research paper on social hierarchy

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17 Competition, Dominance, and Social Hierarchy

University of Oregon, Eugene, OR, USA

University College London, London, England, UK

• Published: 09 May 2019
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This chapter reviews literature having to do with the social-behavioral neuroendocrinology of competition, dominance, and status hierarchies in humans. After defining these terms, their importance, and everyday relevance, the chapter discusses the major research findings that suggest a bidirectional influence between these social behaviors and the steroid hormones cortisol and testosterone. Specifically, the association between cortisol and social rank and cortisol’s reactivity to social challenges are discussed. Further, this chapter discusses research that tests the predictions that basal testosterone is related to status-motivated behavior, that testosterone levels are transiently altered during contests for status, and that these changes function to promote subsequent status-seeking behavior. Noting the nuance of these findings, the personality and context factors that appear to moderate testosterone–status relationships are highlighted. Finally, this chapter includes both a new theoretical model for the testosterone–social status relationship that captures this complexity and, in closing, summarizes promising areas of future research.

In ancient Egyptian society, formalized social hierarchy was manifested in the ritualistic burial of the dead. Social status was symbolized in almost every aspect of the mortuary arrangement—how close in proximity one’s tomb was to the king, height from the ground, size of the tomb, level of artistic decoration, and material accoutrements (Romano, 1990 ; Taylor, 2001 ). Male members of the elite class would begin constructing a tomb at the peak of their career, even setting aside an endowment for upkeep and offerings after death (Baines & Lacovara, 2002 ). These preemptive structures served as status symbols for the living as the “tomb was a central vehicle for peer competition (Baines & Lacovara, 2002 , p. 10). For Egyptians, death was as important as life; it was imperative that social status attained in life be maintained in death.

Social hierarchy remains a pervasive and fundamental framework of modern human society and social relationships across cultural boundaries (Diefenbach, 2013 ; Sidanius & Pratto, 1999 ). Formalized and highly stratified hierarchies exist within political and government structures, such as traditional monarchies, dictatorships, capitalist systems, and resulting socioeconomic status (SES). There are also less large-scale yet pervasive social hierarchical systems: class or grade in school, workplace title or position, finish place in a competition, veteran or rookie status on a professional sports team, the lead versus backup role in a play or dance assemble, and even the naming system for increasingly luxurious seats on an airplane (first class, business class, and main cabin). These systematic rankings within everyday life are representative of the fundamental human need to formally classify social position based on level of distinction, wealth, talent or ability, and assumed power and privilege within each role. Some evolutionary psychologists even suggest that it was the emergence of increasingly complex social dominance interactions that provided the selection pressure for the more sophisticated human intelligence and capacity for language (Alexander, 1989 ; Flinn, Geary, & Ward, 2005 ).

Importantly, society-level social hierarchies result in stepwise differences among the upper and increasingly lower echelons (Dasgupta, 2015 ; Sidanius & Pratto, 1999 ) in access to resources directly linked to survival, such as food, water, housing, land, and health care (e.g., “food deserts” common in low-income neighborhoods, lack of access to potable water in certain developing nations; Montgomery & Elimelech, 2007 ; Walker, Keane, & Burke, 2010 ). In review of the adverse physiological effects of stress resulting from “dominance hierarchies” in primates, Sapolsky ( 2005 , p. 652) states that despite the complexities of human social structure, “the SES gradient of health among Westernized humans is a robust example of social inequalities predicting patterns of disease.” Indeed, there appears to be a strong positive relationship in primates, including humans, between both objective and subjective social status ranking and psychological stress, health, and, ultimately, the quality of life and survivability of individuals (Demakakos, Nazroo, Breeze, & Marmot, 2008 ; Sapolsky, 2004 ). Thus, aside from the basic science motivation to discover patterns of human behavior and related causes, there is also a perhaps more important ethical responsibility to understand the psychological and interacting biological processes involved in social dominance behavior and resulting social hierarchies.

Not all dominance-motivated social ordering is easily recognized. There are subtler hierarchies that individuals form naturally among social groups in various contexts (e.g., spontaneously emerging leader–follower relationships and popularity among peers). Everyday acts of dominance and deference (e.g., eye contact, the firmness of a handshake, the giving and taking of verbal directives) are used to attain and maintain social standing (Tiedens & Fragale, 2003 ). People also go to great lengths to advertise status ranking with luxury goods like clothes, shoes, jewelry, cars, and houses (a term known as conspicuous consumption; O’Cass & McEwen, 2004 ); we even have objects that serve no other purpose than to advertise status (e.g., trophies, diplomas, and award ribbons). Even more subtle, social media platforms provide individuals far-reaching opportunity (70 percent of Americans polled used some form of social media; Pew Research Center, 2017) to attempt to outrank their peers in quality of life, personal achievements, travel, number of friends, and perceived happiness (Chua & Chang, 2016 ). Though seemingly nonconsequential, these small and sometimes subtle everyday gestures may serve to reinforce larger scale hierarchical social systems and represent the pervasiveness of the fundamental human striving for status.

Like other basic psychological drives, such as thirst, hunger, and sexual desire, the motivation for status has biological underpinnings in the form of a cascade of bidirectional brain–body interactions communicated via chemical messengers, specifically steroid hormones. Testosterone was first recognized for its long-term impact on the development of male secondary sexual characteristics and sexual behavior (August, Grumbach, & Kaplan, 1972 ; Phoenix, Goy, Gerall, & Young, 1959 ; Phoenix, Slob, & Goy, 1973 ). Given the more physically dominant nature of male-typical behavior in male primates, including humans, early research on social hierarchies proposed that testosterone would increase with status rank and social dominance (Ehrenkranz, Bliss, & Sheard, 1974 ; Purifoy & Koopmans, 1979 ; Rose, Bernstein, & Gordon, 1975 ; Sapolsky, 1982 ). Cortisol, well known for its link to physical and psychological stress, was also linked in earlier research to rank within the social hierarchy (Sapolsky, 1982 ). Subsequent research in the decades that followed has revealed a complex relationship between testosterone, cortisol, and dominance rank and related behaviors (Casto & Edwards, 2016a ; Hamilton, Carré, Mehta, Olmstead, & Whitaker, 2015 ; Mazur & Booth, 1998 ).

This chapter will review literature having to do with the social-behavioral neuroendocrinology of competition, dominance, and status hierarchies, first defining these terms and then discussing the major research findings that have emerged in this field. There is a vast research literature on these concepts pertaining to nonhuman animal behavior (e.g., Gleason, Fuxjager, Oyegbile, & Marler, 2009 ); however, this chapter focuses primarily on the research involving human participants with implications for human behavior. The literature discussed here lies at the crossroads of social-personality psychology and behavioral endocrinology (two otherwise quite distinct fields), wherein the complexity of human socially, culturally, and contextually embedded behavior is predicted by elegant, yet primitive, and evolutionarily adaptive hormonal fluctuations.

Defining Social Hierarchy, Dominance, and Competition

That valuable resources (e.g., food, water, land, sexual partners) are limited stratifies social order: Some (dominants/superiors) have more access to these resources than others (subordinates). Those who command these resources do so because they possess qualities that are attractive to or more dominant than others, qualities that confer greater attention, influence, or power over those that are lower ranking (Anderson, Hildreth, & Howland, 2015 ; Berger, Cohen, & Zelditch, 1972 ; Berger, Rosenholtz, & Zelditch, 1980 ; Kemper, 1990 ). The compilation of these relationships within social group results in social hierarchy , the rank ordering “of individuals or groups on a valued social dimension” (Magee & Galinsky, 2008 , p. 354; as cited by Cheng, Tracy, Foulsham, Kingstone, & Henrich, 2013 ). A higher rank order indicates that an individual or group has higher social status (Berger et al., 1972 , 1980 ; Ellyson & Dovidio, 1985 ). An individual’s actual rank is not necessarily fixed—some hierarchies can be rather unstable—and is often context dependent (e.g., at work vs. among peers or family). Attaining a higher position within a social hierarchy can predict increases in subsequent status-seeking behavior in an effort to maintain this high status and attain even higher positions in the future (e.g., Zilioli & Watson, 2014 ). Because of its notable survival advantages, direct reward value, universality, and long-term impact on psychological well-being, social status is considered a fundamental human motive (Anderson et al., 2015 ). Indeed, social hierarchies are a universal and evolutionarily conserved characteristic of most human societies (Chiao, 2010 ; Diefenbach, 2013 ; Gledhill, Bender, & Larsen, 1988 ). Additionally, neuroscientists have revealed distinct neural networks in the brain for the perception and maintenance of social hierarchy (Chiao, 2010 ).

Status is often decided through competition , “a social interaction in which access to something valued is contested between individuals and groups” (Casto & Edwards, 2016a , p. 21). The “something valued” could be the resource that is in limited supply or the simple feeling that one possesses rank or power over others. Prevailing over one’s opponent signals dominance , whereas failing to prevail or conceding defeat signals deference (Mazur, 1985 ). In an everyday sense, the motivation for engaging in competition may be the simple joy of winning—undoubtedly resulting from the inherent reward of status gained by the demonstration of dominance. However, the precise definition of dominance and related terms appears to depend on the specific literature within which it is discussed—social psychology, personality psychology, sociobiology, or evolutionary psychology (Cheng et al., 2013 ). One can possess dominance (i.e., have physical or psychological qualities that others defer to or admire, such as strength or competence), be considered dominant (i.e., rank higher), behave dominantly (e.g., verbal and nonverbal signaling of dominance, acting as a leader, refusal to submit), or dominate an opponent (soundly defeat, outperform, or display considerably greater strength than an opponent; Burgoon, Johnson, & Koch, 1998 ; Cheng et al., 2013 ; Cheng, Tracy, & Henrich, 2010 ; Cheng, Tracy, Ho, & Henrich, 2016 ; Ellyson & Dovidio, 1985 ). Additionally, dominance is often used to describe a general style of relating to others that expresses the explicit and implicit motivation for status and status-seeking behavior more broadly (Anderson & Kilduff, 2009 ): A social hierarchy may be a “status hierarchy” or a “dominance hierarchy” (Chase, Tovey, Spangler-Martin, & Manfredonia, 2002 ; see Sapolsky, 2005 ; Mazur, Welker, & Peng, 2015 ). Figure 17.1 visually demonstrates the relationships between the terms competition, dominance, deference, social hierarchy, status ranking , and status-seeking behavior .

Visual description of terms related to status and social hierarchy.

Although “dominance” is typically used as the catch-all concept for general behavior involved in the attainment and maintenance of high status, recent research has determined that there are at least two distinct, yet equally viable, cognitive and behavioral strategies for gaining influence—dominance and prestige (Cheng et al., 2010 , 2013 ; Cheng & Tracy, 2014 ). In this view, dominance refers to a more coercive, physical or psychologically aggressive, intimidating, and conflict-based style in which an individual attains power by demanding deference. Eminence (Kemper, 1990 ) or prestige (Cheng et al., 2013 ) is a style of achieving high status through the demonstration of competence, likability, and prosociality and is characterized by the voluntary deference, attention, and respect of others. Given this distinction, continued research on the complexities of competition- and status-related behavior, including the underlying biological influence, will expand understanding of these important and consequential social interactions. Due to the more broad interpretation of “dominance” (meaning status-seeking behavior) in the social neuroendocrinology literature, the remainder of this chapter will operate under this generalized definition.

Cortisol, Stress, the Social Hierarchy, and Competition

Cortisol is well known for its positive relationship with stress, both psychological and physical. Acute psychological experiences of stress, particularly social-evaluative stress combined with a perceived lack of control over one’s environment and outcomes, produces reliable and transient increases in cortisol (reviewed by Dickerson & Kemeny, 2004 ). Short-term elevations in cortisol, secreted as an end-product of the hypothalamic-pituitary-adrenal axis and via rapid and direct sympathetic stimulation of the adrenal gland (Engeland & Arnhold, 2005 ; Tsigos & Chrousos, 2002 ), provide immediate advantages for escaping or managing a stressful stimulus (e.g., mobilization and breakdown of glucose). Long-term activation or dysregulation of this system is energetically costly and can result in deleterious effects on physical and psychological health and immune system functioning (Cohen et al., 2012 ; McEwen, 1998 ; Whitworth, Williamson, Mangos, & Kelly, 2005 )—a process known as “allostatic load” (McEwen, 1998 , 2000 , 2004 ). Thus, individual differences in basal cortisol are interpreted as a reliable predictor of allostatic load (e.g., Goymann & Wingfield, 2004 ).

Basal Cortisol and Social Rank

Social hierarchy rank is thought to be inversely related to basal cortisol due to the increasing life adversity and resulting allostatic load experienced by increasingly lower ranking individuals (Knight & Mehta, 2014 ; Sapolsky, 2005 ). Early studies with nonhuman animals supported this notion; for example, Sapolsky ( 1982 ) showed that high-ranking males demonstrated significantly lower baseline cortisol in baboons than their more subordinate counterparts. One of the first studies in humans corroborated this finding: Among a relatively small sample of male Dominican villagers, lower cortisol levels were related to higher peer ratings of likability and influence (Decker, 2000 ). At the society level, there is a reliable and robust negative relationship between basal cortisol levels and SES (Cohen, Doyle, & Baum, 2006 ; reviewed by Knight & Mehta, 2014 ).

Recent research in humans has expanded on the cortisol–status relationship. In a relatively large sample of individuals enrolled in an executive education program, Sherman et al. ( 2012 ) showed that leaders (those responsible for managing others) had significantly lower cortisol and anxiety than nonleaders. In a follow-up study among leaders, leadership level (having and managing more subordinates and with greater authority) produced the same effect. Additionally, sense of control (generated using a measure of personal sense of power) mediated the relationship between leadership level and cortisol, as well as anxiety. However, studies of collegiate athletes have shown no direct relationship between cortisol and peer ratings of leadership ability, a proxy for status (Edwards & Casto, 2013 ; Edwards, Wetzel, & Wyner, 2006 ). Using social network analysis, Kornienko, Clemans, Out, and Granger ( 2014 ) demonstrated that among a competitive pool of first-year nursing students, high cortisol levels were associated with low gregariousness, the number of perceived friends within the network of nursing students, but not popularity, the number of network members who perceived them as friends. In a follow-up study among members of a large mixed-sex collegiate marching band, higher basal cortisol was also related to an inability to maintain friendships within the network (i.e., greater turnover in friendships over a two-month period; Kornienko, Schaefer, Weren, Hill, & Granger, 2016 ).

Cortisol Reactivity to a Status Challenge

The direction and strength of the cortisol response to stress depends on a multitude of psychological and contextual factors (Dickerson & Kemeny, 2004 ; Knight & Mehta, 2014 ; Kudielka, Hellhammer, & Wüst, 2009 ). Cortisol increase in response to a stressor could be considered adaptive (e.g., benefiting social status) or maladaptive (e.g., a sign of dysregulation or overreactivity) depending on the context (e.g., competition) and the timing and magnitude of the response (Aschbacher et al., 2013 ; Shirtcliff, Peres, Dismukes, Lee, & Phan, 2014 ).

Social rank appears to be one important factor for predicting patterns of cortisol reactivity to stress (e.g., Hellhammer, Buchtal, Gutberlet, & Kirschbaum, 1997 ; Sapolsky, 1982 ), with higher social status producing what researchers interpret as more adaptive responses, depending on the task and context (Akinola & Mendes, 2014 ; Shirtcliff et al., 2014 ). In one study, women whose cortisol levels did not habituate after repeated exposure to a stressor (i.e., a maladaptive response) subjectively rated themselves lower in SES (Adler, Epel, Castellazzo, & Ickovics, 2000 ). Under the condition of social-evaluative threat, men and women with high subjective social status (perceived rank among dormitory peers) showed significantly larger cortisol increases in a single session of the Trier Social Stress Test (TSST) than men and women who rated themselves low in status (Gruenewald, Kemeny, & Aziz, 2006 ). Cortisol increase, in this sense, could reflect a greater mobilization of energy and activity required to defend one’s status when status is indeed at stake, with a nonincrease in cortisol reflecting a more maladaptive (i.e., blunted) physiological response threat. However, the adaptive function of elevated cortisol would likely depend on the social context. For example, increased cortisol in response to social stressors, in some studies, appears to predict subsequent risky decision making (van den Bos, Harteveld, & Stoop, 2009 ; reviewed by Starcke & Brand, 2012 ), a behavior that could be beneficial in contexts where risk taking is advantageous (i.e., choosing to fight rather than flee when status is relatively high), but could be deleterious in other contexts (e.g., choosing to fight rather than flee when status is relatively low). However, there is some evidence in men and women dyads that transiently increased cortisol levels predict subsequent prosocial behaviors and cognitive states such as affiliation (in women dog handlers, Sherman, Rice, Jin, Jones, & Josephs, 2017 ) and feelings of interpersonal closeness (in men participating in the TSST, Berger, Heinrichs, von Dawans, Way, & Chen, 2016 ) that effectively buffer the negative psychological and physiological effects of stress (Cohen & McKay, 1984 ).

Situational factors also appear to moderate the relationship between status and cortisol reactivity. In at least two studies, individuals who were high on indices of dominance motivation (basal testosterone; implicit power motivation) showed greater cortisol increases across a competition for status, but only when that competition resulted in defeat (Mehta, Jones, & Josephs, 2008 ; Wirth, Welsh, & Schultheiss, 2006 ). Directly testing the moderating effects of social context on the status–cortisol reactivity relationship, Knight and Mehta ( 2017 ) manipulated both the status position and hierarchy stability of men and women competing in a mock job interview. Participants assigned to a high-status role (“manager”) showed reduced cortisol reactivity and better performance compared to participants assigned to the low-status role (“builder”), but only if the assigned roles were fixed (i.e., a stable hierarchy). When told that the assigned roles could change based on performance in the interview (i.e., an unstable hierarchy), high status increased cortisol reactivity and did not result in better performance compared to low status.

In real-world athletic competition, where status is formally contested, cortisol significantly increases over the match period in men and women (e.g., Casto, Elliott, & Edwards, 2014 ; Edwards et al., 2006 ; Edwards & Kurlander, 2010 ; reviewed by Casto & Edwards, 2016a ), an effect that is likely influenced, to some degree, by the physical stress of exercise (Copeland, Consitt, & Tremblay, 2002 ; M. S. Tremblay, Copeland, & van Helder, 2005 ; Viru et al., 2010 ). Contrary to the aforementioned study where high rank predicted a higher cortisol increase across the TSST, competition losers appear to show relatively higher increases in cortisol compared to winners (e.g., Bateup, Booth, Shirtcliff, & Granger, 2002 ; Jiménez, Aguilar, & Alvero-Cruz, 2012 ).

Testosterone, Status, and Status-Seeking Behavior

Several decades of research have revealed a generally positive and bidirectional relationship between testosterone, social status, and status-related behavior. Initial observationally and empirically based models of this relationship, the biosocial model of status (Mazur, 1985 ) and the challenge hypothesis (Wingfield, Hegner, Dufty, & Ball, 1990 ), have provided a theoretical basis for this research. Detailed historical accounts and an extended description of these models are published elsewhere (biosocial model of status—Mazur & Booth, 1998 ; Mazur, 2017 ; challenge hypothesis—Archer, 2006 ; Wingfield, 2017 ). The challenge hypothesis asserts that baseline levels of testosterone in monogamously mating birds regulate reproductive development during breeding season, increase with periods of territorial aggression under conditions of social instability (in response to male–male contests for status and sexual partners), and decrease with the expression of parental care (Wingfield, 2017 ). This hypothesis has been extended to humans to explain, more generally, testosterone increases in response to “competitive situations between young men” (Archer, 2006 , p. 322).

The biosocial model of status, initially informed by empirical studies with nonhuman primates and men engaged in athletic competition, proposes that stable baseline levels of testosterone predict status-related behavior. Furthermore, this model proposes that testosterone should increase in response to status gained and decrease in response to status lost. Specifically, high-testosterone individuals are expected to behave more dominantly and, as a result of status gained from this dominance, demonstrate rising levels of testosterone to promote future competitive behavior. Likewise, low-testosterone individuals are expected to behave more submissively and, as a result of status lost from this deference, demonstrate falling levels of testosterone. That is, transient shifts in testosterone represent the physiological mechanism underlying each stage of the competition, status-sorting, and status-seeking processes depicted in Figure 17.1 . Although originally thought to be an acute mechanism regulating even subtle dominance and deference signals in everyday interactions with others (Mazur, 1985 ), evidence of substantial testosterone fluctuations resulting from status contests may be specific to more formal competitive contexts (Mazur et al., 2015 ).

The decades that followed the dissemination of the biosocial model for status and the challenge hypothesis have produced an abundance of empirical research testing specific predictions from these models in the context of human competition. There are several recent comprehensive reviews and meta-analyses that summarize the findings from this literature (Carré & Olmstead, 2015 ; Casto & Edwards, 2016a ; Geniole, Bird, Ruddick, & Carré, 2017 ; Hamilton et al., 2015 ; G. A. Oliveira & Oliveira, 2014 ). Table 17.1 provides a list of specific hypotheses that can be derived from the biosocial model of status and challenge hypothesis regarding the relationships between testosterone and status ranking, status-seeking motivation, and status-seeking behavior. Some of these hypotheses have been tested empirically more than others (e.g., #1 and #2 more than #5). Among the specific hypotheses that have been well tested, nearly all of them have been supported by some studies but also not supported by others (#1: Burnham, 2007 ; Cashdan, 2003 ; Dabbs & Morris, 1990 ; Josephs, Sellers, Newman, & Mehta, 2006 ; Mehta, DesJardins, van Vugt, & Josephs, 2017 ; R. E. Tremblay et al., 1998 ; van Bokhoven et al., 2006 ; Wirth & Schultheiss, 2007 ; #2—Carré, Putnam, & McCormick, 2009 ; Grant & France, 2001 ; Schultheiss, 2007 ; Sellers, Mehl, & Josephs, 2007 ; Welker & Carré, 2015 ; #3—Cashdan, 1995 ; Purifoy & Koopmans, 1979 ; Zyphur, Narayanan, Koh, & Koh, 2009 ; Apicella, Dreber, & Mollerstrom, 2014 ; Bateup et al., 2002 ; #4—Carré, Campbell, Lozoya, Goetz, & Welker, 2013 ; Costa & Salvador, 2012 ; Jiménez et al., 2012 ; Norman, Moreau, Welker, & Carré, 2015 ; #5—Peters, Hammond, Reis, & Jamieson, 2016 ; #6—Apicella et al., 2014 ; Carré & McCormick, 2008 ; Casto & Edwards, 2016b ; Casto et al., 2014 ; Edwards et al., 2006 ; Guezennec, Lafarge, Bricout, Merino, & Serrurier, 1995 ; Steiner, Barchard, Meana, Hadi, & Gray, 2010 ; van der Meij, Buunk, Almela, & Salvador, 2010 ; #7—Schultheiss, Campbell, & McClelland, 1999 ; Schultheiss & Rohde, 2002 ; Schultheiss et al., 2005 ; #8—Bos, Hermans, Ramsey, & Van Honk, 2012 ; Carré, Baird-Rowe, & Hariri, 2014 ; Carré & McCormick, 2008 ; Hermans, Ramsey, & van Honk, 2008 ; Mehta & Josephs, 2006 ; Mehta, van Son, et al., 2015 ; Welling, Moreau, Bird, Hansen, & Carré, 2016 ). To clarify general concepts that have emerged from the research testing these hypotheses, we summarize the three main empirically supported predictions from this literature next.

Prediction #1: Basal Testosterone Is Related to Status-Motivated and Dominance Behavior

One of the original and more simplistic predictions of the biosocial model of status is that individuals in high-status positions in a social hierarchy would have relatively higher baseline testosterone levels. Higher testosterone would not only motivate behaviors to achieve such a high-status position but also increase further as a result of its attainment. Although some earlier research supported this prediction in humans and other species (Dabbs & Morris, 1990 ; Purifoy & Koopmans, 1979 ; Sapolsky, 1982 ), subsequent research has shown that hierarchical social ranks such as socioeconomic status and peer-rated ranking are not directly related to testosterone levels in men and women (Cashdan, 1995 ; Edwards et al., 2006 ; Mehta & Josephs, 2010 ; earlier work reviewed by Mazur & Booth, 1998 ; Newman, Sellers, & Josephs, 2005 ). However, those who want status may not always have it, and the complexities (e.g., context dependence) of a social hierarchy may make it difficult to directly link social status and absolute testosterone levels. Indeed, a recent study of male employees working for corporate businesses revealed that testosterone levels among these men were positively related to self-reported authoritarian leadership style, but only for those who were not in management (leadership) positions (van der Meij, Schaveling, & van Vugt, 2016 ). Consistent with meta-analytic data on the relationship between actual leadership position and testosterone (van der Meij et al., 2016 ), managers did not have higher testosterone levels on average than their subordinate workers. Rather than directly predicting status position, basal testosterone appears to predict how individuals respond to shifts in status. That is, high-testosterone individuals respond negatively (e.g., poorer performance on a spatial or verbal test) to a drop in status, whereas low-testosterone individuals respond neutrally or negatively to a rise in status (Josephs, Newman, Brown, & Beer, 2003 ; Josephs et al., 2006 ; Mehta et al., 2008 ; Newman et al., 2005 ).

Due to the complexities of achieving high-status or leadership positions, basal testosterone may better predict personality characteristics and behaviors motivated toward achieving status (whether or not those efforts are successful). Indeed, basal testosterone is considered an important “personality variable” that predicts dominance behaviors in various contexts (Newman & Josephs, 2009 ; Sellers et al., 2007 ; Mehta et al., 2008 ). In competition, testosterone levels have been positively related to perceptions of one’s personal success (Casto, Rivell, & Edwards, 2017 ), competitive decision making and subsequent task confidence (Eisenegger, Kumsta, Naef, Gromoll, & Heinrichs, 2017 , but see Apicella et al., 2011 ), and competitive task persistence (Welker & Carré, 2015 ). However, testosterone has been found to be negatively related to the ability to accurately judge the thoughts and feelings of others (empathic accuracy) in laboratory and real-world settings, an aspect of cognition that may have negative consequences on other’s perceptions of one’s leadership ability (Ronay & Carney, 2013 ). A recent study employing an economic decision-making task in which dyadic status relationships are determined during play (dominant–submissive, dominant–dominant, and submissive–submissive; the hawk-dove game) showed that baseline testosterone was positively correlated to taking a dominant position (Mehta et al., 2017 ). Although there are mixed reports on the direct relationship between testosterone and self-reported dominance (e.g., Cobey, Nicholls, Leongómez, & Roberts, 2015 ; Grant & France, 2001 ; Neave, Laing, Fink, & Manning, 2003 ), trait dominance has emerged as an important moderator of testosterone’s relationship to dominant, competitive, or aggressive behavior (Mehta, van Son, et al., 2015 ; for review, Carré & Archer, 2017 ). For example, in one study of men competing for the affection of a woman, high-testosterone individuals displayed more dominant behaviors, but the relationship was specific to those who self-identified as dominant (Slatcher, Mehta, & Josephs, 2011 ). Furthermore, the relationship between testosterone and social status or dominance behaviors appears to also be moderated by basal levels of cortisol (see the section on the dual-hormone hypothesis).

Prediction #2: Testosterone Levels Are Transiently Altered During Contests for Status, More Often in the Positive Direction for Winners Than Losers

A second main prediction of the biosocial model of status is that gaining status through winning a dominance contest should increase testosterone levels from baseline, whereas losing should decrease testosterone levels (i.e., the winner–loser effect , also referred to as the winner effect ; reviewed by Casto & Edwards, 2016a ). Support for this prediction has been found in studies of laboratory competition, where testosterone levels increased across competition for those who won but decreased for those who lost (e.g., Apicella et al., 2014 ; Carré et al., 2013 ; Costa & Salvador, 2012 ; Norman et al., 2015 ). However, other studies have found that testosterone increases across competition regardless of the outcome (Carré & McCormick, 2008 ; Henry, Sattizahn, Norman, Beilock, & Maestripieri, 2017 ; Steiner et al., 2010 ; van der Meij et al., 2010 ). Summarizing the extant literature on the winner–loser effect, Carré and Olmstead ( 2015 ) concluded that a number of studies have reported that male winners had elevated testosterone levels relative to losers, but that a nearly equal number of studies have failed to find such an effect. For women, less studied in general, the proportion of studies that show null findings is even greater (Carré & Olmstead, 2015 ). However, a recent meta-analysis of 60 effect sizes on the winner–loser effect (Geniole et al., 2017 ) determined that winners do in fact show an increase in testosterone compared to losers, who on average experience no change in testosterone. Although the effect was not moderated by sex, the effect was only significant in men (men Cohen’s d = 0.23; women d = 0.14).

However, the winner–loser effect in this meta-analysis appears to depend on important contextual factors; average effect sizes were moderately large only when the studies were conducted in nonlaboratory testing locations (e.g., athletic competitions), when the outcome (win or loss) was determined naturally (not contrived or manipulated), when the competition duration was greater than 15 minutes, and when the precompetition saliva sample was taken more than 10 minutes before competition (Geniole et al., 2017 ). Even though each of these moderators independently accounted for effect size differences in the winner–loser effect, they are not mutually exclusive factors; studies of the hormonal response to naturally occurring athletic competition are always outside the laboratory, where the competition outcome cannot be manipulated, and usually last longer than 15 minutes. Thus, the winner–loser effect is less well supported in experimental designs outside of formal athletic competition. As a matter of convenience and limited access to athletes immediately prior to and following a match, participants in studies of athletic competition are often asked to give their precompetition sample more than 10 minutes before the match begins and more than 10 minutes after the competition has ended (e.g., Jiménez et al., 2012 ). These studies have been more likely to show dramatic win–loss differences in testosterone change. For studies that acquired samples immediately prior to and following competition (mostly with women athletes), testosterone increases significantly across the competition period regardless of outcome (e.g., Casto et al., 2014 , 2017 ; Casto & Edwards, 2016b ; Edwards et al., 2006 ; Edwards & Kurlander, 2010 ).

The issue of lab versus nonlab competition in determining the emergence of the winner–loser effect is currently unresolved. What the laboratory context gains in experimental control, it perhaps loses in being able to sufficiently activate competitive motivation and in being able to create a more realistic social setting where actual status is at stake. What field studies gain in ecological validity, they lose in experimental control. One of the most important potential confounds in the hormonal response to athletic competition is physical exertion, a factor that can elevate testosterone levels independent of competition and the psychological experience of gaining or losing social status (Copeland et al., 2002 ; M. S. Tremblay et al., 2005 ; Viru et al., 2010 ). However, efforts to quantify physical exertion in studies of the testosterone response to athletic competition (e.g., blood lactate, number of minutes played, self-reported physical exertion, observer-rated physical exertion) have not found any significant correlations between these metrics of exertion and testosterone (Aguilar, Jiménez, & Alvero-Cruz, 2013 ; Casto & Edwards, 2016b ; T. Oliveira, Gouveia, & Oliveira, 2009 ; Suay et al., 1999 ). Furthermore, one of us (KVC) has recently collected saliva samples from trained men and women rifle shooters competing in intra- and intersquad rifle competition, a sport that requires the athletes to remain as still as possible. As with other sporting competitions that require more movement, rifle competition results in significant elevations in testosterone on average and for the majority of the athletes sampled over the course of competition (+6 to 41 percent change, Casto & Edwards, unpublished).

Prediction #3: Transient Increases in Testosterone (Associated With a Status Challenge) Promote Future or Subsequent Status-Seeking Behavior

More recently, researchers have begun to explore the functional significance of transient increases in testosterone, that is, the adaptive consequence on subsequent cognition and behavior (for initial review, see Carré, McCormick, & Hariri, 2011 ). If, in fact, testosterone increases in some individuals and under certain contexts, does this change serve a beneficial purpose for attaining or maintaining status in the future? The desire to compete again may reflect elevated dominance motivation when the individual has the potential to improve status rank (i.e., when the potential benefits outweigh the risks). In men, increases in testosterone during competition predict the subsequent decision to compete again against the same opponent in losers (Mehta & Josephs, 2006 ) and aggressive individuals (Carré & McCormick, 2008 ), or a different opponent in decisive winners (Mehta, Snyder, Knight, & Lasseter, 2015 ).

Testosterone increase during competition may also predict subsequent aggressive behavior in men (Carré et al., 2009 ), an effect that appears to be moderated by trait anxiety (Norman et al., 2015 ). Also in men, testosterone increases associated with monetary wins and losses relate to future financially risky choices (Apicella et al., 2014 ). Increased testosterone has also been related to a more positive subsequent performance in athletic contexts (Cook & Crewther, 2012a , 2012b ). In one study of women soccer players, the higher an athlete’s testosterone remained within the 30 minutes after competition, the greater her willingness to reconcile with her opponent—a prosocial strategy for status maintenance (Casto & Edwards, 2016c ). Extending beyond correlational evidence, studies of exogenously administered testosterone have shown that, for periods of time up to four hours after administration, participants administered testosterone demonstrate altered cognitions and behaviors that may promote status seeking or aid in status maintenance in competitive contexts (e.g., Bos, Terberg, van Honk, 2010 ; Eisenegger, Naef, Snozzi, Heinrichs, & Fehr, 2010 ; Hermans, Putman, Baas, Koppeschaar, & van Honk, 2006 ; Mehta, van Son, et al., 2015 ; Radke et al., 2015 ; van Honk, Montoya, Bos, Van Vugt, & Terburg, 2012 ; reviewed by Eisenegger, Haushofer, & Fehr, 2011 ). For example, a series of studies by van Honk and colleagues suggests that testosterone administration, after several hours, increases threat vigilance and reduces fear-potentiated startle, responses that are matched, in some cases, with activation of brain areas associated with emotional reactivity (Hermans et al., 2006 , 2008 ; van Honk et al., 1999 ; for review, see Carré & Olmstead, 2015 ). Testosterone administration also may reduce cognitive reflection in men (Nave, Nadler, Zava, & Camerer, 2017 ) and behaviors thought to reflect empathy in women (Hermans et al., 2006 ; van Honk & Schutter, 2007 ; van Honk et al., 2011 ; Wright et al., 2012 ). Additionally, other research has shown that after receiving a dose of exogenous testosterone, men appear to perceive themselves as more physically dominant (Welling et al., 2016 ).

Testosterone and Cortisol Interact to Predict Social Status: The Dual-Hormone Hypothesis

Early predictions that testosterone should directly and positively predict social status have failed to garner unanimous empirical support (e.g., Neave et al., 2003 ; review by Mazur & Booth, 1998 ; Mehta & Josephs, 2010 ). Evidence that cortisol inhibits, suppresses, or otherwise antagonizes testosterone secretion and action at target tissues (Burnstein, Maiorino, Dai, & Cameron, 1995 ; Chen, Wang, Yu, Liu, & Pearce, 1997 ; Johnson, Kamilaris, Chrousos, & Gold, 1992 ) combined with initial behavioral evidence that these two hormones might interact to predict aggression (Dabbs, Jurkovic, & Frady, 1991 ; Popma et al., 2007 ) suggests that a more integrative approach to the hormone–social status relationship was necessary. Following these early indications, Mehta and Josephs ( 2010 ) proposed that “only at low levels of cortisol should higher testosterone encourage higher status” (p. 898)—a statement that serves as the basis of the dual-hormone hypothesis . Testing this hypothesis, Mehta and Josephs ( 2010 ) indeed showed that only if an individual was relatively low in cortisol did testosterone positively relate to dominance behaviors when instructed to act as a leader (Study 1, in men and women) and when deciding to compete again following a defeat in a rigged puzzle competition (Study 2, in men).

Several novel replications of the dual-hormone effect have been published (for an earlier review, see Mehta & Prasad, 2015 ). In varsity women athletes, Edwards and Casto ( 2013 ) showed that testosterone positively related to actual status as ranked by teammates, but only if the athlete had relatively low levels of cortisol. As with the original studies conducted by Mehta and Josephs, the relationship trended toward an inverse relationship between testosterone and status for those with high levels of cortisol. In a sample of male business executives, testosterone positively predicted the number of subordinates over which an executive had authority (but not income or education level) only if the individual had relatively low cortisol (Sherman, Lerner, Josephs, Renshon, & Gross, 2016 ). Using social network analysis, Ponzi, Zilioli, Mehta, Maslov, and Watson ( 2016 ) showed that among professional male rugby players, participants with low cortisol (but not those with high cortisol) demonstrated higher network centrality (a proxy for social status) in measures of “betweenness” and popularity. Recently published research has even extended the dual-hormone hypothesis to collective hormone profiles in groups. Akinola, Page-Gould, Mehta, and Lu ( 2016 ) measured basal testosterone and cortisol in a large sample of MBA students organized into (diverse and mixed-sex) groups of three to six members and asked to compete against other groups in a business-related decision-making task. Collectively high testosterone was significantly and positively related to group performance, but only if that group had relatively low collective cortisol. Although null findings are less likely to be published, there have been other studies that have also failed to support the dual-hormone hypothesis (e.g., Geniole et al., 2013 ; Mehta et al., 2017 ).

A high-testosterone/low-cortisol profile has also been found to relate to antisocial attitudes (Sollberger, Bernauer, & Ehlert, 2016 ) and externalizing psychopathology in adolescents (Tackett, Herzhoff, Harden, Page-Gould, & Josephs, 2014 ), factors that would seem to be detrimental to social status in modern society. However, other studies have shown that a high-testosterone/high-cortisol profile (the inverse of the original dual-hormone effect) predicts deviant or psychopathic traits (Welker, Lozoya, Campbell, Neumann, & Carré, 2014 ), reactive aggression, and self-reported feelings of dominance (Denson, Mehta, & Ho Tan, 2013 ). However, at least one study has shown that the interaction between testosterone and cortisol has no relationship to antisocial, socially deviant behavior in a large sample (4,462) of male U.S. Army veterans (Mazur & Booth, 2014 ).

How a high-testosterone/low-cortisol individual (or group) behaves in everyday interactions with others to result in higher status is not yet fully understood. This hormone profile may manifest in a style of interacting that balances a desire for status with a desire to affiliate and promote social bonding among others in the group or network. Or these individuals could have a personality that balances status and power motivation with relaxed confidence and low anxiety—a personality profile that is likable, and therefore more likely to garner the support of others required to attain and maintain status. Perhaps high-testosterone/high-cortisol individuals interact with others with high anxiety, low confidence, aggression, or desperation, effectively thwarting attempts to actually achieve status. Future research should explore, more in depth, personality correlates and behavioral interaction styles of high-testosterone/low-cortisol and high-testosterone/high-cortisol individuals. It is also possible that the mechanism explaining the dual-hormone effect is an interaction between the negative effects of chronically elevated cortisol and physiological processes involved in testosterone’s ability to drive status-seeking behavior. That is, perhaps allostatic load–related stress and resulting high basal cortisol dampens or inhibits mechanisms for status motivation and related behaviors (i.e., a high-testosterone–status relationship). Future research should expand the theoretical basis and practical applications of the dual-hormone hypothesis. Additionally, although this hypothesis originally concerned the interaction between basal testosterone and basal cortisol, how testosterone and cortisol changes interact to predict status and performance-related behavior (e.g., Mehta, Mor, Yap, Prasad, 2015 ) is a topic of interest for future research.

Moderators of the Relationships Between Competition and Testosterone

Recent research has exposed increasingly complex qualifiers, moderators, and extenuating circumstances that impact both the effect of competition on testosterone change and the effect of testosterone change on subsequent behavior. These moderating variables fall under two broad categories: (1) person and (2) context factors (reviewed by Carré et al., 2010 ; Casto & Edwards, 2016a ; Hamilton et al., 2015 ; G. A. Oliveira & Oliveira, 2014 , Salvador & Costa, 2009 ). Table 17.2 provides a categorical list of the person and context factors that have been studied and appear to be important for influencing either the relationship between basal testosterone and dominance behavior, the testosterone response to competition, or the effect of testosterone change on subsequent status-seeking behavior.

Edwards and Casto ( 2013 ); Mehta and Josephs ( 2010 ); Ponzi et al. ( 2016 ); Sherman et al. ( 2016 ); Wu et al. ( 2017 ).

van Honk et al. ( 2012 ).

Carré et al. ( 2013 ).

Schultheiss and Rohde ( 2002 ); Schultheiss et al. ( 2005 ).

Mehta, van Son, et al. ( 2015 ).

Carré and McCormick ( 2008 ).

Costa and Salvador ( 2012 ).

Maner et al. ( 2008 ); Norman et al. ( 2015 ).

Welker et al. ( 2017 ).

Mazur and Lamb ( 1980 ); Mehta and Josephs ( 2006 ); Mazur et al. ( 1997 ); Zilioli and Watson ( 2013 ).

Costa, Serrano, and Salvador ( 2016 ); Salvador and Costa ( 2009 ).

Casto et al. ( 2017 ); Gonzalez-Bono et al. ( 1999 ); Oliveira et al. ( 2014 ).

Mehta, Snyder, et al. ( 2015 ).

Apicella et al. ( 2014 ); Carré et al. ( 2013 ); Costa and Salvador ( 2012 ); Jiménez et al. ( 2012 ); Norman et al. ( 2015 ); Zilioli and Watson ( 2014 ).

Zilioli and Watson ( 2014 ).

Mehta, Snyder, et al. ( 2015 ); Zilioli, Mehta, and Watson ( 2014 ).

Oxford, Ponzi, and Geary ( 2010 ); Wagner, Flinn, and England ( 2002 ).

Mehta, Wuehrmann, and Josephs ( 2009 ).

van Anders and Watson ( 2007 ).

Carré ( 2009 ); Carré et al. ( 2006 ); Neave and Wolfson ( 2003 ).

Early studies of the testosterone response to competition included mood as an additional variable based on the notion that testosterone should increase when dominance is achieved through winning, but only if the individual experienced high positive emotions regarding the win (Gladue, Boechler, & McCaul, 1989 ; Mazur & Lamb, 1980 ; McCaul, Gladue, & Joppa, 1992 ; Mazur, Susman, & Edelbrock, 1997 ). In the first study published on the testosterone response to status enhancement and competition in humans, Mazur and Lamb ( 1980 ) stated that “when a man achieves a rise in status through his own efforts, and he has an elation of mood over the achievement, then he is likely to have a rise in testosterone” (p. 236). Although subsequent research has shown the importance of mood as an intervening factor (Mehta & Josephs, 2006 ; Zilioli & Watson, 2013 ), state-level mood may serve only as a proxy for a more important personality factor: dominance motivation. Arguably, only those who have a strong motivation for dominance would experience elation upon achieving it through competition. Indeed, Schultheiss et al. ( 1999 ) wrote, “It would seem reasonable to assume that personality factors may moderate individuals’ testosterone responses to succeeding or failing at a dominance contest” (p. 234).

Implicit power motivation is the degree with which an individual derives reward from “having physical, and mental or emotional impact” on others (Stanton & Schultheiss, 2009 , p. 942). Those higher in implicit power motivation tend to be more likely to show an increase in testosterone (and cortisol) in response to competition, particularly under the context of a win (Schultheiss & Rohde, 2002 ; Schultheiss et al., 1999 , 2005 ; Wirth et al., 2006 ). However, this relationship appears to depend on sex—with stronger, more consistent relationships found for men than for women (Stanton & Schultheiss, 2007 ). Although implicit power motivation has received the most empirical attention, self-reported trait dominance (e.g., Carré et al., 2009 ; Mehta, van Son, et al., 2015 ) and competitiveness (Casto, 2016 ; Costa & Salvador, 2012 ) may also play an equally important role in regulating testosterone–status relationships and testosterone response to competition. In Mehta, van Son, et al. ( 2015 ), testosterone administration in women resulted in increased competitive decision making (i.e., a greater percentage of the trials in which the competitor chose to compete again afterward), but only if the participant scored high on a personality measure of dominance motivation and also won the competition. For participants who lost the competition, testosterone administration decreased competitive decision making regardless of individual differences in dominance motivation.

Social context also moderates the relationship between testosterone and competitive performance. In one of the first demonstrations of this, Newman et al. ( 2005 ) showed that high-testosterone individuals placed in a high-status position performed well on a spatial and verbal task, but high-testosterone individuals placed in a low-status position performed poorly in comparison. This effect, dubbed the mismatch effect (Josephs et al., 2006 ), explains how situational constraints that contrast with self-perceptions hamper status-seeking efforts or competitive performance when status is threatened. The mismatch effect is also relevant for group-level performance. Among college students assigned to work in a group on class assignments for an entire semester, the greater the mismatch between testosterone and status rank within the group (i.e., the more negative the relationship), the lower the group’s collective self-efficacy (i.e., the lower their shared confidence in ability to succeed), a measure that is indicative of group functioning and performance (Zyphur et al., 2009 ).

Another social context factor that could affect that relationship between testosterone and status is hierarchy stability, the degree with which one’s status could readily be changed. Zilioli, Mehta, and Watson ( 2014 ) proposed that under conditions of status instability, there should be a reversal of the winner–loser effect, whereby winners should decrease in testosterone and losers should increase. This amendment to the biosocial model of status, termed the status instability hypothesis , is based on the notion that if the function of a testosterone increase is to promote future status-seeking behavior, then winners who just barely won should be less motivated to compete again because of a high chance of moving down in status. However, losers who just barely lost should be more motivated to compete again because of a high chance of moving up in status. Testing this hypothesis with women, Zilioli et al. ( 2014 ) showed that women competing in a number-tracing task who won by only a small margin (given feedback of their narrow win throughout multiple trials in the task) decreased in testosterone from before to after the task, whereas women who lost by a narrow margin slightly increased in testosterone (Study 1). In a second companion study, under conditions of relative performance uncertainty, women competing in Tetris showed greater competition-related declines in testosterone than women who lost (Study 2). A simultaneous study in men competing in Tetris on two consecutive days (Zilioli & Watson, 2014 ) found evidence that day 2 testosterone increased significantly more across the competition period for those whose status reversed from the day 1 competition (i.e., a day 1 win followed by a day 2 loss, or vice versa; an unstable hierarchy) compared to men who won both days or lost both days (a stable hierarchy). When wins and losses where manipulated to be either “clear” or “narrow,” Wu, Eisenegger, Zilioli, Watson, and Clark ( 2017 ) showed that testosterone levels decreased significantly for narrow winners, but only if their basal cortisol levels were relatively high. Despite some empirical support in these studies for the status instability hypothesis with regard to the direction of testosterone change, no studies have demonstrated the reverse win–lose effect pattern consistent with this hypothesis nor have any studies shown that this pattern of testosterone change predicts subsequent motivation to compete again—a main tenet of the hypothesis. To make more informed inferences about the functional significance of testosterone increases and decreases resulting from status gained and lost within stable and unstable social hierarchies, future research should attempt to corroborate testosterone reactivity with postcontest motivational states related to social status.

The wide array of moderating factors complicates attempts to understand relationships between hormones and competition, but is rightly indicative of the complexity of human nature and the intricate social context of status striving. Figure 17.2 displays an updated theoretical framework for future studies testing both basal and dynamic testosterone–status relationships. For a more complete understanding of the role of person and context factors, future research will require relatively large sample sizes to properly test moderation and other complex interactions.

Theoretical model for the testosterone–social status relationship.

Additional Future Directions

The field of social-behavioral neuroendocrinology is a burgeoning area of scientific inquiry. Although the last three decades have produced foundational discoveries, there are many unanswered questions and new directions for future research.

The Role of Trait Competitiveness in Predicting Testosterone Response to Competition

According to social comparison theory , the drive to compete is derived from the basic human need to reduce uncertainty between one’s own performance and the performance of others in order to maintain superior relative position (Festinger, 1954 ; Garcia, Tor, & Schiff, 2013 ). For some, this drive is sufficiently strong to prompt greater efforts to engage and succeed in situations where relative judgments about performance are made. Because comparison to others through competition is how relative social status is determined, individual differences in competitiveness, the “desire to win in interpersonal situations” (Smither & Houston, 1992 , p. 408), may be a direct predictor of relative social status or the motivation to acquire it. Given the apparent connection between status seeking and testosterone, competition-related changes in testosterone levels may depend on individual differences in trait competitiveness. Welker and Carré ( 2015 ) recently reported that basal testosterone in men correlates with persistence in attempting to solve puzzles made intentionally unsolvable by the experimenters. Although conceptually different than competitiveness per se, task persistence is a core quality of highly competitive individuals. Future research should consider including competitiveness as a person factor regulating hormonal response to competition and explore conceptual and statistical relationships between competitiveness and trait dominance, as well as power motivation.

Group Dynamics and Social Network Analysis

Real-world and everyday contests for social status occur in the context of complex social networks involving multilayered social group interactions with others. Despite the importance of group dynamics in status attainment and maintenance, previous research on the social neuroendocrinology of dominance has largely focused on the individual—testing participants in solitary rooms with little to no interaction with their “opponent.” It is reasonable to assume that status lost or gained in contexts where participants have no current or future potential interaction with each other would have, at best, a minor impact on one’s physiology or behavior (i.e., relative standing to an unknown person may have little relevance to perceptions of social status). Indeed, individuals who are more characteristically similar and closer in a social network typically have amplified social comparison and competitiveness with each other (e.g., sibling rivalry; Garcia et al., 2013 ). Thus, behaviors driven by status motivation are most effectively employed when relevant to others in a social group, one that the individual identifies with. Recognizing the role of group processes and social influence on competitive efforts and hormonal responses will be important for a comprehensive understanding of the social neuroendocrinology of competition. Status motivation and underlying hormonal correlates should be considered within the context of social groups (e.g., Oxford, Ponzi, & Geary, 2010 ) with reference to factors such as an individual’s level of group identification, status ranking within the group, inter- and intragroup competition, and need for affiliation. Social neuroendocrinologists could consider implementing group social psychology techniques (e.g., Cheng et al., 2010 , 2013 ; Ronay, Greenaway, Anicich, & Galinsky, 2012 ) to explore relationships among group members and group-level interactional characteristics of social hierarchies. Advanced statistical techniques could also prove helpful in implementing more comprehensive social group dynamics (e.g., social network analysis; Kornienko et al., 2014 , 2016 ).

The Functional Significance of Transient Elevations in Testosterone

Another fruitful area of future research involves increasing our understanding of the adaptive purpose of testosterone reactivity in status-relevant contexts. Thus, it is important to expand research on the immediate and subsequent benefits of rapid and transient elevations in testosterone during competitive behavior and as a result of status shifts. As discussed earlier, previous research has shown that testosterone elevations, under certain contexts, increase subsequent willingness to compete again and appear to alter cognitions in ways that could benefit status seeking. Going further, researchers could explore testosterone-related competitive decision making when manipulating aspects about one’s opponent (e.g., photos of potential opponents could be shown with aspects such as gender and dominance-related facial features, and information about that opponent’s skill level and history of success in competition could be manipulated). Another area of interest involves moving beyond just the categorical choice of willingness to compete again or not and devising a quantifiable measure of competitive effort following testosterone elevation (i.e., a subsequent task of competitive persistence). As researchers expand the postcompetition repertoire of status-related behaviors, both antisocial and prosocial means of status maintenance should be considered, as these are opposing yet equally viable strategies (Cheng et al., 2013 ). It is also important to understand the potential benefit of testosterone change, in relation to both physiology and psychology, with the purpose of expanding the relevance, reach, and applicability of social neuroendocrinology more generally. That short-term testosterone elevations can advantageously alter future behaviors and cognitions suggests that there could be some marketable use in developing evidence-based interventions, tasks that could reliably produce endogenous testosterone increases in contexts where this response could prove beneficial for performance. Researchers in other fields, policymakers, and organizations interested in how social groups function (e.g., corporate business and sports teams) would better understand the value of implementing social neuroendocrinology methods with proper tools for taking advantage of the testosterone–status relationship.

Other Areas of Future Research

Other future directions include pursuing a better understanding of hormonal modulation of reward circuitry in the brain (e.g., Bless, McGinnis, Mitchell, Hartwell, & Mitchell, 1997 ; Frye, Rhodes, Rosellini, & Svare, 2002 ; Hermans et al., 2010 ; Montoya, Bos, Terburg, Rosenberger, & van Honk, 2014 ; Packard, Cornell, & Alexander, 1997 ; Salvador & Costa, 2009 ). Although researchers have a growing understanding of how testosterone and other sex steroids relate to behavior, less is known about the neural mechanisms of this relationship. Additionally, social neuroendocrinology, discipline-wide, is challenged with the vital goal of improving assay methodologies to increase the validity of hormone measurement (Granger, Shirtcliff, Booth, Kivlighan, & Schwartz, 2004 ; Granger et al., 2007 ; Welker et al., 2016 ). Perhaps there is no more pressing task than this given that the accuracy of hormone data is the basis of all other assumptions about hormone–behavior rela tionships. Finally, future research should make greater efforts to explore how the research discussed in this chapter is dependent on sex, gender, gender identity, and gender socialization and expand theoretical models to include a more comprehensive understanding of the endocrinology of status motivation and social hierarchies among women (Casto & Prasad, 2017 ).

Competition, dominance, and social hierarchy are aspects of human behavior, past and present, that represent a fundamental human need to attain and maintain social status. The core of these behaviors is driven and can be explained by complex endocrinological processes interacting with the social environment. Understanding these relationships helps explain the causes and consequences of social stratification—why, for example, those born in conditions of poverty face harsher realities and have more barriers to educational achievement, job advancement, and health care. Recognizing how our biology influences the struggle to have and be more than others at the society level, and the individual differences of this drive, is perhaps one vital step in altering perceptions and creating greater awareness for the disparate situations of others. Through the lens of individual and group functioning and performance, there is also great opportunity. By seeking out the best of human potential, hormonal–status relationships can be used to discover interventions beneficial to performance in competitive tasks and functioning within social networks.

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Social Meritocracy and Unjust Social Hierarchies: Three Proposals to Limit Meritocracy’s Erosion of Social Cooperation

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• Elena Ziliotti   ORCID: orcid.org/0000-0002-8929-9728 1  

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A well-functioned society depends on its ability to nurture, attract, and deploy talents in critical sectors. However, the implementation of some meritocratic principles to allocate positions often leads to unjust social hierarchies. Is there, then, a solution to meritocracy’s dysfunctional hierarchical effects? This paper attempts to answer this by drawing on the real-world cases of Singapore and the USA to investigate the relationship of toxic social hierarchies with meritocracy. It proposes three solutions to curb the unjustifiable social stratifications and the erosion of social cooperation often associated with social meritocracy. These reflections could help to shed light on the grounds for the ongoing debates on social hierarchies and provide valuable insights into how to weigh up existing socio-political structures.

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1 Introduction

Just Hierarchy (2020) is a thought-provoking attempt to justify certain social hierarchies by going against the common view in contemporary political philosophy that any social hierarchy must be disallowed. The authors, Daniel Bell and Wang Pei, contend that some forms of social hierarchy are justified in modern societies if they can serve morally desirable goals. Footnote 1 Bell and Wang’s stand has the potential to create new normative spaces: it can generate a novel conceptual distinction between ‘just’ and ‘unjust’ social hierarchies and pave the way for new investigations on the realisation of just social hierarchies in contemporary societies.

I have argued elsewhere against the possibility of ‘just’ social hierarchies (Ziliotti 2022 ). What makes me particularly sceptical about Bell and Wang’s claim that some social hierarchies can serve morally desirable goals is the significant amount of empirical evidence indicating that social hierarchies have detrimental effects, especially on those at the bottom of the hierarchy. Leaving my scepticism on just hierarchies aside, the ongoing debate on social hierarchies in the modern world raises an intriguing question on the forms of social hierarchy that Bell and Wang condemn. Although I am not aware of any scholars defending what Bell and Wang call ‘unjust’ social hierarchies (that is, forms of social hierarchy that serve immorally desirable goals), it remains unclear how these toxic forms of social hierarchies can be checked or, at least, limited.

The issue of toxic social hierarchies has recently gained traction in debates on the consequences of meritocratic selection systems. The meritocratic ideal aspires to oppose unjustifiable social hierarchies by allocating positions based on individual qualities, but several scholars view meritocratic systems as intrinsically hierarchical. For example, Kwame Anthony Appiah argues that “in a meritocratic system, status is achieved in ways that reflect relevant achievements or capacities” ( 2016 : 29). In a similar vein, Michael Sandel claims in his latest book, The Tyranny of Merit , that in contemporary American society “today’s meritocracy has hardened into a hereditary aristocracy” ( 2020 : 32). This is because “the principle of merit can take a tyrannical turn, not only when societies fail to live up to it, but also─indeed especially─when they do” (2020: 42).

In focussing on dysfunctional social hierarchies, Sandel and Appiah raise a problem that should be of interest to both scholars who oppose social hierarchies of any sort and those supporting ‘just’ social hierarchies. How can we limit the effects of dysfunctional unjust social hierarchies on contemporary societies? Furthermore, the issue raised by Sandel and Appiah is not unique to Western societies. Heated debates on meritocracy’s hierarchical effects have gained public attention in other East Asian societies, like the city-state of Singapore. Meritocratic ideologies have shaped Singapore’s culture since 1965, the time of Singapore’s independence from Malaysia. However, according to Singaporean political theorist Benjamin Wong, meritocratic ideology is now synonymous with social stratification and economic inequalities in Singapore (2013).

The undesirable implications of meritocracy pose a difficult conundrum to scholars of social hierarchies and meritocracy. On the one hand, unjust social hierarchies and dysfunctional competing social cultures must be opposed while on the other hand, some meritocratic allocation of positions are inevitable in key sectors of societies. After all, the well-functioning of societies depends on their ability to nurture, attract, and deploy talents in critical sectors. So, what is the solution to meritocracy’s dysfunctional hierarchical effects? Drawing on the debate on equality of opportunity, I suggest reconceptualising the notions of meritocracy as a notion of substantive equality of opportunity to make meritocratic practises fairer. Furthermore, I recommend (a) delinking meritocratic selection in the private sector from neo-liberal principles of reward allocation, (b) coupling meritocratic selection principles in the education sector with substantial diversity standards, (c) reconceptualising the idea of merit accepted in the public sphere.

The next section of the paper elucidates the relationship between meritocratic systems and toxic social hierarchies. Debates on the meritocracy’s hierarchical effects are discussed in the context of contemporary Singapore and American society. Assuming that some forms of meritocracy are instrumental to contemporary societies, Section three advances three proposals to make meritocracy less hierarchical. Finally, Section four summarises the main argument of the paper and suggests future directions for academic debates.

2 Toxic Hierarchies and Social Meritocracy

The concept of ‘social hierarchy’ commonly refers to a social structure in which individuals have different social statuses. According to this definition, social hierarchies entail a distinction of rank or social status, where some persons are morally superior to others or are regarded differently. As Bell and Wang state in their recent book, a hierarchy is a relation that is characterised by (a) difference and (b) ranking according to some attribute. Social hierarchies tend to have a normative dimension: “They are social systems in which there is an implicit or explicit rank of individuals or groups with respect to a valued social dimension” (Bell and Pei 2020 : 8).

A difference in social recognition or social status is a necessary condition for any form of social hierarchy. This claim is also supported by Ricardo Blaug. Drawing from Radner ( 1992 ) and Cloke and Goldsmith ( 2002 ), Blaug argues that “[a]s a way of organising human affairs, hierarchy is a set of arrangements bearing a strong family resemblance; all are structured as a ‘ranked tree’. It signifies a stack of power asymmetries, each featuring differentiated levels of status and degrees of power; layered, as it were, one above the other” ( 2009 : 87). Thus, unequal distribution of decisional power and responsibility is insufficient to establish a social hierarchy, unless accompanied by social stratification and unequal distribution of decisional power and responsibility.

Recently, some scholars have argued that instances of social meritocracy have led to reprehensible social hierarchies in contemporary societies (Markovits 2019 ; Sandel 2020 ). Meritocracy is usually understood as the idea that positions should be allocated according to an individual’s relevant merits (e.g. individual qualities or abilities). Meritocracy maintains that (a) positions and posts should be open to all members of society, (b) applicants are assessed based on their merits, and (c) the posts must be assigned to applicants with relevant superior qualities. Meritocracy is practised in three dimensions: social meritocracy, economic meritocracy, and political meritocracy.

Social meritocracy concerns the distribution of positions in a given societal context. To this end, it is often advocated as a criterion to allocate positions in the private or educational sectors, such as scholarships or seats in prestigious educational institutions. Often, social meritocracy is practised together with a form of economic meritocracy that distributes material resources according to the results of the meritocratic competition for positions.

Unlike social meritocracy, political meritocracy requires distributing political positions, or political power, among members of society according to their political expertise and moral virtues (Bai 2008 , 2013 , 2019 ; Bell 2006 , 2015 ; Chan 2013a , b ; Fan 2013 ; Jiang 2012 ). Footnote 2

While political meritocracy has become a central topic of debate in contemporary Confucian political theory, most Western political philosophers are concerned with social meritocracy and economic meritocracy (Williams 1962 /1997; Rawls 1971 /1999; Sandel 2020 ; Appiah 2016 ). This paper focuses primarily on social meritocracy. Economic meritocracy will be discussed only in relation to social meritocracy. Such a view of meritocracy hinges on the principle of ‘equality of opportunity’, according to which “each must face an array of options that is equivalent to every other person in terms of the prospects for preference satisfaction” (Arneson 1989 : 85). This idea reduces meritocracy to a form of formal equality of opportunity , which is popularly known as ‘career open to talents’.

This idea of “career open to talents” sits well in well-functioning modern complex societies as such societies need to adopt some division of labour; social meritocracy is therefore, often considered an effective and fair way to ensure good outcomes. It aims to oppose unjust social hierarchical structures and ethnic and gender discrimination. While hierarchical social structures (like aristocracy) assign positions based on the implicit or explicit social ranks of individuals or groups that characterise the hierarchical structure, social meritocracy rejects this view by aiming to give individuals equal opportunities to compete for a position regardless of social status, religion, gender, or other aspects that are irrelevant to the job.

Despite its egalitarian and anti-hierarchical aim, social meritocracy ignores the fact that typically members of society have different starting points in the race for a position. In a society characterised by socio-economic inequalities, worse-off families have fewer means than better-off families to help their children develop the skills required to pass the meritocratic selection (Williams 1962 /1997). Thus, if the socio-economic and political circumstances in which individuals live are paramount aspects associated with talents, social meritocracy risks justifying the perpetuation of old discredited hierarchies and social inequalities. Footnote 3

Another problem is that because social meritocracy is often deployed in contemporary societies to distribute positions with significant social and economic advantages, meritocratic structures can also generate new social stratifications and mark out the members of society who have ‘failed’ the meritocratic test. Similar negative consequences are visible in societies like Singapore and the USA, where meritocratic ideologies have shaped the countries’ cultures and served as the main ideological drive for their economic growth. For example, at the time of independence from Malaysia, Singapore’s adoption of meritocracy as an ideology and policy was an obvious choice given the multicultural composition of the society and its geopolitical conditions. As Singaporean Sociologist Chua Beng Huat explains, Singapore needed “to demonstrate the absence of racial discrimination in the Malay dominant region, in contrast to Malaysia’s insistence on policy supremacy and privileging of Malays as its indigenous people” (Chua 2017 : 8). With a majority of Chinese-origin people and significant Indian, Eurasian and Malaysian minorities, meritocracy appeared to be the only ‘neutral’ ideology that could unite the people of that young country. Notably, in modern Singapore, meritocratic principles were implemented to reorganise the civil service. Under British rule, the civil service apparatus was quite dysfunctional and accessible only to the British (Ho et al. 2017 : 93). In the 1960s, the government extended the recruitment to all Singaporeans of any race and religion.

Yet, meritocracy has become a dirty word in modern Singapore, and many Singaporeans now associate it with systemic economic inequalities and social injustice (Ong 2018 ). According to Wong, “[s]ince 1994, it has been used as a means to justify the market-based salaries of ministers and top civil servants, who are the highest paid in the world” ( 2013 : 289). For example, after the 2011 salary review, the annual pay of entry-level ministers in Singapore is S$1.1 million (around US$850.000), whereas that of the Singaporean prime minister is S$2.2 million (approximately US$1.7 million). This, in turn, has contributed to creating a perception of a “great affective divide” between the political elite and the people (Wong 2013 : 288).

In a similar vein, American philosopher Michael Sandel claims that “today’s meritocracy has hardened into a hereditary aristocracy” in contemporary American society (2020: 32). Drawing from Max Weber’s The Protestant Ethic and the Spirit of Capitalism , Sandel maintains that the Protestant work ethics gave rise to the meritocratic ideology that characterises contemporary American society. Based on the Calvinist notion of predestination, protestant ethics began considering worldly success as “a good indicator of who is destined for salvation” (Sandel 2020 : 49). Subsequently, “[w]orking and striving became imperatives of their own, detached from Calvinist notions of predestination and the anxious search for a sign of salvation (Sandel 2020 : 51). According to Sandel, these beliefs have shaped the mindset of many contemporary Americans:

Today’s secular meritocratic order moralises success in ways that echo an earlier providential faith: Although the successful do not owe their power and wealth to divine intervention—they rise thanks to their own effort and hard work—their success reflects their superior virtue (Sandel 2020 : 52).

For Sandel, this meritocratic logic has eroded American society’s social cohesion and social capital. The rich believe that they ‘are rich because they are more deserving than the poor’, while the poor resent ‘the winners’ (Sandel 2020 : 52). These common attitudes have influenced recent events in American politics. Commenting on Trump’s victory of the 2016 US elections, Sandel claims that “the populist backlash was provoked, at least in part, by the galling sense that those who stood astride the hierarchy of merit looked down with disdain on those they considered less accomplished than themselves. […] They embraced meritocracy, but believed it described the way things already worked. They did not see it as an unfinished project requiring further government action to dismantle barriers to achievement” (Sandel 2020 : 85–86).

Wong’s and Sandel’s analyses indicate that social meritocracy can create social stratifications with pernicious effects. On the one hand, they foster frustration and resentment in those at the bottom of the hierarchy while on the other hand, they feed the sense of entitlement in those at the top of the social hierarchy, preventing them from developing compassion for fellow citizens. These effects are intolerable even for supporters of just hierarchies because the latter oppose social hierarchies that are detrimental to the moral and material well-being of society’s members.

The negative implications of meritocracy pose a difficult conundrum. On the one hand, social hierarchies and a dysfunctional competing social culture must be opposed, and on the other hand, certain meritocratic allocation of positions seems to be inevitable in crucial sectors of societies. After all, there are compelling reasons why hospitals should hire their staff primarily based on their merits and expertise; airlines choose the most experienced pilots, and governments dealing with the covid-19 pandemic must seek the advice of the most competent virologists. So, what can be done? The following section explores a prominent solution to this problem.

3 Substantive Equality of Opportunity: Possibilities and Limits

One workable solution to the harmful implications of meritocracy proposes a reconceptualisation of meritocracy to make it more egalitarian. According to this view, the meaning of meritocracy must be revised to avoid some of the paradoxical consequences of meritocratic practises. For instance, if socio-economic inequalities are part of many societies and they create unfair chances in meritocratic selections, then, meritocracy should be understood as a form of substantive equality of opportunity , not formal equality of opportunity. Footnote 4 According to the principle of substantive equality of opportunity , achieving equality of opportunity requires a meritocratic selection system whereby a genuine chance to become qualified is presented to all members of society. In the egalitarian literature, this concept is known as equality of access (Mason 2001 : 762). Following John Rawls ( 1971 /1999), one could argue that equality of access is achieved only when members of society with the same native talents and the same ambition have equal prospects of success in relation to competitions for positions (Rawls 1971 /1999: 63). Thus, to the extent that meritocracy concerns fairness, equality of access must be considered a constitutive part of meritocracy.

Rawls’ conception of equality of access, or ‘fair equality of opportunity’, aims to correct formal equality of opportunity by requiring equality of endowment , that is, equal chances to the equally well-endowed members of societies regardless of parents’ incomes and social contingencies (Freeman 2007 : 98). It does so by requiring society to develop structural conditions to prevent inequalities from becoming significant social inequalities. For example, “[f]ree market arrangements must be set within a framework of political and legal institutions which regulates the overall trends of economic events and preserves the social conditions necessary for fair equality of opportunity. […] So, the school system, whether public or private, should be designed to even out class barrier” (Rawls 1971 /1999: 63).

As indicated in the last part of the above quote, achieving equality of access would require substantial restructuring of the educational system. Education is a paramount means through which an individual can acquire the skills and knowledge necessary to compete in the meritocratic allocation of several social positions. This would require allowing as many students as possible equal opportunities for a competitive education. As Rawls suggests, in the American context, this can be achieved by narrowing the gap between the job opportunities of graduates from private and public schools.

Several critics have objected that equality of access requires significant intervention by the government in the citizens’ private life. According to Houlgate ( 1980 ), Goldstein et al. ( 1986 ) and Fishkin ( 1983 ), if Rawls’ idea of equality of access were implemented in education, it would violate the liberty of individual parents to influence considerably the development of their children’s life, as “parents who failed to ensure such prerequisites for their children could justifiably be subject to state interference” (Fishkin 1983 : 36). Therefore, according to Fishkin, Rawlsian equality of access does not only indicate how wealth should be redistributed among the members of society, but it can also constrain some parents to spend their money and time on their children in a specific way.

Fishkin’s claim regarding the tension between liberty and equality is unfounded. Equality of access may impact the liberty of some parents to a certain extent. However, critiques have argued that Fishkin assumes without proving that, from a liberal perspective, parents have the right to liberty in general. In other words, it remains to be proved that they must have the full freedom to substantially affect their children’s life as they see appropriate (Boxill 1984 : 618).

Another objection to fair equality of opportunity comes from Brighouse and Swift ( 2008 ). They contest the radical application of fair equality of opportunity to all aspects of parent–child relationships. According to them, fair equality of opportunity should not and cannot apply to other parental behaviours, such as reading bedtime stories to children that are essential to achieving the particular values of parent–child relations that are made for the sake of the child (2008: 54). But, concerning the other kinds of choice, such as the parents’ choice of sending their children to private schools, Brighouse and Swift argue that the state can try to discourage them by using taxes. In response to this objection, Segall ( 2011 ) maintains that luck egalitarianism would indeed requires neutralising parental partiality (such as reading bedtime stories to their child) if this gives these children an undue advantage later in life. However, this is a pro tanto reason that, in practise, can be overridden by other considerations. For instance, because the parents’ activity of reading bed time stories to their children can contribute to the cultivation of good family relations in a unique way, the luck egalitarian parent has strong reason to engage in this practise, even though they are fully aware and regret that it could lead to unfair socio-economic advantages (Segall 2011 : 29–20).

Sandel formulates one of the most compelling objections to the proposal of reconceptualising meritocracy as a form of substantive equality of opportunity. According to Sandel, revisiting the concept of equal opportunity is a non-starter to overcoming meritocracy’s social stratification and erosion of commonality. For Sandel, one of the main problems of meritocratic practises is that those at the top feel entitled to their economic remuneration and status in virtue of their sacrifices, self-discipline, and so on. This leads them to look down on the rest and justify their worst-off positions. The problem with substantive equality of opportunity is that it may even increase this sense of entitlement: “[i]f opportunity are truly equal, then not only will people rise as far as their talents and hard work will take them; their success will be their own doing, and they will deserve the rewards that come their way” (Sandel 2020 : 82).

Sandel is correct: substantive equal opportunity makes meritocracy fairer but does not diminish its hierarchical effects and erosion of commonality. However, this does not mean that the idea of social meritocracy has no room for further discussions. It simply suggests that substantive equality of opportunity is necessary but insufficient to reform social meritocracy. Other solutions can be formulated to solve the above issue. In the next section, I will propose three solutions to mitigate meritocracy’s detrimental hierarchical effects.

4 Three Proposals to Make Social Meritocratic Practises Less Hierarchical

This section proposes three integrated solutions to contain the hierarchical effects of social meritocratic practises. I contend that it is crucial to (1) delink meritocratic selections from market-based compensation structures, (2) integrate meritocratic selection principles into a pluralistic-value selection system that promotes diversity, and (3) decouple ‘merit’ from the concept of ‘desert’.

4.1 Delinking Meritocracy from Market-based Compensation Structures

To prevent meritocratic practises from contributing to the establishment of toxic social hierarchies, it is paramount to delink meritocracy from market-based compensation structures. Wong’s discussion on meritocracy in Singapore and Sandel’s analysis of meritocracy in the American context indicate that most contemporary debates focus not on the meritocratic ideal alone but on a neo-liberal version of meritocracy in which meritocratic selections parallel market-based compensation structures. The latter further intensifies meritocracy’s social erosion because the creation of a fierce competition for the so-called ‘top jobs’ is often due to the social and economic advantages attached to these positions, not these positions in themselves. To understand this point, it is helpful to consider recent debates on the salaries of civil servants as well as politicians in Singapore.

Many Singaporeans do not question the need to select civil servants and politicians based on their specific merits or expertise. The need for skilled and capable people in charge of policymaking is not a politically sensitive issue in Singapore. For many Singaporeans, the point of contention is the need to pay civil servants and politicians an extraordinary amount of money for their service. Singapore’s government has responded to this criticism by pointing out that market-based pays are required to attract highflyers that otherwise would go into the private sector. In fact, the city-state of Singapore is a global business hub and many international companies have their regional headquarters in Singapore. However, many Singaporeans are unpersuaded by this argument because the paramount quality that civil servants and political leaders ought to have is a strong sense of civic duty, not greed (Wong 2013 : 296).

I believe a similar problem is rooted in the American-style meritocratic job-market systems. From 2002 to 2007, the top 1% secured two-thirds of all gains from the growth in the USA economy. In particular, the “tech sector has created many wealthy entrepreneurs and investors” and “IT companies tend to pay their CEO more” (Brynjolfsson et al. 2015 : 10). One of the reasons for this trend is that new technologies have amplified the effects of the CEO’s decisions. For example, making copies of digital goods has almost zero costs, so who gets the right idea first on how to innovate the market is likely to acquire most of that market (Brynjolfsson et al. 2015 : 9). This has created a new group of extremely wealthy persons because tech companies compete with each other to hire the best people (Brynjolfsson et al. 2015 : 9). As in the Singapore case, selecting CEOs’ based on their creativity and vision is not the leading cause of rising economic inequalities. What is problematic is their excessive compensations, especially if these are compared to the economic situation of the American low-skilled information workers who are losing their jobs due to the increasing automation of the work sector.

In other words, I agree with Sandel that “[m]orally, it is unclear why the talented deserve the outsize rewards that market-driven societies lavish on the successful” ( 2020 : 32). However, Sandel is mistaken in thinking that meritocracy also entails a specific principle of market-based reward allocation. While the two have often gone together in practise, social meritocratic principles can be decoupled from neo-liberalism by putting caps on salaries in the key sectors.

Decoupling meritocracy from neo-liberalism could lead to a type of social meritocracy that is less objectionable. Reconsidering the economic rewards attached to prestigious positions may contribute to reducing the social pressure for competing for these positions, since winning the race will create less economic, and therefore also less social, empowerment. One limit of this proposal is that high performers can move to other tax jurisdictions if such caps are viewed as too heavy. Thus, the success of this proposal depends on the possibility of multiple tax jurisdictions to applying the same policy and also to calibrate the caps sensibly. Footnote 5

Besides putting caps on salaries in key sectors, ensuring decent minimum wages may help control the winners’ social power while fostering the dignity of the lower income groups. This can contribute to keeping the effects of meritocratic practises in check, but it requires a reckoning with neo-liberalist reform policies which are at the heart of both Singapore and American economic systems.

This is not to say that Singapore’s use of neo-liberal reforms is similar to the one in the USA. On the contrary, the Singapore government practises what many call ‘State capitalism’ in which the government has the ultimate say on what sector of the economy should be liberalised (Chua 2017 ). However, if multiple reasons are behind the hierarchical effects of meritocracy in contemporary, establishing whether the problems lie in the meritocratic principle itself or its application in a specific socio-economic context can help formulate successful solutions.

4.2 Redefining the Justiciability of Meritocratic Practises

Besides delinking meritocratic selection in the private sector from neo-liberal principles of rewards allocation, it is paramount to clarify where and to what extent meritocratic practises are necessary. As I said, all well-functioning modern societies must nurture, attract, and deploy talents in critical sectors to ensure efficiency and fulfilment of societal needs. However, this does not mean that individual merits are the only criteria that matter for a well-formed society; other criteria should be considered in the selection mechanisms. This is crucial because opening the selection system to standards other than merit may lessen meritocracy’s hierarchical effects.

Take, for example, the staff hiring mechanisms in the educational sector. Efficiency and competence are not the only selection standards that must matter because these selection systems aim to create communities of inquiry, where individuals can develop meaningful social relationships and mental and moral growth. For instance, gender balance and ethnic diversity are increasingly regarded as essential criteria for developing conducive educational communities and in the hiring of academic staff (Fradella 2018 ; Irby and Brown 2022 : 45). Therefore, more selection criteria must be considered.

However, diversity remains undervalued in the educational sector. Globally, the percentage of women in academia has increased from 39.2% in 2001 to 43.1% in 2019 (Calderon 2022 ), but this trend is not uniform. For example, the proportion of women academics in regions like Sub-Saharan Africa accounts for just 24.3% of academics (Calderon 2022 ). Female Professors in UK Universities continue to remain underrepresented, as male professors outnumber females by three to one (Adams 2020 ). Furthermore, elite Universities remain “white spaces” both in the UK and the USA (Bohpal 2022 ). Black academics accounted for only 2% of all academic staff in the UK, with only 25 black women professors across all UK Universities (Adams 2020 ). In the USA, black academics are approximately 5.7% of all full-time faculty members at Colleges and Universities, while white academics make up 73.2% of the total (American Council of Education 2016 ).

It is open to question whether this data reflects racial and gender bias or whether many Universities are ‘overusing’ meritocratic selection principles. However, while qualifications and competence count, integrating academic merits in a pluralistic-value system, where more standards guide the selection of staff members can help limit meritocracy’s hierarchical effects. First, the inclusion of diversity principles in forming communities erodes the sense of entitlement in those at the top of the social hierarchy. As the hiring decisions do not depend exclusively on the candidate’s talents and skills, it would be unreasonable for the latter to believe that “their success will be their own doing” (Sandel 2020 : 82). Second, because affirmative action breaks social barriers and increases the opportunity of members of marginalised groups, it can diminish the frustrations and the resentment of the latter.

Some meritocrats would worry about the potential threat this affirmative action poses to the quality of staffing in the private sector. After all, the argument goes, private companies are not ‘communities of inquiry’ like educational institutions; their overall goal is profit. Thus, there are good reasons to believe meritocracy trumps diversity in the private sector. In response to this objection, it should be observed that studies have shown that a selection system in the private sector can yield more positive results if it is open to individuals with different skill sets and backgrounds. For example, the presence of qualified women on a company board can bring significant value because women tend to have different leadership skills than men. Female leaders tend to be more transformational (Eagly et al. 2003 ) and more attentive than men to the ‘human side’ of enterprise, meaning that female leaders tend to base judgments more on intuition and emotions than on rational assessments of the relationships between means and ends (Bass and Avolio 1994 ). Other studies found that women’s decision-making styles tend to be more participatory than those typically adopted by men (e.g. Mertz and McNeely 1997 : 8); female leaders are usually more collaborative than male leaders, and they also engage in more contingent reward behaviours (Eagly et al. 2003 ). This suggests that diversity can increase efficiency even in the private sector.

4.3 Decoupling the Notion of ‘Merit’ from ‘Desert’

This brings me to my third and final point to prevent meritocratic practises from contributing to establishing toxic social hierarchies. Another way to lessen the entitlement of those who won the meritocratic race is to re-evaluate the concept of ‘merit’. Part of the problem of meritocratic practises concerns a certain ambiguity in the everyday use of the concept of ‘merit’. Since ‘merit’ and ‘desert’ are often used interchangeably in public speeches, some people may think that winning the meritocratic race for a particular position also entails deserving its economic condition and social status.

However, different linguistic uses suggest the possibility of partly decoupling the concept of ‘merit’ from ‘desert’. For instance, we say that a naturally talented singer has the merit of singing well, but they do not deserve their ability since they did nothing to have it. After all, they were born with her talent. Similarly, a gifted financial mathematician may merit her job as a top executive at J.P. Morgan because they have all the necessary qualities to do an excellent job. However, it is possible that one of the candidates who did not get that job deserved it more because this person had always struggled with financial mathematics and spent double the number of hours as the gifted financial mathematician to build up their knowledge in the subject and prepared for the interview.

In both cases discussed above, an element of desert is still likely to be needed because natural talent is insufficient for success. As in the case of the singer, usually it can take several years of practise and dedication. Footnote 6 However, my point is that these linguistic uses indicate that ‘merit’ does not always correspond to ‘desert’ in English. Merit can refer to the qualities (some may even say the ‘virtues’) that an individual possesses. In contrast, desert concerns the deeds an individual has performed to obtain a particular position or title. Therefore, according to this linguistic usage, a person can merit something without deserving it. The distinction between merit and desert entails that in a meritocratic society, it is false that winners “have earned their success through their talent and hard work” (Sandel 2020 : 21) because (as we have seen in the case of mathematicians at J.P. Morgan) meritocratic selection principles track merit, not effort. In other words, in a meritocratic system, individuals are hired or promoted based on how well their profiles match the required qualifications. Because we are born with different natural inclinations, we must dedicate different efforts to cultivating the same talents even under substantive equality of opportunity.

If this distinction between merit and desert was emphasised and reiterated in a meritocratic society, it could bring a new perspective on what individuals owe to each other and help limit meritocracy’s hierarchical effects. Decoupling merit from desert in public speeches can help the winners of the meritocratic lottery be more aware of the impact of luck in their lives instead of cultivating a sense of entitlement. They may merit their jobs, but it is questionable how much they deserve them. Thus, reviewing the meanings of meritocracy and merit is not just philosophically relevant; it can help limit the sense of entitlement of those who won the meritocratic race.

5 Conclusion

The meritocratic ideal has emerged in opposition to social hierarchy. It reflects the aspiration to create a functional society that efficiently uses the talents and skills of its members. However, meritocracy can generate unjustifiable social stratifications and erode social cooperation. This poses pressing questions for contemporary societies: to what extent should meritocracy be blamed for its adverse effects? Can modern societies do without meritocracy, and if so, how?

This paper has investigated the relationship of toxic social hierarchies with forms of social meritocracy. I have proposed a multipronged approach to curb the unjustifiable social stratifications and the erosion of social cooperation often associated with meritocracy. In addition to reconceptualising meritocracy as a form of substantive equality of opportunity, I have proposed (a) delinking meritocratic selection in the private sector from neo-liberal principles of reward allocation, (b) coupling meritocratic selection principles in the education sector with diversity standards, (c) reconceptualising the concept of merit. While these proposals are not a fixed set of policy proposals, they can be a starting point for a conversation on how we can make meritocratic practises result in less toxic hierarchical relationships while promoting the socio-economic conditions required for social cooperation.

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Code availability

Bell and Wang’s view is in sharp contrast with Kolodny ( 2023 ), which contests the justifiability of any social hierarchy.

Since these two forms of meritocracy apply to different position distributions, it is possible to advocate social meritocracy while rejecting political meritocracy. For instance, by defending a distribution of political power that follows democratic principles.

Here, I am drawing from my analysis of the debate on equality of opportunity in Western literature in Ziliotti ( 2017 ).

I have advanced this proposal in  Ziliotti ( 2017 ).

An alternative solution would be to impose high taxes on high earners, not capping salaries (I am grateful to an anonymous referee for suggestion this point). But this proposal has similar limitations: high performers can move to other tax jurisdictions if such taxes are viewed as burdensome.

I am grateful for an anonymous review for helping me see this point.

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Funding was provided by the Research programme ‘Ethics of Socially Disruptive Technologies’, the Gravitation programme of the Dutch Ministry of Education, Culture, and Science and the Netherlands Organization for Scientific Research, (Grant No. [024.004.031]).

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Ziliotti, E. Social Meritocracy and Unjust Social Hierarchies: Three Proposals to Limit Meritocracy’s Erosion of Social Cooperation. Fudan J. Hum. Soc. Sci. (2024). https://doi.org/10.1007/s40647-024-00400-9

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The establishment and maintenance of dominance hierarchies

Elizabeth a. tibbetts.

Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, USA

Juanita Pardo-Sanchez

Chloe weise, associated data.

This article has no additional data.

Animal groups are often organized hierarchically, with dominant individuals gaining priority access to resources and reproduction over subordinate individuals. Initial dominance hierarchy formation may be influenced by multiple interacting factors, including an animal's individual attributes, conventions and self-organizing social dynamics. After establishment, hierarchies are typically maintained over the long-term because individuals save time, energy and reduce the risk of injury by recognizing and abiding by established dominance relationships. A separate set of behaviours are used to maintain dominance relationships within groups, including behaviours that stabilize ranks (punishment, threats, behavioural asymmetry), as well as signals that provide information about dominance rank (individual identity signals, signals of dominance). In this review, we describe the behaviours used to establish and maintain dominance hierarchies across different taxa and types of societies. We also review opportunities for future research including: testing how self-organizing behavioural dynamics interact with other factors to mediate dominance hierarchy formation, measuring the long-term stability of social hierarchies and the factors that disrupt hierarchy stability, incorporating phenotypic plasticity into our understanding of the behavioural dynamics of hierarchies and considering how cognition coevolves with the behaviours used to establish and maintain hierarchies.

This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.

1.  Introduction

Many animal social interactions are organized hierarchically based on dominance rank. Dominance is typically defined as asymmetry in aggression by one animal towards another animal [ 1 , 2 ]. However, the term dominance is used in different ways across taxa and contexts. For example, in social insects, dominant individuals monopolize reproduction within their group, but may not be involved in aggressive competition [ 3 , 4 ]. More typically, dominance within groups is defined as a long-lasting position associated with asymmetric aggression and priority access to physical or social commodities that increase fitness, including food, water, shelter, space, receptive mates, alloparental care, etc [ 5 ]. In other cases, dominance refers to individuals that win short-term dyadic contests [ 1 ]. The winner of dyadic contests gains priority access to resources or reproduction, but there may be no long-term hierarchical relationship between the two competitive individuals. In groups, there is variation in the structure of dominance hierarchies [ 6 ]. Many social groups have linear or near-linear dominance hierarchies. However, hierarchies need not always be linear [ 2 , 7 ]. Nonlinear hierarchies with dominance reversals and intransitivities can also occur [ 8 ].

Much research on dominance relationships focuses on the initial establishment of hierarchies. Initial rank is based, in large part, on intrinsic individual attributes like resource holding potential (RHP) or motivation [ 9 – 11 ]. Individual attributes are assessed via direct agonistic interactions, signals of fighting ability and social information acquired by watching the interactions of others [ 12 , 13 ]. Other factors such as conventions and social dynamics also play a role in hierarchy formation [ 14 , 15 ]. The relative importance of individual attributes, conventions and social dynamics in the establishment of hierarchies vary across taxa and social contexts.

In large or unstable groups with repeated, low-stakes competitive interactions, individuals repeatedly establish dominance relationships without necessarily forming persistent hierarchies. However, in taxa that live in smaller groups with consistent membership, hierarchies are typically maintained over the long term. Continued conflict over rank is costly (e.g. time, energy, risk of injury) [ 16 , 17 ]. As a result, stable hierarchies are favoured. Hierarchies are maintained via social dynamics like punishment and threats [ 18 , 19 ] as well as signals that provide information about rank, including signals of individual identity and dominance [ 20 , 21 ].

In this review, we describe the behaviours used to establish and maintain dominance hierarchies ( figure 1 ). Some behaviours are involved in both establishing and maintaining hierarchies. For example, differences in fighting ability influence initial rank formation and longer term rank maintenance [ 9 ]. Other behaviours are only involved in a single context. For example, signals of fighting ability influence dominance rank establishment, but are not involved in maintaining hierarchies within groups of known individuals [ 28 – 30 ]. We also describe how factors like group size, consistency of group membership, the costs and benefits of hierarchy position and individual cognitive capacity influence the behaviours involved in hierarchy formation and maintenance. Finally, we highlight several approaches and opportunities for future research on dominance hierarchies.

Mechanisms involved in establishing and maintaining dominance hierarchies. Establishing dominance . Individual attributes are traits such as RHP and/or motivation, assessed via aggressive interactions, signals of fighting ability and social information (e.g. claw waving displays provide information about fighting ability during contests between rival crabs [ 22 ]). Conventions are unique attributes that single out an individual as the next dominant without reflecting intrinsic characteristics that allow individuals to win contests (e.g. maternal rank inheritance in baboons [ 23 ]). Self-organizing social dynamics are social processes at the group level that increase hierarchy linearity (e.g. fishes that interact in groups are more likely to form transitive dominance relationships than fishes that only interact in dyadic contests [ 14 ]). Maintaining dominance . Signals of dominance provide information about dominance rank (e.g. dominant ant queens have cuticular hydrocarbons that provide information about rank and influence queen/worker interactions [ 24 ]). Signals of individual identity are unique phenotypes that receivers learn and associate with individual-specific information about the sender like dominance rank. (e.g. Polistes fuscatus wasps learn the dominance rank of individuals via aggressive contests and social eavesdropping and associate rank information with the unique facial patterns of conspecifics [ 25 , 26 ]). Behavioural mechanisms help dominants maintain their rank like threats, punishment and self-reinforcing behavioural mechanisms (e.g. in meerkats, dominants aggressively evict subordinates that try to reproduce [ 27 ]. (Online version in colour.)

2.  Establishing dominance relationships

(a) individual attributes.

The individual attributes of interacting individuals have a strong and consistent effect on dominance rank ( figure 2 ). The most straightforward way to establish dominance is through competitive dyadic interactions with rivals. Individuals with greater RHP and/or motivation are more likely to win fights and become dominant than individuals with lower RHP and/or motivation [ 9 , 33 ]. RHP, or fighting capacity, is based on a composite of many morphological, physiological and behavioural traits (e.g. weapons, body size, skill, hormones, fat reserves, etc.) [ 34 ]. As a result, many RHP-linked attributes are associated with dominance rank, including body size [ 35 , 36 ], age [ 35 ], personality [ 37 ], hormone titres [ 38 ] and physical condition [ 39 ]. Motivation in contests is influenced by individual state, context and the pay-off individuals receive for winning a contest. Highly motivated individuals will invest more in attaining high dominance rank than less motivated individuals. For example, a hungry individual will be more motivated and fight harder than a satiated individual during competition over food. An individual defending a nest with offspring will be more motivated than an individual defending a potential nesting site. Establishing dominance ranks through direct contests is costly in terms of time, energy and potential physical damage, or death [ 40 , 41 ]. Nevertheless, dyadic contests ultimately guide the establishment of social dominance hierarchies in many species.

Three common ways animals assess the individual attributes of conspecifics. ( a ) Signals of fighting ability. Platysaurus broadleyi signal fighting ability with UV throat coloration [ 31 ]. ( b ) Competitive interactions. Panthera leo fight with rivals. ( c ) Social information. Xiphophorus helleri gain information about the fighting ability of potential rivals by observing and remembering how conspecifics behave during contests with other individuals [ 32 ]. (Online version in colour.)

The outcome of contests over dominance rank may be influenced by an individual's own RHP or the difference in RHP between rivals [ 42 , 43 ]. Some animals persist in contests based on their own abilities such that individuals with low RHP give up before individuals with high RHP (self-assessment). Other animals compare their own abilities with the abilities of their rival (mutual assessment). Self-assessment is less accurate than mutual-assessment but is also simpler [ 42 , 43 ]. Mutual assessment may require substantial time, energy and cognitive resources to process the cues and signals that provide information about an opponent's fighting ability [ 43 ]. A large body of literature has investigated the mechanisms used to determine contest outcomes, finding that self-assessment is more common than mutual assessment [ 43 – 45 ], but even animals thought to have limited cognitive capacity are capable of mutual assessment [ 45 , 46 ].

Some animals minimize the cost of conflict by using signals of fighting ability to directly assess rivals. Signals of fighting ability provide information about RHP and/or motivation during a contest and the signals alter receiver behaviour ( figure 3 ) [ 28 , 47 ]. Receivers are more likely to avoid or submit to rivals that signal high fighting ability than individuals that signal low fighting ability. Signals of fighting ability are widespread across taxa and sensory modalities. For example, fighting ability is signalled by call frequency in frogs, black facial spots in paper wasps and claw size in crabs [ 46 , 48 , 49 ]. Some signals are inherently linked with fighting ability. For example, weapons like antlers or claws signal high fighting ability and weapons also directly influence an individual's ability to win a fight. As a result large weapons always signal high fighting ability. Other signals of fighting ability are termed ‘conventional signals' because they provide information about fighting ability but there is no logical, a priori , link between the signal phenotype and the information the signal conveys [ 50 ]. For example, soft song is a signal of aggressive intent in some songbirds, though there is no required link between aggression and singing quietly [ 51 ]. Some taxa even have multiple different traits that are used as conventional status signals in different contexts [ 52 ]. Although there has been some debate about the accuracy of conventional signals, there is growing evidence that conventional signals provide reliable information about fighting ability and/or aggressive motivation and often mediate conflict over rank [ 28 ].

Signals involved in dominance hierarchy establishment and maintenance. (Online version in colour.)

The individual attributes of rivals can also be assessed indirectly using social information. For example, during social eavesdropping, bystanders gain information about the agonistic ability of particular rivals by observing and remembering how conspecifics behave during contests with other individuals [ 53 , 54 ]. For example, contest behaviour of Polistes fuscatus wasps is influenced by observing fights between conspecifics. Wasps are less aggressive towards individuals they observed initiate more aggression and receive less aggression. Control trials illustrate that the link between bystander behaviour and observed aggression is caused by social eavesdropping rather than alternative behaviours like winner/loser effects or cueing on physical traits (e.g. size) [ 25 ]. Social eavesdropping occurs in many birds, fishes and primates. Individuals often change their contest behaviour based on information about fighting ability obtained by watching or listening to interactions between others [ 32 , 55 , 56 ].

Social eavesdropping may be facilitated by ‘victory displays’, stereotyped behaviours performed by winners following fights that advertise the outcome of a contest to observers [ 57 , 58 ]. For example, little blue penguins give victory calls after they win a fight. Experimental playback of victory calls suggests that advertising victories may help males develop a reputation for winning fights within the social group, potentially reducing the likelihood of being challenged by eavesdroppers in future contests [ 59 ]. Victory displays are widespread, though less research has tested the functional consequences of these displays. Additional research will be useful to assess whether victory displays commonly play a role in social eavesdropping.

Some taxa combine social information and transitive inference to make inferences about the likely fighting ability of individuals without directly observing all potential fights. For example, if A dominates B, and B dominates C, then you can make the inference that A will probably dominate C, even if A and C have never been observed interacting. Transitive inference was originally thought to be based on logical deduction and confined to taxa with ‘advanced’ cognitive abilities like primates [ 60 ]. More recent work has shown that transitive inference occurs in many social species, including primates, birds, fishes and paper wasps [ 60 – 62 ]. Transitive inference may be favoured in social species with linear dominance hierarchies because it allows animals to keep track of dominance relationships while minimizing direct conflict. Further, hierarchies may form much more quickly when animals use transitive inference and social observation to assess rival ranks than when ranks are determined via direct aggressive competition [ 63 ]. Consistent with transitive inference providing a benefit during hierarchy formation, social complexity is linked with the capacity for transitive inference [ 64 , 65 ]. For example, highly social pinyon jays have more accurate transitive inference abilities than less social western scrub-jays [ 64 ].

Although many taxa use social eavesdropping and transitive inference to minimize conflict over dominance, there are cognitive costs associated with social information use. Social eavesdropping involves learning and remembering unique individuals, observing interactions, making appropriate deductions and remembering those deductions [ 53 , 54 ]. Social eavesdropping also requires animals to adopt a non-egocentric perspective and assess interactions they do not directly participate in. Keeping track of a broad network of social interactions has potential to dramatically increase the cognitive challenges of social life compared with only keeping track of personal interactions [ 66 ]. As a result, social information use influences cognitive evolution. Species that keep track of many individually differentiated social relationships often have larger brains and/or enhanced cognitive capacity compared with other species [ 67 – 70 ]. Within-species variation in cognitive performance may also be linked with social knowledge. For example, Australian magpies that live in larger social groups performed better on multiple intelligence tests than birds from smaller groups, suggesting that keeping track of others' social relationships may influence general intelligence [ 71 ].

Thus far, it is unclear how cognitive costs limit which taxa use social information. Social information use is common in taxa with ‘advanced’ cognitive capacity like primates, but taxa with relatively small brains like paper wasps and fishes are also capable of social eavesdropping and transitive inference [ 25 , 54 , 60 – 62 ]. Therefore, brain size does not strictly limit social information use. There are exciting opportunities for future research exploring the relationships between dominance, social information use and cognitive evolution across taxa and social contexts [ 72 ].

(b) Conventions

Dominance rank can also be acquired through conventions. In societies with convention-based dominance, individuals have unique attributes that single them out as the next dominant, (e.g. age, tenure in a group or maternal rank) without reflecting intrinsic characteristics that allow individuals to win contests [ 73 , 74 ]. For example, some social insects determine dominance based on the seniority convention. The oldest worker is the most dominant and will take over if the queen disappears [ 15 , 75 ]. ‘Nepotistic hierarchies' are common dominance conventions where dominance rank is inherited from the mother. Juveniles acquire status immediately below their mother, with younger offspring outranking older siblings [ 76 ]. Although nepotistic hierarchies are considered convention-based, rank inheritance depends on support from mother, kin and coalition members to ensure offspring acquire the appropriate rank [ 77 , 78 ].

There has been a long-standing interest in the evolutionary stability of dominance conventions. If high dominance rank is beneficial and rank is not based on intrinsic attributes, what prevents individuals with high fighting ability from ignoring the convention and asserting their dominance? One explanation is that conventions are more likely to occur when the costs of competition over dominance rank outweigh the benefits of high dominance rank [ 79 ]. For example, conventions are particularly common in taxa that live in long-term social groups with many relatives because there are both direct and indirect fitness costs to group conflict (e.g. primates, social insects) [ 75 , 78 ]. Conventions may also be more likely to mediate dominance when there are weaker benefits of high dominance rank. For example, meerkats aggressively compete over rank rather than relying on conventions. Reproductive success is strongly influenced by rank because dominant meerkats kill the offspring of subordinates [ 80 ]. By contrast, savannah baboons use convention-based hierarchies and individuals of all ranks reproduce [ 81 , 82 ]. In chimpanzees, females largely use a seniority convention for social status in the female hierarchy, while males aggressively compete for status in the male hierarchy. The sex difference in behaviours chimpanzees use for hierarchy formation may occur because male reproductive success is strongly influenced by rank, while female social status is more strongly influenced by longevity than rank [ 83 ].

In some societies with convention-based hierarchies, aggressive competition also influences hierarchy position [ 78 ]. For example, rank in hyenas is inherited from the mother, but maintaining a particular rank requires support from kin and other coalition members. Without sufficient coalitionary support, dominance reversals can occur [ 77 ]. Therefore, rank may often be influenced by a combination of conventions and competition rather than purely conventions.

(c) Self-organizing social dynamics

Recent research suggests that self-organizing social dynamics play a role in establishing linear dominance hierarchies. In a dominance hierarchy, individuals are arrayed in a line from most to least dominant; individuals are dominant to those below them in the hierarchy and subordinate to those above them in the hierarchy. In most social groups, dominance hierarchies are more linear than expected by chance [ 84 ]. Some of this linearity is owing to differences in individual attributes, as described above. However, a growing body of literature suggests that at least some of the linearity in dominance hierarchies occurs because of social processes that go beyond dyadic contests [ 8 ]. For example Chase et al. [ 14 ], experimentally demonstrated that social interactions between group members facilitate the formation of highly linear hierarchies in Metriaclima zebra fish. When fish interact in groups, nearly all groups formed linear dominance hierarchies. When fish interact in dyadic, round-robin competition alone, without any group interactions, only half the hierarchies were linear. Field studies in Pukeko birds also suggest that social interactions enhance the formation of orderly dominance hierarchies. Birds are more likely to form transitive dominance relationships (A dominates B, B dominates C, A dominates C) than cyclical relationships (A dominates B, B dominates C, but C dominates A). Their results suggest that dominance ranks are at least partially based on structural dependence between relationships rather than individual attributes alone [ 85 ].

Winner–loser effects are one key mechanism by which social interactions facilitate the formation of linear dominance hierarchies. Winner–loser effects mean that winners are more likely to win future encounters with any individual and losers are more likely to lose [ 86 ]. Winner–loser effects can occur when contest experience influences fighting behaviour or physiological traits like androgen titers [ 86 ]. Theory suggests winner–loser effects will generate linear dominance hierarchies via self-organizing dynamics, even without individual differences in intrinsic abilities [ 87 ]. Empirical work has shown that winner–loser effects occur in many species [ 86 , 88 ] and may contribute to stable dominance hierarchies [ 89 , 90 ]. Initial differences between individuals based on intrinsic attributes or random conditions are amplified with each social interaction, leading to stable, linear rankings. Intriguingly, the occurrence of winner and loser effects varies within and between species based on factors like social context [ 91 , 92 ] and genotype [ 89 ], suggesting that winner–loser effects may not be an inevitable consequence of competition. Instead, winner–loser effects may be an adaptive mechanism that minimizes conflict and stabilizes social relationships in situations where they are beneficial.

Winner–loser effects are unlikely to account for all patterns of self-organization in dominance hierarchies, so additional behaviours probably play a role. Bystander effects and localized social network properties have been proposed as potential mechanisms [ 8 ]. In taxa with bystander effects, an animal that observes a dominance contest between two others behaves differently than a non-observer when it meets the interactants [ 93 ]. Bystander effects are common across diverse taxa and encompass multiple specific behaviours. Priming occurs when observing any contest alters future contest behaviour. For example, bystander fish that observe contests between conspecifics are ‘primed’ to be more aggressive and more dominant than naive fish that do not observe contests [ 94 ]. Social eavesdropping occurs when individuals observe and remember the behaviour of specific individuals [ 13 , 53 , 94 ]. Some bystander effects, like social eavesdropping, may primarily influence dominance relationships by providing information about individual attributes, while other bystander effects, like priming, may mediate dominance relationships without involving assessment of individual attributes. Highly localized social network properties also mediate dominance hierarchies. Factors like aggressive state, fighting success, social status, motivation and random conditions influence rank via social feedback loops rather than having static effects on rank [ 95 , 96 ]. As a result, initial small differences between individuals are amplified over time via feedback loops, leading to differentiated ranks.

Given the accumulating evidence that dominance is influenced, in part, by self-organizing social dynamics, additional research will be important to understand both the behaviours that underlie these patterns [ 95 ] as well as the circumstances where self-organizing social dynamics are more versus less important. For example, social dynamics may be more likely to influence dominance hierarchies when orderly dominance hierarchies are beneficial [ 97 ]. In group-living taxa, orderly, stable dominance relationships minimize aggression and improve efficiency [ 16 , 17 , 98 ], so self-organizing dominance hierarchies may be common. Large groups that lack orderly hierarchies or groups where rank instability is less costly may be less likely to have self-organizing dynamics.

3.  Maintaining dominance relationships

After dominance hierarchies are established, other mechanisms are used to maintain hierarchy stability. Maintaining stable dominance hierarchies is particularly valuable in taxa that live in small groups with consistent membership. Social instability has negative effects on traits like reproductive success, offspring survival, stress responses and longevity [ 16 , 17 ]. For example, zebra finches in flocks with an established dominance hierarchy are more efficient foragers because stable social relationships improve the coordination and synchronization of foraging groups [ 99 ]. Subordinates sometimes aggressively test dominants because subordinates would benefit if they rose in rank [ 100 ]. Accordingly, individuals may preferentially direct aggression towards individuals immediately below themselves in the hierarchy [ 101 ] to avoid such rank reversals. However, individuals in stable groups typically recognize and abide by established dominance relationships because of the costs of repeated conflict.

Empirical work indicates that dominance hierarchies are largely stable. Challenges by subordinates are rare and dominants often maintain their status for much of their lifetime [ 81 , 102 – 104 ]. Often, long periods of hierarchy stability are interspersed with short periods of competition following the death of dominant or other changes in group membership [ 105 ]. Strauss & Holekamp [ 77 ] used 27 years of field data to show that rank reversals are relatively uncommon in hyena groups, but some individuals can improve their position in the hierarchy by working together with top allies. The stability of dominance hierarchies has also been studied by experimentally inducing conflict over rank. Cant et al. [ 106 ] experimentally induced conflict in groups of queen wasps by removing the dominant queen, allowing a subordinate to take over as the new dominant, then releasing the original dominant. In 34 of 35 trials, the original dominant retook her dominance position. Future research using long-term field observation and experiments inducing conflict over rank will provide additional insight into the long-term stability of dominance ranks and the factors that influence hierarchy stability [ 79 ].

There are two types of mechanisms that maintain hierarchy stability. First, multiple behaviours maintain the supremacy of the dominant, including punishment, threats and self-reinforcing behavioural differences. Second, social taxa have signals that provide information about rank and maintain stable hierarchies, especially individual identity signals and signals of dominance.

(a) Behaviours that maintain hierarchies

Dominants maintain their rank through threats, punishment and self-reinforcing behavioural mechanisms. A range of threat behaviours including dominance displays and minor aggression may maintain dominance relationships by providing information about an impending threat of eviction or attack by the dominants toward subordinates [ 18 , 107 ]. Threats are easily overlooked by researchers because many threat behaviours are subtle and effective threats rarely need to be carried out. Therefore, although threats may play key a role in maintaining dominance hierarchies [ 18 , 107 ], we have less direct experimental evidence of threats than other dominance maintaining behaviours.

While subtle threats can be difficult to quantify, there is strong evidence that punishment is involved in maintaining dominance hierarchies. We use Clutton-Brock's definition of punishment [ 19 ], retaliatory infliction of fitness reduction. Dominants punish subordinates via numerous behaviours, including eviction, aggression, social stress and infanticide. For example, some fishes have size-based dominance hierarchies where subordinates restrain their growth to maintain a minimum size difference between dominants and subordinates. When researchers created groups where the difference between dominant and subordinate size was smaller than the minimum difference observed in natural groups, dominants responded by forcibly evicting subordinates [ 108 ]. In taxa where dominants monopolize reproduction, dominants may punish subordinates attempting to maintain higher reproductive potential with aggression or by killing their eggs or offspring [ 19 , 39 ]. For example, subordinate wasps experimentally forced to maintain high reproductive potential are aggressively attacked by dominants [ 109 ]. Finally, subordinates that fail to contribute to the group by avoiding alloparental care or foraging may be aggressively attacked by dominants [ 110 ].

Dominance hierarchies can also be maintained without direct punishment or threats by dominants. In fact, dominance hierarchy position is often self-reinforcing because there are rank-linked disparities in feeding, work, and physical condition [ 111 ]. Dominants typically have priority access to food resources [ 102 , 103 , 112 , 113 ]. Access to more, higher quality food will accentuate initial differences in RHP or fertility between dominants and subordinates. In many taxa, subordinates also perform energetically expensive tasks like foraging more frequently than dominants [ 102 , 103 , 114 ]. The disparities between dominants and subordinates maintain reproductive and behavioural dominance hierarchies, so it is more difficult for individuals to increase their rank after a prolonged period as a subordinate.

(b) Signals that provide information about dominance rank

(i) individual identity signals.

The most common way to identify dominance rank in stable groups is individual recognition ( figure 3 ) [ 20 ]. During individual recognition, receivers learn the unique phenotype of conspecifics (termed individual identity signals), associate the phenotype with individual-specific information and recall the phenotype-information link during subsequent interactions [ 20 ]. Learning the unique phenotype and dominance rank of conspecifics provides a precise, non-cheatable method of assessing dominance relationships [ 115 ]. As a result, individual recognition plays an important role in dominance interactions in diverse taxa [ 20 ].

Individual recognition is a nearly universal mechanism involved in dominance relationships in vertebrate taxa where there are repeated interactions between a limited number of individuals. For example, individual recognition via plumage and calls is thought to maintain hierarchies in many stable bird flocks [ 116 ]. Primates individually identify group members using visual, olfactory and acoustic information [ 66 ]. Many taxa have surprisingly sophisticated knowledge about other individuals and their dominance relationships. For example, ravens know the unique calls and relative dominance ranks of individuals in their own group and neighbouring groups [ 117 ]. Baboons learn the individual identity of all group members and classify individuals according to both rank and kinship [ 118 ]. Individual recognition also occurs in invertebrates, though it is less widespread than vertebrates. Lobsters and crayfish use unique odours to maintain stable relationships with territorial neighbours [ 119 ]. Polistes fuscatus wasps learn the unique facial patterns and of many rivals and remember individual rivals over a week of separation [ 26 , 120 ].

Using individual recognition to keep track of dominance relationships is thought to have substantial cognitive costs. Individual recognition may be more costly than other types of recognition because it requires precise perception, learning and memory. Developing and maintaining the sensory and nervous tissues required to assess and remember individual identity signals involves constitutive costs [ 121 , 122 ]. For example, P. fuscatus wasps have visual and cognitive adaptations to facilitate individual recognition [ 123 – 125 ]. There are also operating costs associated with learning the unique features of many individuals, including the time, energy and resources required to collect, store and recall information. Long-term memory formation also reduces immunity, survival and fecundity [ 126 , 127 ].

The cognitive challenge of keeping track of many individuals has long been thought to play a key role in the evolution of brain size and cognitive abilities (social intelligence hypothesis). The social intelligence hypothesis proposes that enhanced cognition is favoured in species that live in complex societies because individuals with superior cognitive capacity can keep track of more individual relationships and respond appropriately during interactions [ 66 , 128 , 129 ]. Consistent with the social intelligence hypothesis, species that keep track of more individually differentiated social relationships often have larger brains, brain regions or enhanced cognition compared with species that have less complex social behaviour [ 68 – 70 ]. Although there is strong evidence that social behaviour influences cognition, social behaviour is only one of multiple selection pressures that shapes cognitive evolution [ 130 , 131 ].

Individual recognition often maintains dominance relationships in vertebrate groups when a limited number of individuals interact repeatedly, but the use of individual recognition is highly context-dependent. For example, Gelada baboons individually recognize close group members, but do not individually identify all the individuals they regularly encounter [ 118 ]. Many birds use individual recognition to mediate dominance relationships in small, stable groups, but the same species use status signals to assess rivals in large, unstable groups [ 29 , 30 ]. Polistes fuscatus nest-founding queens use individual recognition in multiple social contexts, but workers are much less adept at individual recognition and rely primarily on signals of dominance [ 132 ]. These studies highlight the importance of testing recognition in multiple social contexts instead of assuming that recognition measured in a single situation is universally applicable. As a result, there is much room for future work exploring what animals know about themselves and other individuals as well as how this social knowledge varies across contexts.

(ii) Signals of dominance

Signals of dominance provide information about the signaller's dominance status and also influence receiver behaviour ( figure 2 ). Chemical signals of dominance are ubiquitous in social insects. Dominance in social insect colonies is commonly defined as the individual that monopolizes reproduction [ 3 ], so dominance signals provide information about queen fertility and dominance rank [ 133 – 135 ]. Dominance signals also influence the behaviour of nest-mates, as workers and subordinate queens limit their own fertility in response to the queen's dominance signal [ 21 ]. Dominance is inherently relational, so the phenotype of reliable signals of dominance change as an individual's rank changes. For example, if a new individual takes over as the dominant queen, her chemical profile becomes queen-like [ 136 ].

Signals of dominance are a straightforward mechanism to rapidly and accurately assess dominance rank with relatively little cognitive investment. Responding appropriately to signals of dominance requires little learning and memory. Instead, individuals may treat any individual with a particular signal phenotype as the dominant without needing to learn the specific phenotype of their dominant. Consistent with a conserved receiver response, dominance signals in social insects are similar across at least three independent origins of eusociality and may have originated as fertility cues in the common solitary ancestor of all ants, bees and wasps, which lived approximately 145 Ma [ 135 ]. Although a few insects are capable of individual recognition [ 26 , 137 ], current work suggests that social insects primarily rely on dominance signals to maintain stable hierarchies on nests. Dominance signals provide a straightforward way to identify rank with lower cognitive cost than individual recognition [ 132 ].

‘Dominance signals’, ‘status signals’ and ‘signals of fighting ability’ are terms that are sometimes used interchangeably, but they are quite distinct ( figure 3 ). Signals of dominance are flexible traits that provide information about current dominance status, but not an individual's intrinsic fighting ability or RHP. Status signals or signals of fighting ability provide information about aspects of intrinsic ability like RHP [ 28 , 47 ]. Status signals are used to minimize conflict during dominance establishment and are often correlated with dominance rank. However, status signals do not provide information about rank per se . Polistes dominula paper wasps have both chemical signals of dominance and visual signals of status. On stable nests, Polistes use chemical signals to assess which individual is the dominant queen and chemical signals of dominance change as an individual's rank changes [ 138 , 139 ]. Polistes dominula also have visual signals that are used to assess the fighting ability of unknown rivals. The visual signals of fighting ability probably minimize the cost of conflict by allowing individuals to avoid escalated conflict with strong rivals [ 46 , 140 ]. Signals of fighting ability are one of multiple factors that mediate dominance hierarchy establishment, but they are not involved in maintaining hierarchies on nests. In fact, wasps ignore visual status signals on stable nests [ 141 ].

4.  Opportunities for future research

Dominance hierarchies have been the focus of much research in the 100 years since Schjelderup-Ebbe's [ 2 , 142 ] pioneering work in domestic fowl. It has become clear that dominance relationships are common in social taxa, as they provide a way for interacting animals to manage the trade-offs inherent in social interactions. New methods and analyses have facilitated recent advances in our understanding of the diversity and complexity of dominance interactions [ 77 , 143 , 144 ]. Here we review several approaches and opportunities for future research on dominance hierarchies.

(a) Self-organizing social dynamics

While it is clear that individual attributes have a huge effect on dominance hierarchy position, less is known about how other factors, especially self-organizing social dynamics, interact with individual attributes to influence rank [ 8 ]. This topic was tackled in wild baboons, finding that baboon hierarchies are based on both individual differences in fighting ability and winner–loser effects [ 89 ]. Notably, individual susceptibility to both mechanisms may have a genetic basis, suggesting that self-organizing dynamics co-evolve with individual attributes. Additional theory and experiments will be useful to test whether there are certain types of societies where self-organizing dynamics are more versus less important. For example, self-organization may be more important in societies where orderly hierarchies with transitive properties are most beneficial. By contrast, large or unstable social groups may be less likely to have self-organizing social dynamics. Newer network analysis methods will allow us to consider the role of self-organization in a way that was not previously possible [ 145 ].

(b) Hierarchy stability

Thus far, there has been more research on the factors that influence dominance hierarchy establishment than the factors involved in maintaining hierarchy stability [ 146 ]. Both theoretical and empirical work suggests that maintaining stable social hierarchies is fundamental to successful, long-lasting social groups. Hierarchy stability reduces conflict, saves energy, promotes survival and increases reproductive success [ 16 , 17 , 99 , 147 ]. At the same time, it is clear that there is extensive variation in both the stability of hierarchies and the factors that maintain stability. We have much to learn about the types of hierarchies that are the most stable as well as the factors that disrupt stability. Experiments can provide insight into both the behaviours that maintain social stability and the potential costs of instability. For example, when the queen Streblognathus peetersi ant is treated with a hormone that alters the queen's signal of dominance and fertility, queens are aggressively removed from their position as dominant and a new individual takes over as queen [ 148 ]. Network analyses also offer new opportunities to analyse the nuances of hierarchy structure as well as how and why stability varies [ 84 , 144 , 149 ].

(c) Plasticity

Animal behaviour is highly plastic, with individuals expressing different behavioural phenotypes in response to differences in the social environment, ecology or an individual's internal state [ 150 ] but see [ 151 ]. The mechanisms involved in dominance hierarchies are unlikely to be static. Instead, the way animals establish and maintain hierarchies may vary with traits like ecology (e.g. habitat saturation, food availability), social behaviour (e.g. group size, group consistency, costs and benefits of dominance rank) or individual characteristics (e.g. age, RHP, cognition, experience, genotype). Although relatively little is known about intra-specific plasticity in hierarchy formation and maintenance, there is some intriguing evidence of plasticity [ 152 ]. For example, the strength of winner/loser effects varies with the social environment [ 89 , 91 , 92 ], suggesting that the role of self-organizing social dynamics in hierarchy formation may also vary. In addition, animals communicate about dominance in different ways across different social contexts [ 29 , 30 , 116 ]. For example, P. fuscatus nest-founding queens use individual recognition to identify the rank of conspecifics prior to nest foundation [ 25 , 26 ], while workers largely ignore individual identity signals [ 132 ]. Instead, workers probably use chemical signals of dominance to assess dominance rank on stable nests [ 139 ]. Plasticity can be an experimental challenge because it is difficult to draw broad conclusions based on research conducted in a single situation. However, plasticity also presents exciting experimental opportunities to examine how and why dominance behaviour varies.

(d) Cognition

Cognition has an important role in the behaviours used to establish and maintain hierarchies. Although there is debate about what constitutes ‘advanced’ versus ‘simple’ cognitive tasks, it is clear that there is variation in how much learning, memory, sensory perception, abstraction and deduction are required for different dominance behaviours. For example, self-assessment during fights and signals of dominance are traditionally considered to be cognitively simple behaviours, while individual recognition, transitive inference, social eavesdropping and mutual assessment during fights are typically considered more cognitively challenging tasks. The capacity for cognitively sophisticated behaviour was originally thought to be limited by taxonomic group or brain size. However, there is growing evidence that behaviours traditionally considered to be complex are used by diverse taxa [ 141 , 153 ]. For example, Mantis shrimp use mutual assessment [ 154 ]. Astatotilapia burtoni fish are capable of social eavesdropping and transitive inference [ 62 ]. Polistes paper wasps use individual recognition, social eavesdropping, transitive inference and mutual assessment to manage dominance relationships [ 25 , 46 , 61 ]. Therefore, we need to think about the cognitive challenge of apparently complex behaviours in more nuanced ways.

There is much room for future work at the intersection of dominance and cognition [ 66 , 72 , 143 ]. First, we have much to learn about animals' capacity for complex social knowledge and how this knowledge influences social relationships. For example, what do animals know about themselves and other individuals? How does this social knowledge vary across contexts? How do differences in social knowledge influence the way animals form and maintain hierarchies? Second, we need a more nuanced perspective on the cognitive costs or limitations associated with the behaviours involved in hierarchy formation and maintenance. It is clear that learning, memory and sensory systems involve substantial costs [ 122 , 126 ] and the challenge of keeping track of dominance relationships can influence cognitive evolution [ 67 – 70 ]. However, animals could also minimize the costs by using simple mechanisms to mediate apparently complex behaviours [ 153 , 155 ].

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• Published: 10 June 2020

Social dominance hierarchy: toward a genetic and evolutionary understanding

• Bruce T. Lahn 1  

Cell Research volume  30 ,  pages 560–561 ( 2020 ) Cite this article

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In social animals, the formation of dominance hierarchy is essential for maintaining the stability and efficacy of social groups. A study by Wang and colleagues employ a combination of comparative genomic and functional approaches to shed new light on both the genetic mechanisms and the evolutionary histories of dominance behavior.

Many animals display social behavior of one sort or another, ranging from the relatively simple (e.g., food sharing in wolf packs) to the extremely complex (e.g., the formation of human societies). While social behavior is widespread up and down the animal kingdom, there is a clear evolutionary trend wherein the more derived species on the phylogenetic tree such as placental mammals tend to display more sophistication in their social behavior. 1 This trend reflects the fact that positive Darwinian selection generally favors greater sociality as it tends to confer enhanced fitness to the species.

A prevailing theme in highly social animals is the formation of social groups. Within-group cooperation on many activities such as the search for food, procurement of shelter, and provision of common defense, can enhance the fitness of all group members. Social groups are commonly structured as a dominance hierarchy based on a ranking system whereby higher-ranked individuals have better access to valuable resources such as food and mates but they also tend to assume greater responsibilities in providing leadership and maintaining order. 2 , 3 The formation of a hierarchical ranking system requires the dominant-subordinate relationship to be established between individual group members. This can be achieved by physical fights, proxy contests such as sizing each other up and visual displays of strength, and in cognitively advanced animals, political maneuvers. Importantly, in order for a hierarchy to remain stable, each individual needs to remember and follow its dominance rank vis-à-vis other members of the group until such time when challenging the established rank is deemed beneficial.

The first scientific investigation of dominance behavior can be traced to Thorleif Schjelderup-Ebbe about a century ago, when he used the now popular colloquial term “pecking order” to describe the dominance hierarchy in domestic chickens. 4 He noted that dominance in chickens is asserted by various types of behavior, including pecking by the dominant chickens on the subordinate ones, which he used as a measure of dominance rank. He proposed that such a hierarchy, while arising from and maintained by some degree of conflict (e.g., pecking), could enhance social stability and actually reduce overall conflict.

To date, the study of social dominance has mostly focused on the neurocircuitry and neurochemicals underlying dominance behavior. 3 Many brain regions have been associated with social decision-making including dominance behavior such as the prefrontal cortex and the amygdala. Several hormones and neurotransmitters such as testosterone, dopamine and serotonin have also been implicated. By contrast, not much is known about the genetic basis of dominance behavior and how it evolved.

A recent paper in Cell Research by Wang et al. offers a nice inroad into the genetics and evolution of dominance behavior. 5 This study took a somewhat unusual approach. The authors began by performing a whole-genome sequence comparison across 16 amniotes (12 placental mammals, one marsupial, two birds and one reptile) to screen for genomic regions exhibiting accelerated evolution in the common ancestral lineage leading to placentals. The most accelerated region identified in the screen was named placental-accelerated sequence 1 (PAS1). Among the 16 species used in the original screen plus three additional species added to the comparison (two marsupials and one monotreme), PAS1 is found to be strongly conserved in the placentals and be also well conserved among the non-placentals, but shows substantial divergence between placentals and non-placentals. This indicates dramatically accelerated evolution of this otherwise highly conserved region in the ancestral placental lineage. Indeed, PAS1 displays a rate of sequence change in this lineage that is over one order of magnitude greater than the null expectation of constant rate.

PAS1 is a putative enhancer located upstream of the LIM homeobox 2 ( Lhx2 ) gene. Lhx2 is a key regulator of embryogenesis including brain development. 6 The authors confirmed the enhancer activity of PAS1 on Lhx2 expression in the embryonic nervous system, and showed that PAS1 orthologs from different species can drive different patterns of gene expression in cell culture and in transgenic mice. They then generated three modified PAS1 alleles in mice, including knockout of endogenous PAS1 (PAS1 – ), knockin of wallaby PAS1 to replace the endogenous copy (PAS1 w ), and knockin of chicken PAS1 (PAS1 c ). The authors measured the dominant-subordinate relationship of mice carrying various combinations of these modified PAS1 alleles and the wild-type allele (PAS1 m ) using the classic tube test. In this test, two rodents are allowed to enter a tube from opposite ends. When they meet in the middle, the animal that manages to push forward in the tube is scored as being dominant, whereas the animal that retreats is considered subordinate. 7 PAS1 w/m mice are found to be more dominant than PAS1 w/w , whereas PAS1 c/m mice are less dominant than PAS1 c/c . It is important to note that in these mice, once a dominance rank is established between two individuals, it is stably maintained over time as would be expected in nature. Interestingly, tests of PAS1 –/m against PAS1 –/– mice showed that they failed to establish a stable dominance rank. One mouse could be dominant over another on one day, but the relationship could change on the next day. The authors concluded that PAS1, through its enhancer function on Lhx2 , acts as an essential regulator of social dominance behavior, and furthermore, PAS1 has played a role in the evolution of social hierarchy in amniotes.

The authors suggested that the accelerated evolution of PAS1 in the ancestral lineage leading to placentals might have contributed to the emergence of social hierarchy broadly observed in placentals. While this is a reasonable hypothesis, it should be pointed out that sociality is not a clear-cut defining trait demarcating placentals from non-placentals as prolonged gestation. While a high degree of sociality is found in many placentals, it is far from universal as many placental species are solitary. Additionally, some non-placentals are quite social, such as kangaroos and chickens. As such, there may be alternative interpretations for the accelerated evolution of PAS1 in the ancestral placental lineage. Another caveat of the study is that, even though the authors generated multiple PAS1 alleles in the mouse that could potentially give rise to many genotype combinations, only a few genotypes were used in the assessment of dominance rank (i.e., PAS1 w/m vs PAS1 w/w , PAS1 c/m vs PAS1 c/c , and PAS1 –/m vs PAS1 –/– ). This could be due to the difficulty in performing large amounts of animal breeding and testing. Nevertheless, with limited genotypes used in the study, it is difficult to interpret how PAS1 in different species modulates dominance behavior.

Notwithstanding the above caveats, the study is significant in that it provides the first example of an enhancer involved in the modulation of social dominance and it hints at the tantalizing possibility that the sequence evolution of this enhancer could have contributed to the phenotypic evolution of social dominance behavior in amniotes. Future studies are needed to bring more granular insights into the functional and evolutionary significance of this enhancer.

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Lahn, B.T. Social dominance hierarchy: toward a genetic and evolutionary understanding. Cell Res 30 , 560–561 (2020). https://doi.org/10.1038/s41422-020-0347-0

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