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The neuroeconomics of cooperation

Through cooperation we are able to thrive, build societies, culture and technology. But history also reveals our
potential for selfishness, spite and prejudice. Studying the neural processes that drive choice behaviour is essential
to understand this paradox and develop means to curb greed and extend the limits of cooperation.

Carolyn H. Declerck and Christophe Boone

A
s with all social mammals, human
behaviour is guided by core motives
that fulfil survival and belonging
needs. Meeting both these needs at the
same time, however, creates a cooperation
problem because different motives may
drive behaviour in opposite directions:
social motives aimed at group integration
make us care about the fate of others and
encourage actions that benefit the group,
while self-serving motives tempt us into
free-riding. In situations where individual
and group interests conflict, a cooperative
decision favouring the group is therefore
vulnerable to abuse by selfish others.
To avoid deceit, cooperation in this case
must hinge on trust and expectations of
reciprocity, which may seriously confine the
Credit: Science Photo Library/Alamy Stock Photo
range in which we are willing to exert effort.
How is this dilemma resolved? When do we
decide to cooperate, and which information
do we use in the process? When we associate that region with a Hence ventral striatum activation by itself
Unveiling motives is challenging because particular function, sensation or motive is not sufficient to differentiate between
motives are not readily observable, and that was elicited by the task, we must the motivating forces of greed versus care.
simply asking for them is not a reliable remember that all brain regions fulfil many People also rely on external information to
method to understand the process of functions, and that the observed pattern form expectations of rewards, beliefs about
decision-making because we are not of activation could therefore be the result others and impressions of trustworthiness,
always aware of or willing to share the of several causes. But as the number of all of which help to decide whether or not to
reasons for why we behave the way we independently conducted neuroimaging cooperate. This information, processed in
do. A neuroeconomic approach offers a experiments is growing, we begin to have sensory regions of the brain that project to
fuller understanding of the motives behind converging evidence showing that it is the striatum, has furthermore been shown
decisions, and can inform us about possible possible to unravel motives from observed to influence the satisfaction obtained from a
interventions to effectively expand and brain–behaviour relationships. cooperative decision3.
sustain large-scale cooperation. In a recent book, Neuroeconomics of
Neuroeconomics combines research Imaging the brain Prosocial Behaviour4, we summarized the
paradigms from behavioural economics and One of the first studies investigating how evidence for the proposition that different
psychology with neuroimaging techniques. the brain responds to a dilemma eliciting decision rules for (non)cooperation are
In a typical experiment, participants are conflicting motives indicated that mutual supported by distinct neural networks
conducting a choice task with various cooperation corresponds with activation that integrate different sources of sensory
conditions that have consequences for self in the ventral striatum1, a region of the input, and we provide a testable mechanism
and others. Their brain is scanned using brain lying at the crossroads of limbic and for how it might work: when, on the one
functional magnetic resonance imaging cortical neural circuits that is implicated hand, self-interest is elicited, the decision to
(fMRI), making it possible to reveal the in motivational learning and reward cooperate (or not) depends on information
neural underpinnings that link a particular processing. Interestingly, it does not regarding extrinsic incentives, which
motive (triggered by a task condition) matter whether the reward is lucrative we expect to be processed by regions in
to the actual choice a person makes. (self-serving) or social. A systematic review the lateral prefrontal cortex (LPFC), the
An important caveat, however, is that fMRI indicates that there is much overlap in the evolutionary most recent part of the human
cannot reveal cause and effect relations. sites of ventral striatum activation when cortex known from previous research to play
It conveys the extent to which an event participants in the scanner are either an important role in computation, working
during the task correlates with the metabolic receiving a sum of money or experiencing memory and self-control. Consistently,
needs of a particular region of the brain. the ‘warm glow of giving’2. experiments during which participants
438 Nature Human Behaviour | VOL 2 | JULY 2018 | 438–440 | www.nature.com/nathumbehav

(in the scanner) needed to evaluate the
presence or absence of extrinsic cooperative
incentives showed increased activation in
the LPFC that correlated with a calculated
decision, namely to cooperate when it
was satisfactorily remunerated, and to
defect otherwise5.
When, on the other hand, social
motives are triggered, we expect decisions
to be supported by different neural
networks involved in processing social and
emotional information4. A particularly
important region in this regard may be the
temporoparietal junction (TPJ). It projects
to the ventral striatum and is known to
be involved in decisions that require trust
or perspective taking. Morishima and
colleagues showed that, when motives
conflict, an increase in TPJ activation
corresponded with a shift towards the social
side: activity in the TPJ predicted generosity,
but only when the conflict of motives was
peaking; when the cost of being generous
was either very cheap or too expensive, TPJ
activity was unrelated to decision-making6.
While activity in isolated brain regions
such as the TPJ and LPFC appears to
be necessary to solve a cooperation
dilemma, the process is certainly more
complex, involving the interplay of several
interconnected brain regions. This was
convincingly shown by Hein et al. who
scanned the brain of benefactors in a
donation game7. The decision to donate
was motivated either by empathizing
with someone’s pain, or by reciprocating
someone’s kindness. Depending on which
of these two motives was triggered, the
strength of the correlations between
activated brain regions that fed into the
ventral striatum changed substantially. Thus,
a cooperative decision can be supported
by different patterns of brain activation
depending on the motive that provoked it.
Neural signature differences
People vary in their intrinsic inclination
to cooperate. Some are primarily driven
by greed, while others reciprocate, and
still others are willing to bear the cost of
another’s well-being. These inclinations
are also reflected in neural networks.
Individuals who consistently seek self-
interest and who cooperate only when there
are extrinsic incentives to do so tend to have
an enlarged and more active LPFC5,8. In
contrast, individuals with a stable prosocial
disposition (those who consistently favour
equal over selfish decisions) tend to have an
anatomically enlarged TPJ6 and a stronger
connection between the ventral striatum and
the anterior insula7, an emotion-processing
hub in the brain with an important role in
generating empathy and developing trust.
Nature Human Behaviour | VOL 2 | JULY 2018 | 438–440 | www.nature.com/nathumbehav
The examples above suggest that intuitive
prosocial or selfish tendencies can become
reinforced to the point that some neural
pathways are more chronically activated
than others, leaving a neural signature in
the brain corresponding to values. Such
internalized values are the compass by
which to arrive at quick and frugal solutions
in ambivalent social situations: individuals
with a chronically active LPFC may have
a heightened focus on incentives by which
they fulfil their self-serving needs and favour
selfish decisions. In contrast, individuals
with a chronically active insula and TPJ are
more likely driven by compassion and social
belonging motives. They will cooperate
readily, provided they can trust4.
Not surprisingly, most societies and
institutions attach much importance to
nurturing their children’s social values,
teaching them to share and care in the hope
to tilt the compass away from greed and
towards more cooperation. However, by
triggering the social motives underscoring
belonging needs, prosocial values may also
inadvertently foster parochialism, or bias
decisions to favour those who are close to
us at the expense of distantly related others.
By contributing to ingroup/outgroup feuds,
and in extreme cases, to ethnocentrism,
radicalism and religious warfare, narrow-
minded prosociality has a dark side.
Given today’s tensions in geopolitics
and global economy, a major challenge
for future research in the social sciences
is to understand how to overcome
parochialism without undermining
cooperation. Economists have long
emphasized the role of incentives and
advocated inducements for improving
desirable behaviour. But the insights from
neuroscience discussed here suggest that
there may be other routes to cooperation
that bypass pecuniary gratification.
Improving cooperation
The human brain is remarkable for its
neuroplasticity, by which it is able to alter
the strength of its synaptic connections
between neurons, both in response to
short-term environmental stimuli (activity-
dependent plasticity) as well as during
long-term learning processes (cortical
remapping). This gives us an immense
capacity to flexibly switch behaviour as we
see fit, and also to cultivate the enduring
behavioural changes necessary to implement
collectively beneficial and sustainable
cooperative goals. How can we engage this
chameleon-like capacity to tap our multiple
motives and use them to expand the
boundaries of cooperation?
First, we could learn from behavioural
economists how to ‘nudge’ cooperative
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decisions by creating environments that
elicit social motives9. Popularized by the
2017 Nobel Laureate Richard Thaler,
‘nudging’ refers to a cheap and non-coercive
behavioural modification tool that changes
a person’s field of view, introducing a bias
in automatic decision-making processes.
Such tactics might work especially for
individuals with self-serving values. In the
donation study by Hein and colleagues7,
activating the empathy motive by drawing
attention to someone’s suffering not only
had a proportionally greater impact on
selfish individuals, but also changed the
connectivity pattern of the ventral striatum
so that it resembled that of prosocials.
Similarly, highlighting partnerships and the
commonalities between people might be an
effective way to bypass the computational
neural circuits of the LPFC and, instead,
foster empathic feelings and trust, which
facilitate our innate capacity to cooperate.
Second, to encourage cooperation
without fuelling parochialism, prosocial
values need to be accompanied by a healthy
dose of reasoning to counterbalance the bias
introduced by motives that satisfy our need
to belong. However, reasoning capacity is
supported by neural networks that include
the LPFC, the region of the brain that
also endorses self-interest! How does the
LPFC support both social and self-serving
motives? Recent work with brain stimulation
techniques reveals an interesting way by
which the LPFC influences reasoning in the
social domain. By experimentally enhancing
activity in the LPFC, Gross et al. increased
participant’s willingness to violate rules
that demanded hurting someone without
additional benefits10. Training the reasoning
capacities associated with the LPFC may
thus be an effective way to implement a
more inclusive cooperative goal, because
it would reduce the restrictions imposed
by social values that inadvertently also
sustain ingroup favouritism. This might
help especially prosocial individuals to
refrain from parochialism in situations that
require them to follow a norm that may no
longer be adaptive.
A brain that is paradoxically wired to
motivate both our own as well as others’
well-being is perhaps an asset in sustaining
large-scale cooperation. With the ability
to empathize, share perspectives, compute
trade-offs between incentives and values,
and integrate them with experience and
current conditions, we are well geared to
care for others, trust and achieve outcomes
that benefit the larger collective. Further
research into the factors that trigger
the activation and connectivity of brain
regions such as the LPFC or TPJ will
continue to shed more light on how social
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motives and reasoning interact when
making narrow- versus broad-minded
cooperative decisions. Such knowledge
may help us to create ethical principles
and policies that underscore prosocial
values, yet avoid the perils of parochialism.
In addition, neuroeconomic experiments
that incorporate the influence of values
into the study of decision-making could
inform more personalized interventions by
clarifying why incentives, norms and nudges
do not work for everyone alike.
The tragedy of the commons does not
have to be an irreversible fate. That we
progress in our understanding of how,
when and for whom the brain compromises
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conflicting motives in favour of a
cooperative and open mind is grounds
for optimism.
Carolyn H. Declerck* and Christophe Boone
Faculty of Economic Sciences, University of Antwerp,
Antwerp, Belgium.
*e-mail: carolyn.declerck@uantwerpen.be
Published online: 9 July 2018
https://doi.org/10.1038/s41562-018-0387-3
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Author contributions
The authors contributed equally to this work
Competing interests
The authors declare no competing interests.