OR “high-status

people” [Title/Abstract]) NOT (disorder [Title/Abstract] OR pathology [Title/Abstract] OR

pathological [Title/Abstract] OR symptoms[Title/Abstract])) (see Figure 1 for details).

The 207 publications originally identified were submitted to a screening

procedure to assess their adequacy based on inclusion and exclusion criteria, as well

as assessing the methodological quality of studies. The screening procedure was

simultaneously and independently carried out by two reviewers previously trained for

the task. The following items were used as inclusion criteria: (1) type of publication –

peer reviewed articles based on empirical research and published in scientific journals;

(2) aim of the study – the main objectives, or at least the secondary objectives, should

be related to dominance and anger; (3) subjects – postpubertal healthy humans; (4)

period of publication - articles published from 1990 to 2015; and (5) language - articles

published in English, Spanish or Portuguese. Then, non-relevant publications for this

review were excluded by the following criteria: (1) clinical variables – studies focusing

on diseases and/or pathological behaviors and on the effects of health interventions

and clinical treatments; (2) psychoactive substances – studies on effects of

psychoactive substances; and (3) social discrimination – studies on prejudice and

social discrimination.
6
 Ultimately, 20 articles were reviewed and the concordance rate between

reviewers was 91.4%, calculated after exclusion of duplicated articles (Figure 1). In the

cases of discrepancies, the opinion of a third reviewer was requested. Reviewed

articles were organized by author, publication year, sample size, methods and design

of the study, variables, and main results (Table 1).

3. Results and Discussion

The search of five databases resulted in identifying 207 articles, of which 20

publications met the inclusion and exclusion criteria, as illustrated in Figure 1. These

20 publications, corresponding to 26 empirical studies, evaluated, directly or indirectly,

the relationship between human dominance behavior and anger. Most publications

reported only one study, however, seven publications reported two or three

experiments with different objectives. Any experiments that did not investigate the

relationship between dominance and anger were excluded from this review. In total,

these 26 studies assessed 5,032 healthy adults, from different countries and distinct

sociodemographic and professional profiles. The vast majority of studies were

conducted with mixed-gender samples, while only two studies were carried out with

exclusively male samples and one with exclusively female samples. Studies were

predominantly experimental (i.e., 23 studies), with only three observational studies,

highlighting the current need to understand the causal mechanisms of the relationship

between these variables. Furthermore, 22 of the reviewed articles were published in

the last 10 years. No study from before 2000 was identified by the search strategy and

selection criteria used.

3.1. Relationship between dominance and anger

7
All reviewed studies revealed an association between dominance and anger, and

almost all reported a positive relationship, at least for one measured variable. Thus, it

can be consistently said that there is an association between human dominance

behavior and anger. Nevertheless, contradictory findings were also seen in these

studies. Accordingly, some studies found no significant association of dominance with

anger for a specific gender, especially females, or for some specific ethnic group or

culture. An experiment with dominance self-report measures found no significant

relationship of dominance motivation with stimuli displays of different emotional facial

expressions, however, the authors have found such an association for salivary

testosterone measurement (van Honk et al., 2000), which is considered a biological

marker linked to dominant behavior (Mazur and Booth, 1998; Terburg et al., 2009; van

Honk et al., 2010). Besides, cross-cultural research has found a negative correlation

between angry expressions and self-reported measurements of subjective social

status (Park et al., 2013), and this was the only paper to report a negative relationship

between the variables in reviewed articles. On the other hand, Westhuizen and Solms

(2015) showed a positive association of anger with dominance, despite discussing the

low probability of there being a relationship between these variables. According to

them, dominance plays a central role in establishing social ascendancy, which,

ultimately, has an adaptive effect of stabilizing the social group and its internal

conflicts. For these authors, the involvement of anger in this process of social

hierarchization would be unlikely, and even counterproductive, during social

competition, given the deleterious potential of anger in these contexts.

Although there is a need for further clarification of this relationship and for

deepening theoretical understanding, the current review allows us to warily assume

that dominance and anger are associated, likely through a positive relationship. Since

8
both dimensions (dominance-submission and agonistic emotions, i.e., anger–fear)

produce a strong effect on human social interactions and on group organization, it is

worth noting that the association between dominance and anger can exert a major role

in the understanding of how to develop and maintain human social relations, in

particularly hostile interactions.

3.2. Does dominance have an effect on anger?

Nine studies performed some kind of experimental manipulation on dominance-related

conditions, and seven out of these nine focused specifically on social and/or

occupational status. Most of these studies manipulated the status of a stimulus to be

perceived (pictures, videos, or descriptive vignettes), i.e., the social status of a

character. Moreover, two correlational studies discussed the impact of dominance on

anger. Within the experimental research, almost 90% of studies (eight studies) focused

on perceptual and/or attentional aspects of dominance.

Individuals of high social status or dominance traits are perceived as being more

likely to express anger in negative contexts (Tiedens et al., 2000) and as having

greater intensity in their expressions of anger (Hess et al., 2004). Furthermore, the

perception of time for expressions of anger (stimulus dwell time) are also biased by the

dominance level of who is observed. Participants in the experiment conducted by

Ratcliff, Franklin, Nelson, and Vescio (2012) perceived angry expressions to appear

sooner and persist longer on faces of high-status people compared to low-status faces.

In other words, there is considerable attentional bias for facial expressions of anger,

resulting from the social status of the observed individual. According to the authors

(Ratcliff et al., 2012b), anger strongly signals threat in social contexts, and high-status,

and therefore more powerful, people are more likely and more free to act on their

intentions and goals than low-status people. In this sense, anger expression would be
9
physical evidence of intent to cause harm (i.e., a sign of a feeling of hostility) and, in

turn, high-status would be a clue to the increased capacity and likelihood to act, which

could lead to the occurrence of direct aggression or coercive behavior (de Almeida et

al., 2015; Tiedens et al., 2000).

However, this perceptual bias can also be explained by stereotypes related to

high social status positions. It is socially expected that high-status people feel more

anger in adverse situations (and pride in positive situations) than sadness or fear

(Ratcliff et al., 2012a; Tiedens, 2001; Tiedens et al., 2000). Thus, such social

expectations can be the result of attributional biases related to dominance behaviors.

Stereotypes may play a prominent role, especially by attributing increased ability and

competence in performing tasks to angry individuals, attributions that are also usually

conferred to people with high-status and power (Tiedens et al., 2000). Both

explanations, i.e. the propensity to violent and coercive behaviors, as well as the

attributions of increased ability and competence, for dominant individuals who express

anger can be a mechanism to help to stabilize the group hierarchy. It would occur to

prevent potential challenging behavior of low social status and/or submissive

individuals. Indeed, expectations related to the social roles and power can be relevant

to human perception of emotions, especially for anger.

The direct role of dominance motivation on anger should be further addressed.

Carr, Winkielman, and Oveis (2014) conducted an experiment that induced dominance

(sense of power) in participants, and then, they measured electromyographic activity of

the corrugator supercilii (for anger facial expression) and zygomaticus major (for

happiness) muscles, while exposing participants to emotional expression stimuli. As a

result, the participants responded to the expressions of anger with increased activity of

the corrugator muscle, as compared to expressions of happiness or baseline

10
measures. In this study, participants also showed higher corrugator activity in response

to angry than to happy expressions, when high-status was conferred to targets

(stimuli). Correspondingly, participants (perceivers) of low-power group smile for all

targets, regardless of emotional expression or status conferred to targets. Participants

assigned to high-power group did not mimic high status targets’ smiles, and curiously,

they showed incongruent smiling responses when there were high-status targets

expressing anger. This may be related to social expectation differences of people who

are at different power levels. Certainly, the implicit attributions made by a non-

dominant subject (perceiver) about a dominant one (target), will be considerably

different from an attribution made when both are dominant or subordinate. Therefore,

these results may be explained by a power asymmetry.

In this way, possible attributions for dominance behaviors (propensity to

aggression and attribution of increased ability) can be significantly reduced when there

is a balance in dominance levels between perceiver and perceived (Carr et al., 2014;

van der Ham et al., 2014; Wilkowski and Meier, 2010). Even the aforementioned

incongruous response to a high-status adversary expressing anger (i.e., smiling) may

be evidence of another emotion directly linked to stabilized dominance, pride (Shariff

and Tracy, 2011, 2009), though this incongruity can equally be interpreted as a sign of

submission. It is worth mentioning that, in conflictual social interactions, absence of

mimicry of another person’s behavior does not necessarily imply ignoring the

opponent’s emotional expressions. In fact, Hofman, Terburg, van Wielink and Schutter

(2013) found a linear relation between dominance motivation and attentional bias for

facial expression of anger. The group with high levels of dominance had greater

attentional bias for anger expressions, which may mean that the dominance motivation

causes an increased behavioral vigilance towards facial anger cues (Hofman et al.,

11
2013). Under these circumstances, we can assume that social status and dominance

are key features of social life that have a direct effect on the perception of anger.

However, more studies are required for understanding the role of dominance on other

aspects of its relation with anger.

3.3. Does anger have an effect on dominance?

Among the 26 reviewed studies, 22 (84.6%) carried out some manipulation or

experimental condition related to the expression of anger. Of these, 20 studies used

visual stimuli (e.g., pictures, videos, and virtual simulations), and almost all studies

assessed perceptual and/or attentional aspects linked with emotional expressions.

Only one correlational study discussed the possible effects of personal experience of

anger on dominance behavior. In turn, dominance was evaluated by very different

methods, having some inclination for evaluations through subjective self-report on

perception of dominance.

Anger perception has a consistent effect on attribution of dominance for those

who express this emotion (e.g., Brescoll and Uhlmann, 2008; Hareli et al., 2015;

Hortensius et al., 2014; Terburg et al., 2012a, 2012b, 2011; Tiedens et al., 2000; van

der Ham et al., 2014; van Honk et al., 2000). Subjects expressing anger are perceived

as more dominant or having higher status (Brescoll and Uhlmann, 2008; Flowe, 2012;

Hareli et al., 2015, 2009; Terburg et al., 2011; Tiedens et al., 2000; van der Ham et al.,

2014; Watkins et al., 2013). Conversely, some researches on dominance reported

inconsistent findings when comparing the effects of anger with emotionally neutral

expressions. The most contrasting results are often seen when comparing the effects

of anger with fear and sadness expressions (e.g., Hareli et al., 2015, 2009; Marsh et

al., 2005; van der Ham et al., 2014; Wilkowski and Meier, 2010). Likewise, some

researchers suggest that emotionally neutral facial expressions or lack of emotional

12
reaction to adverse situations can lead one to be seen as more competent and skilled

by their peers, as well as being attributed higher social status (Brescoll and Uhlmann,

2008), at least in a professional context (Hareli et al., 2009). In other words, facial

expressions without emotional changes can communicate significant dominance

signals, depending on the context of their occurrence.

On the other hand, supporting the role of anger on dominance, angry facial

expressions can lead to an increase in sustained eye-contact in subjects (perceivers)

exposed to such stimuli (Hofman et al., 2013; Hortensius et al., 2014; Terburg et al.,

2011), at least for people with a high level of dominance motivation (Hofman et al.,

2013; Terburg et al., 2012a, 2011). Accordingly, dominant subjects show decreased

gaze aversion from angry expressions. This pattern has even been seen for masked

stimuli (which were presented faster than necessary to be consciously perceived) (e.g.,

Hortensius et al., 2014; Terburg et al., 2012a, 2011; van Honk et al., 2000). Enduring

gaze is considered a behavioral marker of dominance motivation, and may even be an

automatic and non-conscious adaptive mechanism (Terburg et al., 2012a; van Honk et

al., 2000). Just as in humans, other mammals with complex social behaviors establish

and maintain social hierarchy through direct and sustained eye contact (Chiao, 2010;

Drews, 1993; Mazur and Booth, 1998). In primates, the dominance hierarchy is

primarily established when the animals stare directly and fixedly at each other,

generally also involving other threat signals, until one animal looks away, signaling

submission, thereby avoiding possible injuries resulting from direct physical conflict (de

Almeida et al., 2015; Mazur and Booth, 1998).

Some lines of research have sustained the notion that the angry facial

expression can serve as a sign of threat in the context of conflict for hierarchy (Mazur

and Booth, 1998; Mellentin et al., 2015; Öhman et al., 2001; Terburg et al., 2012a,

13
2011; Tipples et al., 2002). Once again, this indirect association between anger and

dominance could have their causal mechanisms related to attribution bias for those

expressing anger, both in terms of greater physical strength/hostility and greater

personal skills. According to Sell, Comides, and Tooby (2014), anger expression can

be an adaptation for enhancing cues of strength to possible opponents, which, in turn,

would increase the bargaining power of the anger expresser. During a conflict, such

cues would make the opponent perceive better conditions to fight in one who express

anger. A similar hypothesis was formulated and tested by Tiedens and colleagues

(2000). They affirmed that mutual perceptual effects between anger and dominance

might be explained by the assumptions (i.e., attributions) made by an observer, that is,

angry people and people who have high social position are seen as having higher

skills and abilities. Thus, it is clear that anger has causal effects on dominance

behavior, but this statement can only be sustained for perceptual and attentional

levels. There are no studies that have tested if feelings of anger, or their physiological

responses, can impact personal motivations for dominance.

3.4. What environmental and neurobiological factors are determinants of

these patterns and their functions?

Cultural, personal and physiological differences can be decisive for the occurrence of

this reciprocal effect between dominance and anger. Understanding the cultural

aspects underlying this relationship is crucial to have a complete view of the

phenomenon. Not only are there significant differences in emotional expression

depending on the cultural norms of a country (McLinton and Dollard, 2014), but these

can also alter the causal pattern of dominance and anger (Park et al., 2013). A large

correlational study found, by comparing the relationship of anger with social status

among Japanese and Australian workers, that anger against superiors is significantly

14
attenuated in the Japanese population (McLinton and Dollard, 2014). This pattern was

not seen among Australians. These authors advocated the notion that the cultural

dimension has an undeniable relevance in the experience of anger. However, the

highest levels of anger, in both Japanese and Australian populations, were against

coworkers and subordinates. This cross-cultural pattern corroborates the experimental

findings reviewed here, although it does not ignore that there are cultural differences

for expression of anger and dominance. Another cultural aspect that may have direct

implications on the concept of the functionality of anger is that individualistic cultures

(exemplified by countries such as the United States) tend to understand anger as a

way to relieve frustration (Park et al., 2013). On the other hand, collectivistic cultures

(often attributed to Asian countries) tend to understand this emotion as a way to

demonstrate authority (Park et al., 2013). Further, there are certain cultural norms in

some Asian countries against angry expressions, especially for low status people

(Park et al., 2013), which can directly affect the interaction between dominance and

anger.

Gender and physical features can also have moderating effects on the

relationship between dominance and anger. Women are perceived as less dominant

than men when both express anger (Hareli et al., 2009; Hess et al., 2004; Watkins et

al., 2013). Male expression of anger can lead to an increase in perceived status for

those who express it. However, for women expressing anger, the effect may be the

opposite, being conferred lower status and being perceived as less competent

(Brescoll and Uhlmann, 2008). Brescoll and Uhlman (2008) have argued that this is

due to an attribution bias: angry females were seen as internally motivated, i.e., the

explanation for this emotion was based on personal characteristics and personality.

For angry males, attribution was usually external and contextualized. Nonetheless,

15
when the anger expressed by women had an external explanation experimentally

attributed, there was no impairment in women’s perceived status and competence.

These perceptual and stereotypical differences may originate in the physical

characteristics and appearances related to gender. People with more masculinized

faces are perceived as more dominant (Hess et al., 2004; Watkins et al., 2013).

Furthermore, physiological characteristics linked to sex also play a key role in

the manifestation of agonistic behavior. Androgenic hormone levels (e.g., testosterone)

are consistently associated with the expression of anger and dominance motivation

(Archer, 2006; de Almeida et al., 2015; Mazur and Booth, 1998; Peterson and Harmon-

Jones, 2012; Sewards and Sewards, 2003; van Honk et al., 1999; Wirth and

Schultheiss, 2007), although these results are often not successfully replicated in

women (Cashdan, 2003; Schultheiss et al., 2004; Stanton et al., 2009; Stanton and

Schultheiss, 2009). In both female and male samples, physiological levels of

testosterone are positively associated with attentional bias to angry facial expressions

(Terburg et al., 2012a; van Honk et al., 1999), and exogenous administration of

testosterone can enhance cardiac responses to such facial expressions (van Honk et

al., 2001). Testosterone is a steroid hormone essential for promoting and maintaining

dominant behavior in several vertebrate species (Archer, 2006; Mazur and Booth,

1998; Montoya et al., 2012). In contrast, low levels of cortisol are correlated with

agonistic behavior patterns (Terburg et al., 2009; van Honk et al., 2010). Cortisol is the

main glucocorticoid hormone in humans, and its sustained high levels can suppress

gonadal function and the synthesis of androgenic steroids (Charmandari et al., 2005;

McEwen, 2007). Thus, the combination of low levels of cortisol with high levels of

testosterone has been considered a physiological marker of aggressive behavior and

dominance (van Honk et al., 2010). This pattern also matches the hormonal pattern

16
associated with dominant behavior, known as winner effect, i.e., an increase in

aggressiveness and readiness for new confrontations in individuals that win a

competition (Holekamp and Strauss, 2016). Another steroid hormone, estradiol, which

is considered the main female sex hormone, similarly has a significant impact on the

manifestation of dominance, emotional lability, and aggressive behavior in humans

(Gasbarri et al., 2012; Montgomery et al., 1987; Stanton and Edelstein, 2009; Stanton

and Schultheiss, 2007). Stanton and Schultheiss (2009) argue that estradiol, and not

testosterone, performs a central role in female dominance motivation. In women,

dominance is negatively associated with estradiol and estradiol/testosterone ratio

(Ziomkiewicz et al., 2015), and higher levels of estradiol produce less competitive

interaction (Cashdan, 2003). The effect of aromatase on the conversion of

testosterone to estradiol (Cover et al., 2014) should also be highlighted, since this may

have a relevant function for expression of agonistic behaviors in men too (Unger et al.,

2015). However, there is a need for further studies to investigate a possible sexual

dimorphism for expression of anger and dominance in humans, assessing the effects

of sex hormones and their relations with neurobiological findings regarding agonistic

behaviors.

Indeed, such endocrine profiles are highly correlated with the activation of

subcortical and cortical structures, such as the amygdala (de Almeida et al., 2015;

Hermans et al., 2008; Hofman et al., 2013), a structure responsible for emotional

reactions, including anger (Davis and Whalen, 2001), and the PFC (van Honk et al.,

2010), responsible for impulse control, planning and decision-making, among other

executive functions (Anderson et al., 2006; Miller and Cohen, 2001), respectively.

Clinical and experimental data indicate that the amygdala performs a prominent

function in both recognition and experience of anger, being its medial nucleus primarily

17
relevant for this emotion (Derntl et al., 2009; Öhman et al., 2001; Phillips et al., 2003;

Potegal et al., 2010; Whalen et al., 2001). Similarly, the activation of the amygdala is

also involved in the perception of dominant subjects in a disputed status context

(Chiao, 2010; Zink et al., 2008). Moreover, the PFC activation pattern for high status

cues is similar to the pattern seen for facial expression of anger, mainly in the

ventrolateral PFC (Marsh et al., 2009). Medial, dorsolateral and vmPFC, as well as

anterior and posterior cingulate cortex, are implicated in the perception of dominant

and superior status in a hierarchy (Chiao, 2010; Chiao et al., 2009; Marsh et al., 2009;

Zink et al., 2008). Such PFC regions are also involved with the perception of anger

(Beaver et al., 2008; Bediou et al., 2009; Coccaro et al., 2007; Pawliczek et al., 2013).

Furthermore, OFC is responsible for emotional impulse control, including anger

impulses (Lotze et al., 2007). Going beyond a perceptual level, the experiences of

anger and dominance provoke an increased OFC activity (Bechara et al., 2000; van

Honk et al., 2014). This activation pattern can be explained by social norms, since the

explicit expression of agonistic behaviors is socially undesirable (Hofman et al., 2013).

For instance, there is a decrease in OFC activity when there are unfair offers in the

ultimatum game, which is an economic game associated with both anger expression

and dominance (Mehta and Beer, 2010; Sanfey, 2003).

These patterns are also common in aggressive behavior in human and nonhuman

primates, and its manifestation is characterized by an impaired top-down control of

PFC (de Almeida et al., 2015). Such impaired inhibitory function of PFC and enhanced

amygdala activity is related with high levels of testosterone and low levels of cortisol.

Additionally, the amygdala and PFC are reciprocally connected (Price, 2003;

Stefanacci and Amaral, 2002), and this connectivity can also be influenced by sex

steroids (van Wingen et al., 2011). Testosterone increases the neural activity in the

18
amygdala (Derntl et al., 2009; Manuck et al., 2010; van Wingen et al., 2009) and

hinders its connectivity with PFC (Bos et al., 2012; van Wingen et al., 2010). On the

other hand, cortisol has a negative correlation with amygdala activation (Henckens et

al., 2010) and estradiol seems to enhance the connectivity between the PFC and

amygdala (Peper et al., 2011; Zeidan et al., 2011). Thus, ovarian hormones, such as

estradiol and progesterone, in low levels and in intermediate levels, respectively, can

impair the functioning of these brain regions (Cover et al., 2014). Additionally, steroid

hormones may also impair communication within serotonergic pathways (Martel et al.,

2009).

Indeed, clinical, experimental and observational studies have consistently

implicated low serotonin (5-HT) levels in agonistic behavior patterns and violence,

regardless of gender (Higley et al., 1996; Moore et al., 2002; Oades et al., 2008; Soloff

et al., 2003), since 5-HT can inhibit aggression in many species (Holekamp and

Strauss, 2016). Nevertheless, serotonergic systems can also be positively associated

with such behavioral patterns (Olivier and van Oorschot, 2005; van der Vegt et al.,

2003), including anger (Coccaro et al., 2015; Coccaro and Lee, 2010; Duke et al.,

2013; Suarez and Krishnan, 2006) and dominance (Krakowski, 2003; Larson and

Summers, 2001; Raleigh et al., 1991). The 5-HT1A receptors are distributed in several

brain regions, such as PFC and amygdala (Olivier and van Oorschot, 2005). This

receptor is abundantly expressed in the cingulate cortex and PFC, including vmPFC,

OFC and ACC (Albert et al., 2014; Paus, 2001; Witte et al., 2009), and its activation

inhibits the activity of target neurons (Albert et al., 2014). Thus, both low 5-HT levels

and postsynaptic 5-HT1A receptor agonist can enhance aggressive and impulsive

reactions (de Almeida et al., 2015). Furthermore, several studies have shown an

interaction between 5-HT and steroid hormones (Bonson et al., 1994; Miller et al.,

19
2012; Pinna et al., 2008; Witte et al., 2009). Testosterone, estrogen and cortisol can

modulate 5-HT function, altering density and distribution of 5-HT receptors and 5-HT

levels in the brain, as well as reducing tryptophan (Cowen, 2002; Kuepper et al.,

2010). It has been found that variations in the levels of sex hormones during menstrual

cycles can then change the distribution of 5-HT1A receptors (Witte et al., 2009).

Controversies about the modulating effects of 5-HT on agonistic behaviors certainly

need more studies to be undertaken in order to achieve further clarity, bearing in mind

the possible effects of upregulation and downregulation of 5-HT receptors, and their

interactions with steroid hormones. Moreover, these future studies may have to take

into account the differences between dominance behavior and dominance drive, as

well as the differences between anger expression and feeling of anger. Naturally these

explicit versions (behaviors) may have higher levels of prefrontal activity, given the

increased impulse control demand.

Finally, the brain activity linked to dominance and anger also depends on its types

and categorizations, since reactive (or explosive) aggression follows a similar pattern

seen for anger experience, unlike the pattern seen for proactive (or instrumental)

aggression. While reactive aggression is characterized by impaired top-down control

(higher amygdala activity and lower impulse control areas activity), proactive

aggression is characterized by elevated control exerted through PFC regions, given

that there is a great demand of action planning for these behaviors (de Almeida et al.,

2015). In turn, the neural effects of different possible types of dominance, e.g.,

charismatic dominance and agonistic dominance (Borkowska and Pawlowski, 2011;

Noë et al., 1980), and their associations with anger, need to be empirically assessed in

humans. Furthermore, studies aiming to directly determine the neurobiological basis of

the interaction between dominance and anger are still scarce.

20
 4. Conclusions

In sum, dominance and anger have mutual effects, possibly with a positive feedback

loop. Individually, both behavioral patterns, those related to the dominance–

submission dimension and those related to angry emotional reactions, have a

significant effect on human social interaction. However, understanding the mutual

effect between these variables will allow the development of a broader framework

about how to define social relations of hostility and power in human groups. Reciprocal

causal relationship between anger and dominance, being recurrent and positively fed

back, at least in a perceptual domain, can lead to a stabilization of the hierarchical

structure in social groups. That is, the expression of anger is perceived as more

dominant, and dominant individuals are perceived as more angry in adverse contexts.

This feedback could inhibit the challenge behavior (of subordinate individuals) to the

dominant, and, thereby, increase stability and maintenance of the hierarchy.

Reciprocal perceptual effects of anger and dominance may mean that this relationship

has a phylogenetic basis, which serves as a threat signal, through increased

aggressiveness and perceived strength, ensuring dominance hierarchy maintenance

and privileged access to resources. Although, given the absence of studies that have

empirically assessed the direct relationship of these feelings (i.e., to investigate

whether the personal feeling of anger induces dominance motivation and vice-versa),

we cannot state with certainty that the relation between dominance and anger occurs

only at a perceptual level, since both variables are predispositions to act in a certain

way. Functionally, the main action of these variables is not in perception, but, rather, in

the subjective feelings, personality traits, and ultimately, behavioral manifestations,

both being patterns of agonistic behavior. Therefore, further studies are needed to test

such hypotheses, as well as investigating the underlying neurobiological and genetic

21
mechanisms and several environmental factors that can influence or be influenced by

anger and dominance.

Acknowledgments

This work was supported by grants from Conselho Nacional de Desenvolvimento

Científico e Tecnológico (CNPq).

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Figure caption:

Figure 1. Flow chart of the articles selection process.

37
Table 1. The key features and main results of the selected studies on association between dominance and anger.

First author, n
Design Independent variables Dependent variables Main results [association dominance-anger]
year Population (♀/♂)
Hareli, 2015 94 - MBA students Experimental EE transitions (pictures); Perception of competence, EA in the end ↑ PD; change in emotion has an effect on
(Exp 1) (♀/♂) sociability, and intensity of PD; sign of weakness (sadness) has a greater impact on
emotions; PD; PD than a sign of strength (anger); [Positive]
Hareli, 2015 220 - Participants Experimental EE transitions (videos); Perceived competence, EA at the beginning or end (of an emotional reaction) ↑
(Exp 2) recruited from sociability, and intensity of PD; change in emotion has an effect on PD; end emotion
websites (♀/♂) emotions; PD; affects PD; [Positive]
Westhuizen, 36 - Academic Correlational Salivary C and T; Affect (ANPS); dominance scale; There was no correlation between T and dominance; T
2015 students (♂) correlated - with anger; C was not related to these
measures; T/C ratio + correlated with dominance; [Positive]
Carr, 2014 55 - Academic Experimental Induction of dominance; EMG (corrugator and Perception of EA ↑ corrugator activity; there was interaction
students (♀/♂) Stimuli: EE in combination zygomatic muscles); between emotional valence and power (status) of stimuli;
with high or low social PANAS; LP subjects smiled to all targets and expressions; HP
status (videos);
subjects did not mimic HP targets’ smiles; [Positive]
Ham, 2014 36 - Dutchmen Experimental Role conditions (high and Perception of believability Social context impacts perception of EE; after
(♀/♂) (online) low status for participant and emotional intensity; manipulation, stimuli with EA was perceived as more
and for stimulus); Stimuli: PD; dominant, regardless of stimuli status; [Positive]
EA or sadness;
McLinton, 601 - Japanese Correlational MIDAI scales (superiors, subordinates, co-workers, JAP: ↓ levels of anger towards superiors (and strangers);
2014 (JAP) and family, and strangers); AUS: influence of social status was non-significant; there
Australian (AUS) was no difference between AUS and JAP in anger towards
workers (♀/♂)
co-workers and subordinates; in both cultures, the ↑ levels
of anger were towards co-workers and subordinates;
culture was relevant to EA; [JAP: Positive; AUS: n.s.]
Hortensius, 32 - Healthy Experimental Facial and bodily EE Attentional bias (gaze- There was a slower gaze-aversion from non-consciously
2014 (Exp 1) young people (pictures) aversion); Dominance processed EA compared to happy, being + related to
(♀/♂) (BAS); dominance; there was an effect on dominance provoked by
masked angry body postures; [Positive]
Hofman, 2013 26 - Healthy Experimental Facial EE (pictures); high Dominance (BAS); EEG; High dominance group had ↑ attentional bias for EA;
academic and low dominance; Attentional bias; dominance was related to anger bias; dominance was
students (♀/♂) reflected in the asymmetry in cortical and subcortical
processing; [Positive]

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Park, 2013 2081 - Japanese Correlational Objective social status (occupation and educational Social status were associated with anger, showing that EA
(JAP) and achievements); subjective social status (self- ↓ with subjective status, but ↑ with objective status; there
American (USA) assessment); AE (AEI); personality traits (Big Five); was a moderator effect of culture: USA: ↓ subjective status
community
had ↑ EA; JAP: ↑ objective status had ↑ EA; [Positive;
sample (♀/♂)
Negative]
Watkins, 2013 200 - Participants Experimental Facial EE; stimulus sex PD; PD in ♂ was modulated by stimulus sex, regardless of EE;
recruited via (online) (picture) and level of facial ♀ perceived masculinized faces as more dominant (than
Internet (♀/♂) sexual dimorphism feminized), after EA of male stimuli, but not after female EA
(masculinized-feminized);
or after expression of happiness of both sexes; PD follows
a sex-specific pattern; [Positive (for ♀)]
Flowe, 2012 512 - Academic Experimental Facial EE; stimulus sex Perception of criminal PD was + associated with criminality in ♀ and ♂;
students (♀/♂) (picture) appearance, trustworthiness and dominance underlie the perception of
trustworthiness, criminal appearance; anger was + correlated with
dominance, threatening, dominance in ♀ and ♂; EA was perceived as more criminal
maturity and anger; appearance; [Positive]
Ratcliff, 2012a 48 - Academic Experimental Facial EE; high or low EE identification; SDO; High SDO subjects were better at identifying the emotions
students (♀/♂) social status; of high-status people; EA on high-status faces was more
discernible for perceivers high in SDO; [Positive]
Ratcliff, 2012b 69 - Academic Experimental Facial EE (video); high or EE detection; reaction Anger was perceived to persist longer on high-status faces
(Exp 1) students (♀/♂) low social status; time; compared to low-status faces; [Positive]
Ratcliff, 2012b 39 - Academic Experimental Facial EE (video); high or EE detection; reaction Anger was perceived to appear sooner on high-status
(Exp 2) students (♀/♂) low social status; time; faces compared to low-status faces; [Positive]
Terburg, 20 - Healthy Experimental 0.5 mg of testosterone; Mood (PMD); attentional After T administration, gaze-aversion was slower for EA
2012a volunteers (♀) facial EE; bias (gaze-aversion); than for happiness (unconsciously); anger-specific effect
confirms that T promotes dominance-related gaze
behavior; [Positive]
Terburg, 40 - Healthy Experimental Facial EE; Memory; anger (STAS); Submissive subjects averted their gaze more rapidly from
2012b volunteers (♀/♂) anxiety (STAI); attentional EA and anxious subjects initially gazed more often to, but
bias (gaze-aversion); subsequently avoided looking directly at angry faces; trait
anxiety predicted improved memory for the location of
angry faces; [Positive]
Terburg, 2011 40 - Healthy Experimental Facial EE; Attentional bias (gaze- Slower gaze aversion from masked EA is predicted by the
volunteers (♀/♂) aversion); Dominance dominance scores; speed of gaze aversion from masked
(BAS); EA depends on motives of dominance; [Positive]
Hareli, 2009 210 - Canadians Experimental Facial EE (picture); PD; There were main effects for EA and for stimulus sex, and
(Exp 1) (♀/♂) stimulus sex; an interaction between them; ♂ EA and happiness had ↑
PD, and ↓ PD when expressed sadness or fear, when
compared with neutral expression; ♀ EA and happiness

39
 had ↑ PD and ↓ PD when expressed sadness, when
compared with neutral or fear expression; [Positive]
Hareli, 2009 96 - Managers Experimental EE (videos); Perception of emotional EA was perceived as more dominant than shame; EA was
(Exp 2) (♀/♂) intensity; PD; not significantly different from neutral expression; EA was
perceived as less submissive; [Positive]
Brescoll, 2008 69 - Adults with Experimental EE (videos); stimulus sex; Status conferral; Subjects conferred higher status on the ♂ EA stimulus,
(Exp 1) work experience perception of competence; when compared with ♂ sadness or ♀ EA; ♀ sadness
(♀/♂) assessment of internal or expression had higher status conferred than ♀ EA; ♀ EA
external attributions; was attributed to internal characteristics and ♂ EA reaction
was attributed to external circumstances; [Positive]
Brescoll, 2008 180 - Adults with Experimental EE (videos); stimulus sex; Status conferral; There was an interaction between EA and stimuli gender;
(Exp 2) work experience occupational status of perception of competence; ♀ EA conferred lower status than ♀ expression of sadness;
(♀/♂) stimulus; assessment of internal or ♀ EA was seen as more out of control than ♂ EA and than
external attributions; ♂ or ♀ neutral expressions; [Positive]
Brescoll, 2008 133 - Adults with Experimental EE (videos); stimulus sex; Status conferral; There was an interaction between EA and stimuli gender;
(Exp 3) work experience perception of competence; ♂ EA without external attribution received higher status
(♀/♂) than the unemotional ♂ or ♂ EA with external attribution;
♀ EA with external attribution received higher status than ♀
EA without external attribution; [Positive]
Hess, 2004 44 - Academic Experimental EE (videos); physical Perception of emotional When affiliation and dominance levels were controlled for,
(Exp 1) students (♀/♂) intensity; stimuli sex; intensity; ♀ were perceived as expressing more intense anger, and
♂ were perceived as happier; [Positive]
van Honk, 40 - Academic Experimental Facial EE (picture); Dominance (DPI); T; C; There was no effect between self-reported dominance and
2000 students (♂) supraliminal and subliminal attentional bias; attentional bias for both the unmasked and the masked
stimulus; task; C ↑ in subjects with selective attention towards
unmasked and masked EA; T ↑ in subjects with selective
attention towards unmasked; [n.s.: self-reported
dominance; Positive: T]
Tiedens, 2000 40 - Academic Experimental Vignettes of different Emotion perception; High status character was viewed as angry or proud and
(Exp 1) students (♀/♂) social status characters; attribution of agency; low status character was seen as sad or guilty; [Positive]
Tiedens, 2000 71 - Volunteers Experimental Vignettes of character with Social status; Angry or proud character was viewed as high status and
(Exp 2) (♀/♂) different emotional states; sad or guilty character was seen as low status; [Positive]
Legend: ♀ = Women; ♂ = Men; EA = Expression of anger; PD = Perception of dominance; ANPS = Affective neuroscience personality scales; EMG = Electromyography;
PANAS = Positive and Negative Affect Schedule; LP = Low-power; HP = High-power; + = Positive/positively; - = Negative/negatively; ↑ = Increase; ↓ = Decrease; EE =
Emotional expression; MIDAI = McLinton interpersonal domain-specific anger instrument; BAS = Behavioral activation scale; EEG = Electroencephalography; AEI = Anger
expression Inventory; IV = Independent variable; BIG-5 = five factor personality traits model; SDO = Social dominance orientation scale; PMD = Profile of mood state; T =
Testosterone; C = Cortisol; STAS = State-trait anger scale; STAI = State-trait anxiety inventory; DPI = Dutch personality inventory; n.s. = non-significant.

40
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