1

Evolution, Medicine, and Public Health [2020] pp. 1–6 c o m m en t a r y

doi:10.1093/emph/eoaa034

Evolutionary perspectives
on human behavior during
the coronavirus pandemic
Insights from game theory
Martin Brüne1,* and Daniel R. Wilson2
1
Division of Social Neuropsychiatry and Evolutionary Medicine, Department of Psychiatry, Psychotherapy and
Preventive Medicine, LWL University Hospital, Ruhr-University, Bochum, Germany and 2President, Western
University of Health Sciences, Pomona, CA, USA
*Corresponding author. Division of Social Neuropsychiatry and Evolutionary Medicine, Department of Psychiatry,
Psychotherapy and Preventive Medicine, LWL University Hospital, Ruhr-University Bochum, Alexandrinenstr. 1, D-44791
Bochum, Germany. Tel: þ49-234-5077-4410, Fax: þ49-234-5077-1329; E-mail: martin.bruene@rub.de
Received 10 May 2020; revised version accepted 26 August 2020

ABSTRACT
The coronavirus pandemic constitutes a global challenge to society and medicine. Here, we review
evolutionary insights that are relevant for the understanding of how people respond to the pandemic
and what to expect in the aftermath of the crisis. Specifically, we argue that the behavioral immune
system (BIS) and sickness behavior (SB) comprise two adaptive responses to impending and actual
infection, respectively, and that individuals activating their BIS differ from those showing SB in import-
ant ways that may have implications for the prevention and treatment of COVID-19. Moreover, we
reframe some of the behavioral health issues associated with the pandemic in a game-theoretical
scenario, illustrating the difficulties that arise when public health is treated as a ‘public good’.

Lay summary: The coronavirus pandemic constitutes a global challenge to society and medicine. In
this article, we employ evolutionary theory to improve our understanding of how people respond to
the pandemic. Specifically, we argue that human behavior is guided by ancient mechanisms involving
either the avoidance of infection or defense against attacks in times of enhanced vulnerability.
Moreover, we reframe some of the behavioral health issues associated with the pandemic in a game-
theoretical scenario. This helps understand why most people comply with rules of social distancing,
while a minority fails to do so for very different reasons. The evolutionary perspective also allows mak-
ing some predictions for the course of the pandemic.

K E Y W O R D S : coronavirus; behavioral immune system; sickness behavior; game theory; public

goods game

C The Author(s) 2020. Published by Oxford University Press on behalf of the Foundation for Evolution, Medicine, and Public Health.
V
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.
0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
2 | Brüne and Wilson Evolution, Medicine, and Public Health

INTRODUCTION characterized by fatigue, loss of appetite and drive, psycho-

motor retardation and social withdrawal. SB is also frequently
The current pandemic caused by coronavirus SARS-CoV-2 is an
associated with loss of appetite, which reduces the exposure to
extraordinary global challenge to society and medicine. What
toxic or infectious material, and heightened body temperature.
began as a medical crisis has rapidly and directly now led to crises
As such, it reflects an adaptive evolutionarily conserved defense
of a social, economic and even existential nature. Healthcare sys-
reaction to conserve energy and reduce the risk of being
tems, even in developed countries, are overburdened by increas-
attacked in times of enhanced vulnerability [9]. Some research-
ing numbers of critically ill patients, and in some places at risk of
ers have therefore highlighted the similarities between SB and
collapse. Here, we consider how evolutionary perspectives on
clinical depression, including immunological theories suggest-
health issues associated with the pandemic can explain difficulties
ing that aberrant priming of the immune system could be part
in handling infection risk and in dealing with at-risk populations
of an explanation for the ‘depression pandemic’, because the di-
and manifest COVID-19 patients. This viewpoint is consistent
vergent exposure to certain pathogens in modern environments
with previous work, suggesting that evolutionary insights are
compared to ancestral ones has created an evolutionary ‘mis-
mandatory in infectious disease surveillance, among other topics
match’ [10–12]. Similar to people activating their BIS, individu-
of interest to public health [1]. Specifically, we argue that the be-
als displaying SB avoid contact with strangers (for different
havioral immune system (BIS) and sickness behavior (SB) are im-
reasons, however), but seek proximity to close kin, even though
portant adaptive responses to impending infection and actual
this may impose costs to inclusive fitness, because it increases
infection, respectively, which may be associated with disparate be-
the risk of infection of genetically related individuals [3].
havioral attitudes impacting prevention and treatment of COVID-
With regard to the SARS-CoV-2 pandemic, aside from individ-
19. The problem of how to deal with the crisis, we assert, can be
uals showing signs of BIS or SB, at-risk groups deserve special
reframed in a game-theoretical scenario, such as the ‘Public
Goods Game’, whereby predictable uncertainties arise from con- medical attention. They comprise a heterogeneous array of syn-
ceptual difficulties in defining public health as a ‘public good’. dromal or diagnostic categories, including people with pulmon-
ary or cardiovascular disease, obesity, metabolic syndrome and
diseases associated with compromised immune function.
INFECTIOUS DISEASES AND HUMAN EVOLUTION Accordingly, age is also considered a risk-factor for severe or
Infectious disease has clearly shaped the human immune sys- fatal courses of COVID-19 [13]. In addition, many with psychi-
tem for most of our evolutionary history. It is less widely appre- atric conditions are burdened with one or more comorbid disor-
ciated, however, that infectious disease has also shaped human ders listed above, which makes them vulnerable to poor
behavior, though it has become increasingly obvious how tightly outcome of COVID-19, too. There is also evidence to suggest
the immune system is mutually linked to brain function [2]. that socially under-privileged persons suffer more from the
Humans evolved defense mechanisms against infectious dis- pandemic than economically more affluent individuals [14].
eases, known as the ‘behavioral immune system’ (BIS) [3]. This With respect to public health issues in relation to the SARS-
system is, by no means, specific to humans; even insects and CoV-2 pandemic, populations comprising a relatively large
lobsters avoid contact with infected conspecifics [4, 5], and in number of unaffected individuals, an increasing number of
higher vertebrates, this avoidance seems to be mediated by the infected people and a sizeable proportion of at-risk subjects
emotion of disgust [6], which is closely linked to the physical im- with regard to outcome, may act in quite specific and partly pre-
mune system [7]. The human BIS comprises the avoidance of dictable ways that may require public health measures and
social interactions posing a potential infection risk, conformity intervention programs. The at-risk group is relevant from an
and maintenance of cultural norms, i.e. ingroup coherence, as evolutionary point of view, specifically, because one can expect
well as neophobia. Activation of the BIS is thus associated with BIS and SB to occur more frequently in these patients com-
heightened vigilance toward and avoidance of outgroup mem- pared with the general population, and some (especially those
bers, with linkages to the fear system [8]. Indeed, it has been with psychiatric disorders, and the elderly) may be most
shown that humans are able to recognize even subtle signs of affected from the costs of social distancing, including social iso-
sickness in others, which activates an immune response in the lation, and loss of key benefits of sociality, such as within-group
observer, and fosters rejection and avoidance of the sick indi- cooperation and provision of safety [summarized in 15].
vidual [3].
While the BIS is activated by healthy individuals to prevent in-
A GAME-THEORETICAL APPROACH TO THE
fection, ‘sickness behavior’ (SB) refers to a psychophysiological
SARS-CoV-2 PANDEMIC
state usually caused by infectious agents (though in modern
environments, it can also occur in autoimmune disease, aller- The behavioral attitudes toward the SARS-CoV-2 pandemic can
gies and immune-modulating drug treatment). SB is theoretically be modeled using evolutionary game theory (with
Evolutionary perspectives on the coronavirus pandemic Brüne and Wilson | 3

BIS and SB impacting on social decision-making in manifold protects people from developing infectious diseases; second, im-
ways). Game-theoretical paradigms create situations requiring munization of a person does not preclude others from protec-
individual decisions to cooperate or defect (at the cost of tion; and third, once herd immunity has been established, no
others). Most neuroeconomic games concern the distribution one can be excluded from the protection it provides. Framed in a
of quantifiable (often monetary) resources between two or more PGG, most people comply with mandatory vaccination regula-
parties—individuals, groups or nations [16]. tions (thus contributing to the public good of herd immunity). As
Concerning the topic discussed here, the Public Goods Game herd immunity is already achieved in vaccination programs, if
(PGG) may be most informative. As far as health issues are con- only 90% of the population have been vaccinated, the public
cerned, the PGG has previously been utilized to explain social good of ‘herd immunity’ can even tolerate [17, 23].
dilemmata arising from the pros and cons of vaccination, herd However, as Dees (2018) argues, sanitation and clean water
immunity and use of antiviral drugs [17]; the PGG has, however, would not fall under this definition of a public good, because
not been applied to the spread of infectious diseases, or the neither is non-rivalrous or non-excludable. Indeed, many people
prevention thereof, when no vaccination is available. around the world are denied access to clean water and sanita-
The most common version of the PGG is played by an option- tion, and enjoying either one can be quite competitive.
al number of players who receive a defined amount of money or According to Dees (2018), public health as a public good thus
number of tokens at the beginning of the exchange scenario. warrants a normative component. Put another way, sanitation
Participants are asked to simultaneously invest their money in a and clean water ought to be non-rivalrous and non-excludable to
common pool (the public good), usually without knowing the qualify as a ‘normative public good’.
allowance of the other players. An experimenter multiplies the This distinction is important for the understanding of how
whole sum by a factor that is larger than one but smaller than epidemics and pandemics are dealt with. A normative public
the number of players, and returns an equal share of that money good, as Dees [23] points out, has four elements: first, it is a
to each player. In other words, all players benefit equally, irre- ‘good’; second, everyone has unlimited access to it, that is, it is
spective of how much they have invested before. If someone non-rivalrous and non-excludable; third, it benefits society (the
chooses a free-riding strategy while letting the others make their public) through collective effort; fourth, the good is important
contributions, his or her return will exceed those of the other enough to justify collective effort, a point that is open to debate,
players [18, 19]. If played iteratively, investments usually decline and thus also relevant for public health regulations in relation
over successive rounds, unless non-cooperative behavior can to the SARS-CoV-2 pandemic. In any event, by definition, com-
be sanctioned by the other players [20]. If punishment of selfish batting the SARS-CoV-2 pandemic qualifies as a normative pub-
behavior becomes part of the game, investments increase sig- lic good in every sense.
nificantly and remain stable across trials [21]. Interestingly, re- With this evolutionary background, we may now re-interpret
search utilizing the PGG has also demonstrated the occurrence several social-behavioral aspects of the pandemic:
of ‘antisocial punishment’, that is, the sanctioning of coopera-
tive behavior [22]. Strangely enough, at first sight, some individ-
Cooperation versus ‘defection’
uals tend to punish altruistic acts, particularly if the
punishment can be performed anonymously. The motivations In the absence of a vaccination against SARS-CoV-2, cooper-
for antisocial punishment can be diverse, ranging from a desire ation in this kind of PGG encompasses a variety of behavioral
to dominate others, competitiveness, or derogation of ‘do- means to reduce the risk of virus transmission. These include
gooders’. Conversely, antisocial punishment is constrained by ‘social distancing’, wearing face masks and identifying individu-
strong norms of civic cooperation, but less well contained in als who show signs suspect of infection. This is a mammoth
societies with weaker norms, which is, in part, explained by cul- task for society, because virtually everyone needs to comply to
tural differences in collectivism versus individualism [22]. achieve the public good of slowing the spread of infection.
In contrast to social decision-making in relation to the distribu- While being far from perfect, these measures help slow down
tion of quantifiable goods, the definition of ‘public health’ as a the number of new infections and protect those at increased
‘public good’ has instilled controversy [23]. In fact, it has been risk of detrimental outcomes of COVID-19. Activation of BIS is
argued that a public good comprises three elements: First, it probably supportive in this regard.
must be a ‘good’; second, it is non-rivalrous (i.e. the consump- However, social distancing has predictable downsides. One
tion of the good by someone does not preclude others to benefit is that social isolation is stressful and detrimental to health and
from it); third, it is non-excludable (i.e. one cannot be prevented immune function [9]. Moreover, social distancing does not
to enjoy it). According to these criteria, herd immunity (an im- work the same for individuals showing SB, and it may even not
portant goal in the handling of epidemics or pandemics, usually be required from those who have recovered from COVID-19. In
achieved through vaccination) constitutes a public good: first, it addition, BIS activation may be associated with anxiety to
4 | Brüne and Wilson Evolution, Medicine, and Public Health

spread more quickly as the virus itself. Indeed, it can be danger- observable in many countries around the world that, if political
ous, if not fatal, to overlook someone who is infectious, but in- decision-makers lift public health regulations too early, infection
expensive to believe someone is infectious who, in fact, is not. rates increase rapidly (as well as the death toll of COVID-19). It
Such hypervigilance can expand quickly in populations via so- is thus beyond fortune-telling to predict that almost certainly se-
cial or vicarious learning [24]. Accordingly, the dispersal of cond and third infection waves will hit the globe.
threat-detection mechanisms faster than the infectious agent
may give an additional survival advantage at the cost of false
positives. It is little more than speculation that social prejudices Antisocial punishment
may impact on the number of false positives, which would likely Strangely enough, but also predictably from a game-theoretical
include socially disadvantaged people, racial minorities, and point of view, one can also observe antisocial punishment in
perhaps mentally ill [15]. the current pandemic, at least indirectly and in subtle ways.
In contrast to cooperation in this special PGG, defining ‘free- This is a much more contentious issue compared to the
riding’ or ‘defection’ is much more difficult. Indeed, in neuroe- points discussed above, because, as we will argue, antisocial
conomic games, the failure to cooperate is usually conceived of punishment in relation to the SARS-CoV-2 pandemic comes in
as an intentional act. This is much less clear in a normative different disguises.
PGG where public health is the public good. More specifically, One is the frank denial of the existence of the infectious agent
there is certainly a minority of individuals who deliberately de- itself, or the view that the pandemic is much less dangerous as
fect, because they are willing to take the risk of own infection, suggested by virologists and other medical experts. Denying or
and, at the same time, do not care about the health of others. belittling medical concerns about the significance of the pan-
The percentage of such antisocial behavior may be low, how-
demic by official institutions including governments invites peo-
ever, particularly when compared to those who inadvertently do
ple to not obey to the rules of social distancing, perhaps more
not cooperate. Accordingly, inadvertent non-cooperation per-
so those whose BIS is only mildly activated.
tains to the very young, people with cognitive impairment inso-
Another (indirect) way to undermine cooperation in this par-
far as it compromises rational decision-making, and
ticular PGG is the spread of irrational or even delusional ideas
asymptomatic virus carriers. In the words of PGG, these individ-
about the origin of the virus or the idea that some socially high-
uals would count as ‘defectors’, even in the absence of deliber-
ranking individual utilizes the pandemic to take control over the
ate choice to not cooperate. With regard to a ‘normative public
world. Such bizarre propositions, we believe, arise from ances-
good’ such as public health, however, non-compliance with
tral fears, where invisible agents causing sickness and/or death
rules of social distancing cannot count as ‘free-riding’, because
incite suspiciousness against outgroups or putatively danger-
there is no short or long-term benefit for those who ‘defect’.
ous within-group coalitions [25]. That is, an enemy not from a
Reinforcing cooperation by public health regulations for these
competing group must come from one’s ingroup, usually some-
social groups is thus possible only by incurring costs on third
one who has great power or influence, or someone who lacks
parties, which includes parents (of young children), caretakers
protection from kin (accusation of witchcraft, etc.) [26]. This is
of elderly (e.g. at home or in nursing homes) etc.
a particularly dangerous aspect of the current crisis, because
modern technology accelerates the distribution of misinforma-
Development over time tion without effective control of its veracity. From a psychiatric
Theoretically, achieving herd immunity against SARS-CoV-2 point of view, the distinction between conspiracy theories and
slowly is, aside from the search for a vaccine, one potential delusional ideation may become reduced, idiosyncratic or
strategy to cope with the pandemic. The strategy is, however, volatile.
fraught with unpredictable consequences, particularly in terms In any event, conspiracy theories erode collective efforts to
of casualties and the risk for health services (and the economy) control the spread of the virus, and they can be specifically detri-
that a large proportion of the population could fall ill at the mental to the most vulnerable parts of society, i.e. at-risk sub-
same time. Unlike vaccination campaigns aiming at herd im- jects or individuals presenting with SB. This is particularly the
munity, however, the goal to slow down SARS-CoV-2 infection case, if conspiracy theories include the notion that vaccination
rates has a different temporal dynamic, in fact, an unpredictable against SARS-CoV-2 (once available) may be come at the cost
dimension with respect to time. of being ‘tagged’. Even if we do not know at present, whether or
This aspect is highly relevant, because the PGG logic predicts not vaccination will become available, and how effectively it will
that cooperation decreases over time, particularly, if no sanc- protect against COVID-19, the prospect that a sizeable number
tions have been put into place or, in the case of the pandemic, of individuals may fall prey to conspiracy theories and thus re-
may not work for reasons outlined above. It is already fuse vaccination is grim. Textbox 1 summarizes the most
Evolutionary perspectives on the coronavirus pandemic Brüne and Wilson | 5

Textbox 1. Bullet points derived from evolutionary insights about the SARS-CoV-2 pandemic

• In times of pandemics, people activate ancestral behavioral mechanisms to reduce the risk of infection (i.e. BIS), also impacting
social life.
• Game-theoretical approaches suggest that a significant (and as the pandemic continues) and growing number of people with fail
to comply with social rules to keep infection rates low.
• Accordingly, almost certainly second and third waves of COVID-19 will occur, unless an efficient vaccine will be available on a
mass scale.
• Antisocial punishment poses a threat to public health, as it undermines efforts to control the disease.

Textbox 2. Suggestions for public health measures

• The most relevant issue is continuing medical education of the public about SARS-CoV-2 to minimize the risk of defection and
the spread of false information.
• Regulation of social distancing and other measures that prevent virus transmission (e.g. compulsory face masks) is imperative. A
lesson learnt from game theory is that this must entail punishment for non-compliance.
• Preparation for a second (and third) wave is warranted, as well as for secondary health issues that may affect vulnerable or at-risk
populations most, including psychological problems following prolonged social isolation.

important insights and predictions from evolutionary theory on language and perhaps laughter evolved to allow ‘social groom-
the SARS-CoV-2 pandemic. ing’ at a distance, with remarkable physical effects mediated by
endogenous neuropeptides that are good for positive mood
and thus mental health [31].
SUMMARY
The pandemic will remain a challenge at least for an indeter-
The coronavirus crisis has created a natural experiment that has minate and perhaps lengthy time. Health care systems in devel-
put ancestral means of controlling the spread of contagious dis- oped countries erroneously believed that communicable diseases
ease in small-scale communities to the test in contemporary were primarily a threat to the less developed part of the planet,
mass societies. Individuals greatly differ with regard to their only to learn that the systems are vulnerable to unexpected
susceptibility to COVID-19 with a great many asymptomatic or (though predicted) attacks from the viral world [32]. As the virus
only mildly ill, yet others with severe syndromes that have a continues to evolve during its propagation around the world,
wide range of dramatic organic disease. Although research there is hope it may become more contagious, but less fatal
about the genetics underlying individual differences in vulner- (reduced virulence), though this is at present uncertain [33, 34].
ability is still in its infancy, the human leukocyte antigen (HLA) Unlike previous pandemics or endemic events, this pandemic will
may play a critical role [27]. In addition, there is limited evidence most likely leave no selective mark on the gene pool, because the
to suggest that people with blood group A carry a greater risk vast majority of fatalities occur beyond the reproductive lifespan.
for detrimental outcomes of COVID-19 as opposed to 0 [28]. With the first measures taken by governments to lift the shut-
Recent research indicates, however, that it is possibly not the down, it needs to be kept in mind that decisions over the reduc-
blood group per se that confers risk for COVID-associated pneu- tion of the burden of ‘social distancing’ are largely driven by eco-
monia. Instead, the presence of Anti-A antibodies seems to be nomic policy, not medical advice. Public health authorities need
protective against severe lung affection, particularly IgG [29]. to take action to reduce incipient or expressed policies of Social
Social contact has been crucial to fitness over millions of Darwinism. Textbox 2 comprises suggestions for public health
years in our evolutionary history, and failure to achieve social measures derived from evolutionary insights.
connection can therefore activate stress mechanisms that real- Moreover, in the aftermath of the pandemic there likely will
ize immediate survival at the expense of longer-term health. As be a second epidemic of people who are grieving because they
West-Eberhard put it, ‘individuals of social species having these lost loved-ones to the pandemic, or people suffering from post-
specialized characteristics are in a sense trapped into group traumatic stress disorder (notably among hospital staff).
life, and group living may become virtually ‘obligatory’ for Individuals may develop depression, anxiety disorders, addic-
them.’ [30, p. 224]. Yet, physical distancing is not necessarily tion or other maladaptive stress-coping conditions. This may
equal to social distancing, particularly not in our species, as pose a challenge specifically on mental health workers and
6 | Brüne and Wilson Evolution, Medicine, and Public Health

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