RUNNING HEADING: Partner choice 1

The psychology of partner choice

Justin W. Martin, Liane Young, Katherine McAuliffe

Department of Psychology, Boston College, Chestnut Hill, MA

Correspondence: [email protected] (J.W. Martin)

Word count: 7,966

Keywords: partner choice; cooperation; punishment

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Partner choice

Abstract

Partner choice captures the idea that individuals exist in a biological market of potential

partners, and we can therefore choose or reject our social partners. While prior work

has principally explored the functional basis of partner choice, here we focus on its

mechanistic basis, motivated by a surge of recent work exploring the psychology

underlying partner choice decisions. This work demonstrates that partner choice is

predictably sensitive to a number of factors, including 1) a potential partner’s generosity

and fairness, 2) cooperative disposition, 3) moral decision-making, and 4) intentions.

We then broaden our scope, first reviewing work suggesting that, in some cases, the

psychology underlying partner choice may be distinct from other responses to a

partner’s behavior. We then discuss work demonstrating the sensitivity of partner

choice decisions to market characteristics as well as work that illuminates the neural,

ontogenetic and phylogenetic basis of partner choice. We conclude by highlighting

outstanding questions and suggest directions for future research.

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Partner choice

The importance of partner choice

Human success – as a species and as individuals – depends upon our ability to

interact with others: To work together, to help and be helped and to form social bonds.

Equally important are the mechanisms that exist to maintain these interactions.

Theoretical work has outlined a number of such mechanisms, including indirect

reciprocity (Alexander, 1987), direct reciprocity (Trivers, 1971), kin selection (Hamilton,

1964; all reviewed in Rand & Nowak, 2013), punishment (Boyd & Richerson, 1992; Fehr

& Gächter, 2002; Guala, 2012; Henrich et al., 2010; Yamagishi, 1986) and partner

choice (Barclay, 2016; Baumard, André, & Sperber, 2013; Hammerstein & Noë, 2016;

Noe & Hammerstein, 1994). And, empirical work has shed light on the psychology

supporting many of these mechanisms supporting successful interactions (Raihani &

Bshary, 2015; Rand & Nowak, 2013; Warneken, 2017). However, until recently, less

has been known about the psychological underpinnings of partner choice. Our goal in

this review is to discuss recent work on the psychology of partner choice and to

encourage further work on this emerging topic.

At the most fundamental level, partner choice is the ability to leave or reject a

partner (Barclay, 2016). Individuals exist in a marketplace of potential partners (a

biological market; Hammerstein & Noe, 1994), and this leads market dynamics—supply

and demand—to influence the formation of partnerships. Broadly, this yields three main

goals for individuals: Attracting partners, choosing partners, and maintaining partners

(Barclay, 2013). In this review, we focus on the latter two capacities: How we choose

and maintain partners (see Barclay, 2013, 2016 for a discussion of how to attract

partners). More specifically, we evaluate our relative options and choose the best
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Partner choice

partners we can. Once a partnership is formed, we end or continue it depending on the

relative quality of our outside options. Such decisions may be made based on

characteristics of the population (e.g. the presence of relatively more skillful or

cooperative partners) as well as characteristics of individuals. Here, within this latter

category, our primary focus is on internal characteristics that are relevant to an

individual’s social normative behavior (in the parlance of person perception, those that

lie along the “warmth” dimension; Fiske, Cuddy, & Glick, 2007) and those that may also

be relevant for moral evaluation, though we also offer a brief discussion of internal

characteristics beyond this focus.

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Partner choice

Figure 1: The four major questions about partner choice. The four major questions

about partner choice, from a levels of analysis perspective (Tinbergen, 1963). At the

top are questions regarding the proximate mechanisms of partner choice; on the bottom

are questions regarding ultimate explanations for partner choice. While other work has

focused on a functional account of partner choice (Barclay, 2013, 2016; Baumard et al.,

2013), our review focuses principally on a mechanistic account of partner choice. We

also briefly discuss its ontogenetic and phylogenetic basis.

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Partner choice

Theoretical work on partner choice in humans has mainly investigated partner

choice from a why perspective (Tinbergen, 1963): for instance, considering the

implications of partner choice for cooperation (Barclay, 2013, 2016; Misyak, Melkonyan,

Zeitoun, & Chater, 2014; Warneken, 2017), the evolution of fairness (Baumard et al.,

2013) and moral religions (Baumard & Boyer, 2013). Here, we primarily take a how

perspective, focusing on the mechanistic basis of choosing partners and maintaining

partnerships in humans (Figure 1). Drawing on recent evidence from a range of fields –

including evolutionary biology, evolutionary psychology, moral psychology, behavioral

economics, developmental psychology and judgment and decision-making – we show

that partner choice is supported by a suite of identifiable mechanisms. In particular, we

highlight four factors that partner choice is sensitive to: Indications of 1) generosity and

fairness, 2) cooperative disposition, 3) a partner’s moral decision-making and 4) a

partner’s intentions. The sensitivity of partner choice to these factors provides insight

into the psychological mechanisms underlying partner choice decisions. We argue that

these mechanisms appear to be, in some cases, importantly different from those

regulating social interactions by other means.

We then expand the focus of this review in important ways. First, we consider the

extent to which the psychology underlying partner choice is unique or shared with other

ways of responding to others’ behavior, using sensitivity to intentions and outcomes as

a test case. Second, we review work investigating how the influence of the personal

characteristics highlighted earlier can be modulated by market characteristics. We then

review evidence on the neural correlates of partner choice. Next, we additionally

provide evidence that partner choice is deeply rooted in both phylogeny and
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Partner choice

development, with signatures of partner choice emerging across a range of animal taxa

and in infancy in humans. Finally, we briefly touch on the consequences of partner

choice and conclude by suggesting promising avenues for new research on partner

choice1.

Many modes of partner choice

Partner choice has been studied using a number of different paradigms across a

variety of domains. Here, we bring these paradigms together under one heading and, in

so doing, we hope to provide researchers with a menu of options for studying partner

choice in the future (see Figure 2).

The key feature of partner choice paradigms that we focus on here is that the

target individual is deciding whether or not to continue or enter into a new relationship

with another individual. We focus principally on contexts involving resource distribution,

1
We reviewed a range of articles when writing this article. Our main focus was on recent work (within the
past 5 years), as (1) other reviews on partner choice exist and are referenced in this the article and (2) we
are interested in current approaches to this topic. We conducted our search with 3 main criteria. First, we
targeted empirical work that involves resource distribution. For instance, mate selection can be thought of
as a special case of partner choice (Buss & Schmitt, 1993; Clutton-Brock & McAuliffe, 2009; Dall,
Giraldeau, Olsson, McNamara, & Stephens, 2005; McNamara & Houston, 1986; Noe & Hammerstein,
1994; Rusbult, Zembrodt, & Gunn, 1982), but one that we consider beyond the scope of the current
review. Second, we prioritized incentivized contexts, where participants’ earnings are dependent upon
their choices, though we include especially relevant work in non-incentivized contexts (e.g. Lammers et
al., 2017). Third, we focused on cases in which a single individual was making a choice about interacting
(or not interacting) with another single individual (generally in comparison to one or more other
individuals). We do this because this is a paradigm case of partner choice (Barclay, 2013, 2016). Thus,
we focus less on cases in which groups are deciding whether or not to exclude an individual, though such
contexts are relevant; we include especially relevant work that falls into this category (e.g. Liddell &
Kruschke, 2014; Page et al., 2005; Rockenbach & Milinski, 2011). Our literature search involved the
search terms “partner choice”, “partner selection”, “choice AND cooperation”, “trust game”, “partner
switching”, and “economic game AND partner” and was conducted using PsycInfo and Google Scholar.
For relevant articles, we then searched their cited references, using the same categories. The search was
conducted in November and December, 2017.
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Partner choice

where the choice of partner will affect how much one can earn and is thus an impactful

one for participants. We do this partly out of necessity: Most of the existing work on

partner choice has been in these contexts. However, the incentivized nature of these

decisions gives particular weight to any inferences drawn from them. This is especially

important in the domain of social interaction, given the fact that the factors we say we

care about are sometimes not the ones that influence our judgments or behavior (Blake,

McAuliffe, & Warneken, 2014; Carlsmith, Darley, & Robinson, 2002; Cushman, Young,

& Hauser, 2006). Moreover, the fact that these paradigms generally involve resource

distribution does not indicate that they are uniform. They include studies framed as

highly de-contextualized interactions (Raihani & Barclay, 2016) as well as those

explicitly described to participants as relationships in a hunter-gatherer society

(Eisenbruch & Roney, 2017). And, they investigate partner choice in response to

simple descriptions of past behavior (e.g. Everett, Pizarro, & Crockett, 2016) as well as

in response to observed, concrete behavior (e.g. people assign either themselves or

their partners to an onerous task (Simpson & Harrell, 2013) or donate some amount to

charity (Fehrler & Przepiorka, 2016).

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Partner choice

Choose between potential partners Continue with a partner or switch partners

a b

VS VS

(Aksoy, 2015; Bornhorst, Ichino, Kirchkamp, Schlag, & Winter, 2010; Bostyn & Roets, 2017; (Barclay & Willer, 2007; Coricelli, Fehr, & Fellner, 2004; Eisenbruch & Roney, 2017; Gross, Leib,
Everett, Pizarro, & Crockett, 2016; Fehrler & Przepiorka, 2016; Hardy & Van Vugt, 2006; Offerman, & Shalvi, 2018; Liddell & Kruschke, 2014; Martin & Cushman, 2015; Page,
Pleasant & Barclay, 2018; Raihani & Barclay, 2016) Putterman, & Unel, 2005; Rockenbach & Milinski, 2011; Sylwester & Roberts, 2010, 2013)

Continue with a partner or go it alone Exclude or not

c d

Exclude?
VS

(Barclay, 2006; Fehrler & Przepiorka, 2013; Jordan, Hoffman, Bloom, & Rand, 2016; Jordan,
Hoffman, Nowak, & Rand, 2016; Lammers, Gast, Unkelbach, & Galinsky, 2017; Przepiorka &
Liebe, 2016; Simpson & Harrell, 2013) (van der Lee, Ellemers, Scheepers, & Rutjens, 2017)

Figure 2: The many modes of partner choice. The many modes of partner choice. (a)

Choosing between two potential partners. (b) Interacting with a partner and then

choosing to either continue the partnership or switch to a new partner. (c) Entering into

an interaction (e.g., trust game) or staying out of the interaction, based on some

information about the potential partner. (d) Rejection behavior by members of a group.
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Table 1. Economic games described in this review and their description.

Game Description
Dictator Game (DG) Player One is given an endowment of a resource and

can give any amount of their endowment to Player Two.

Player Two is a passive recipient.

Trust Game (TG) Player One is given an endowment of a resource and

can give some or all of their endowment to Player Two.

Any amount sent to Player Two is multiplied by a

specified factor (often 3). Player Two can then return

any amount of the resource they receive to Player One.

Prisoner’s Dilemma (PD) Player One and Player Two each decide to either pay a

small cost to deliver a larger benefit to the other (i.e.

cooperate) or to earn a resource for themselves at a

cost of a resource to the other (i.e. to defect).

Public Goods Game (PGG) A group of players (often 4) are each given an

endowment of a resource. Each can give some or all of

their endowment to a group pot. The amount of

resources in the group pot is multiplied (often by 1-2x)

and divided evenly among all players (including those

who did not contribute to the group pot).

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Perhaps the most representative paradigm is when a participant chooses

between two (or more) potential partners for an economic interaction, whether that be a

trust game (see Table 1 for a description of this and other economic games referenced

here) (Bornhorst, Ichino, Kirchkamp, Schlag, & Winter, 2010; Bostyn & Roets, 2017;

Everett et al., 2016; Fehrler & Przepiorka, 2016; Pleasant & Barclay, 2018), a prisoner’s

dilemma (Aksoy, 2015), or a dictator game (Hardy & Van Vugt, 2006; Raihani &

Barclay, 2016). Closely related are contexts in which participants interact with a partner

and decide whether they would like to play an economic game with that partner again or

with a different partner, either in a subsequent dyadic context (Barclay & Willer, 2007;

Coricelli, Fehr, & Fellner, 2004; Eisenbruch & Roney, 2017; Gross, Leib, Offerman, &

Shalvi, 2018; Martin & Cushman, 2015; Sylwester & Roberts, 2010, 2013) or a group

context (Liddell & Kruschke, 2014; Page, Putterman, & Unel, 2005; Rockenbach &

Milinski, 2011). We note that these group exclusion decisions have been previously

studied under the heading of ostracism. While our principal interest is in the decision of

one individual to partner (or not) with another, decisions by individuals in a group

context to exclude another member can nevertheless inform our understanding of

partner choice.

Some studies have asked participants whether they would like to enter into a

trust game with an individual based on some information about that individual (Barclay,

2006; Fehrler & Przepiorka, 2013; Jordan, Hoffman, Bloom, & Rand, 2016; Jordan,

Hoffman, Nowak, & Rand, 2016; Lammers, Gast, Unkelbach, & Galinsky, 2017;

Przepiorka & Liebe, 2016; Simpson & Harrell, 2013). The choice here is not which of

two individuals to interact with, but a decision between staying out of the game (and
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keeping one’s endowment) and entering into a game with the potential partner. Finally,

some studies have looked at rejection behavior by groups in non-economic social

contexts (van der Lee, Ellemers, Scheepers, & Rutjens, 2017) and at preferences for

avoiding individuals in non-economic contexts (Kemper & Newheiser, 2017).

In children, while measures are necessarily less sophisticated and less specific

to partner choice (e.g. social evaluation can be different from partner choice), they

nevertheless can serve as early signatures of partner choice. For instance, children can

choose between two puppets about whom they have some information (Hamlin, 2013;

Hamlin, Wynn, Bloom, & Mahajan, 2011; Woo, Steckler, Le, & Hamlin, 2017), or choose

to take an item from one puppet or another (Lucca, Pospisil, & Sommerville, 2018;

Tasimi & Wynn, 2016). They can also be given a resource to distribute between two

partners (A. Dahl, Schuck, & Campos, 2013; Kenward & Dahl, 2011) or asked which of

two partners they like more (Shaw, DeScioli, & Olson, 2012).

Characterizing the psychology underlying partner choice

These paradigms have yielded a number of important insights, allowing us to

offer a preliminary characterization of the psychological mechanisms underlying partner

choice. We organize this characterization around four main features of individuals that

are relevant to their social normative behavior (see Figure 3). Of course, this is not an

exhaustive list of factors partner choice could be sensitive to, and we conclude with a

brief discussion of features beyond these four. Moreover, these factors are related – for

instance, generosity and cooperative disposition co-vary (Peysakhovich, Nowak, &

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Rand, 2014). Thus, this division should not be taken to imply a bright-line demarcation

between them. Rather, focusing on these factors is a useful way of conceptually

dividing the space of features partner choice is sensitive to.

We choose partners who have been generous and fair

Much work suggests that partner choice is sensitive to signals of generosity and

fairness. For instance, we appear to choose partners who donate to others (Fehrler &

Przepiorka, 2013, 2016) or give to others in a dictator game (Przepiorka & Liebe, 2016).

We prefer partners who are generous to us (Eisenbruch & Roney, 2017; Hackel, Doll, &

Amodio, 2015) and as well as to others (Raihani & Barclay, 2016). This applies both

when the generosity cannot be repaid by the participant (Hackel et al., 2015; Raihani &

Barclay, 2016) and when it can (Eisenbruch & Roney, 2017).

However, there are constraints on and moderators of the influence of generosity

and fairness on partner choice decisions. While we prefer those who behave

altruistically, this preference is much stronger when we learn that others (i.e. not the

focal individual) generally behave consistently in that context. That is, when we learn

that people other than the focal individual behave inconsistently (equally likely to be

selfish or cooperative) or consistently (either only selfish or only cooperative) in this

context, we prefer those who have behaved altruistically in the consistent context

(Lammers et al., 2017). And, there is individual variability in preferences for the

generosity of potential partners. For instance, though the majority of individuals prefer

those who donate, some subjects appear to instead choose partners who did not
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donate as much, potentially to avoid feeling any sort of obligation to give more to these

individuals (Fehrler & Przepiorka, 2016). Finally, preferences for generosity in others

can be goal-dependent. When choosing someone to judge us, we prefer an altruistic

individual when we previously behaved fairly and a selfish individual when we

previously behaved unfairly (Melnikoff & Bailey, 2018). Thus, preferences for fairness

and generosity in partners are nuanced and vary across context and individual.
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a b c
Cooperative
disposition Intentions

Moral
Generosity
decision-
VS and fairness
making

Figure 3: How partner choice decisions are made. A schematic of how partner choice

decisions are made. (a) First, individuals for consideration are selected out of a

population of “outside options”. (b) The subsequent partner choice decision is guided by

four key factors: Indications of 1) generosity, 2) cooperative disposition, 3) a partner’s

moral decision-making and 4) a partner’s intentions. (c) Finally, the cooperative

interaction takes place.

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Individuals care about whether someone has been generous or not when

selecting them as a partner, but there are two reasons why this information might be

important. One possibility is that those who are generous provide greater material

benefits or rewards. On the other hand, generosity might signal a deeper prosocial

disposition, indicating that this individual will reliably be prosocial in a variety of

contexts. Recent work has demonstrated that both factors matter. When presented

with potential partners that are equally fair, individuals do choose those who can deliver

greater benefits (Raihani & Barclay, 2016). However, when these two sources of

information are in tension - when forced to choose between a stingy but wealthy

individual and a generous but poor individual - a partner’s demonstrated generosity has

a much bigger influence on how we choose partners than benefits delivered (Hackel et

al., 2015; Raihani & Barclay, 2016).

In sum, while individuals generally choose those who are generous and fair as

partners more than those who are stingy or selfish, these preferences can be nuanced

and influenced by individual differences and contextual factors.

We choose partners who have been cooperative

We also choose partners who have been cooperative2. We prefer those who

play cooperatively or contribute more, both in group contexts (e.g. a public goods game)

(Hardy & Van Vugt, 2006; Page et al., 2005; Pleasant & Barclay, 2018; Sylwester &

2
Here we focus on cooperation as being potentially more strategic than generosity: We are generous in
contexts in which we expect no return from the cost we pay, whereas in cooperative contexts there may
be some expectation of reciprocation. However, this distinction is mainly one of convenience –
cooperation and generosity are strongly related (Peysakhovich et al., 2014).
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Roberts, 2010) and dyadic contexts (Barclay & Willer, 2007; Bornhorst et al., 2010;

Coricelli et al., 2004; Lammers et al., 2017), and we exclude those who contribute the

least (Rockenbach & Milinski, 2011). We pick cooperative individuals for subsequent

cooperative interactions (Hardy & Van Vugt, 2006; Pleasant & Barclay, 2018; Sylwester

& Roberts, 2010), and they receive more in subsequent partner choice contexts (Hardy

& Van Vugt, 2006; Sylwester & Roberts, 2013; but see Barclay, 2004). But, there are

potential limits to the influence of a cooperative disposition on partner choice: Holding

past cooperative behavior constant, we prefer in-group members as future partners over

out-group members (Aksoy, 2015).

When deciding whether to interact with someone by investing money in them (in

a trust game), participants care about whether this potential partner was previously

willing to pay costs to provide benefits to others (Jordan, Hoffman, Nowak, et al., 2016).

And again, similar to signals of generosity, previous work has investigated the degree to

which benefits delivered versus cooperative intent matter more. While benefits

delivered do influence partner choice decisions, what appears to matter more is signals

of a cooperative disposition (Eisenbruch & Roney, 2017).

We choose partners based on how they make moral decisions

It is perhaps unsurprising that we choose to interact with cooperative and

generous partners. We appear to also care about the moral decisions our partners

make, in somewhat surprising ways.

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Partner choice

For instance, we care about the content of potential partners’ moral decisions. In

certain contexts, we prefer partners who make deontological choices rather than

utilitarian ones (Bostyn & Roets, 2017; Everett, Faber, Savulescu, & Crockett, 2018;

Everett et al., 2016). And, we prefer to avoid those who commit purity violations,

relative to those who have harmed others or broken a social norm (Kemper &

Newheiser, 2017). This is consistent with work demonstrating that we view those who

commit purity violations as having particularly abnormal or atypical underlying desires

(Chakroff & Young, 2015). In the context of choosing a partner, this abnormal

underlying disposition may mean that this agent is likely to behave in counter-normative

ways and is thus someone to avoid as a partner. However, preferences for moral

decision making can take a back seat to other considerations – when presented with a

task in which a dishonest partner leads to a better chance of higher payoffs for the

dyad, individuals will switch away from honest partners and toward dishonest partners

(Gross et al., 2018).

Beyond the content of others’ moral decisions, we also care about the strength of

partners’ moral standards. We choose partners with strong and inflexible moral

standards over those with weak or flexible moral standards, and in fact we prefer those

with stronger moral standards than our own (Simpson & Harrell, 2013; van der Lee et

al., 2017). And we care about whether individuals are willing to enforce norms: We

invest more money in individuals who previously punished selfishness on behalf of a

victim (i.e. as a third-party) (Barclay, 2006; Jordan, Hoffman, Bloom, et al., 2016). This

appears to be specific to partner choice decisions: Third-party helpers are rewarded

more than third-party punishers (Barclay & Raihani, 2016). And, the third-party nature
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of such norm enforcement may be important: Those who punish in a public goods

game, in which group members are neither clearly impartial third-parties nor victims, are

not preferentially picked for interactions (Rockenbach & Milinski, 2011).

Thus, when selecting a partner for a future interaction, we care about the kinds of

moral decisions they make: The way they treat others, the kinds of moral preferences

they hold and how strongly they hold them, and their concern for moral norms.

We choose partners based on their intentions (rather than the outcomes they

cause)

As discussed above, in a variety of contexts investigating a variety of factors,

adult partner choice decisions are influenced by the intent with which a partner has

behaved and less by the material benefits delivered (Eisenbruch & Roney, 2017; Hackel

et al., 2015; Liddell & Kruschke, 2014; Martin & Cushman, 2015; Raihani & Barclay,

2016). Even among children, intentions play an important role: Children around eight to

ten months of age preferentially choose to interact with agents who accidentally cause

harm relative to agents who intentionally cause harm (for a review see Van de

Vondervoort & Hamlin, 2016; Hamlin, 2013; Woo et al., 2017). This early incorporation

of information about intentions may be unique to partner choice – other judgments do

not show such intent sensitivity until later in life. For instance, when children around the

age of 4 and 5 are asked about deserved punishment, their judgments focus to a larger

degree on the outcome that was caused (Cushman, Sheketoff, Wharton, & Carey,

2013; Nobes, Panagiotaki, & Bartholomew, 2016; but see Van de Vondervoort &
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Partner choice

Hamlin, 2017). Punishment judgments do not attain adult levels of incorporation of

information about intentions until 12 years of age (Gummerum & Chu, 2014). Of

course, this relative difference in the weighting of intention information could be due to

differences in the methods used, precluding strong conclusions. Nevertheless, these

results do suggest that there is an early-emerging capacity for choosing between

individuals that incorporates information about their mental states in a manner similar to

adults, whereas other responses to a partner’s behavior do not begin to look adult-like

until much later in development. Thus, in terms of the processes partner choice is

sensitive to, information about others’ intentions appears to be weighted more than

information about the outcomes they cause, potentially across the lifespan.

We choose partners based on features outside social normative behavior

Features beyond those related to social normative behavior (those on the

“competence” dimension; Fiske et al., 2007) also influence partner choice. As

discussed above, although intent to deliver benefits carries greater weight, we do select

those who are wealthy, either when that wealth is directly relevant to how much we can

earn presently (Hackel et al., 2015; Raihani & Barclay, 2016) or when it is a more

general signal (Nelissen & Meijers, 2011). Partner selection is also influenced by the

ability to deliver benefits, whether through reputation or prestige (for a review see

Henrich & Gil-White, 2001). Finally, we care about a partner’s ability to generate

benefits, both when this is framed around luck (e.g. being endowed with a greater ability
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to generate wealth) or skill, although to a greater extent in the latter case (Eisenbruch &

Roney, 2017).

The psychological distinctiveness of partner choice

Our highlighting of signals of generosity and fairness, cooperative disposition, a

partner’s moral decision-making and a partner’s intentions as important to partner

choice does not necessarily mean that they are uniquely important for partner choice.

Indeed, it is likely that some mechanisms are more uniquely related to partner choice,

whereas others are less so. Next, we consider when and why we might expect the

mechanisms supporting partner choice to be psychologically distinct. Here, we use the

functional purpose of partner choice as a guide. To the extent that partner choice

serves a particular function—one that may be different from other ways of responding to

a partner’s behavior—we may expect the mechanisms supporting partner choice to be

relatively distinct. In particular, we focus on punishment – sometimes studied under the

heading of partner control (Bshary & Grutter, 2002; Fruteau, Lemoine, Hellard, van

Damme, & Noë, 2011; Martin & Cushman, 2015; Schino & Aureli, 2015) – as an

alternative to partner choice, one which has been traditionally emphasized as a

response to others’ behavior (Clutton-Brock & Parker, 1995; Trivers, 1971). From a

functional perspective, punishment may exist to change others’ behavior (Clutton-Brock

& Parker, 1995; Martin & Cushman, 2016; Trivers, 1971), engage in costly signaling

(Barclay, 2006; Jordan, Hoffman, Bloom, et al., 2016) or enforce norms (Boyd &

Richerson, 1992; Fehr & Gächter, 2002; Henrich et al., 2010; Yamagishi, 1986).
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However, punishment does not serve to select partners and so may serve as a strong

contrast with partner choice.

One possibility is that partner choice and punishment are supported by the same

set of mechanisms – if each is utilized to ensure we are surrounded by cooperative

individuals, for example, they may stem from the same decision-making process.

Indeed, some past work has described or investigated punishment and partner choice

as part of the same continuum, with partner choice viewed as a severe form of

punishment (Cinyabuguma, Page, & Putterman, 2005; Feinberg, Willer, & Schultz,

2014; Hoffman & Goldsmith, 1999; Kerr et al., 2009; Liddell & Kruschke, 2014; Schulz,

2016; Seymour, Singer, & Dolan, 2007). However, partner choice and punishment

represent very different routes to ensuring the presence of cooperative partners: Partner

choice is used to select and be selected as the best partner, whereas punishment is

used to make others behave more cooperatively. Given these different proximate

targets, these two responses could in fact be supported by distinct underlying

mechanisms.

A well-studied feature of punishment decisions is their particular sensitivity to the

presence of bad outcomes, studied in the literature as outcome bias or moral luck (for a

review see Martin & Cushman, 2016). Whereas we do not view agents who accidentally

cause harm as behaving particularly wrong, we do view them as worthy of punishment

(Cushman, 2008). In contrast, outcomes on their own do not appear to matter as much

when choosing a partner. As discussed above, when researchers have pitted

information about benefits delivered (a salient outcome) against information about an

agent’s intended generosity and asked participants to choose a partner, a partner’s

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Partner choice

demonstrated generosity has a much bigger influence (Eisenbruch & Roney, 2017;

Hackel et al., 2015; Raihani & Barclay, 2016). This is true even in contexts in which

preferentially weighting generosity over reward is suboptimal (Hackel et al., 2015).

These results suggest that the inputs important to selecting partners are different

from the inputs important to punishing. Recent work has directly investigated this: in

both a dyadic economic game (Martin & Cushman, 2015) and a variant of the public

goods game (Liddell & Kruschke, 2014), the decider’s intent and the outcome they

caused were dissociated using a stochastic mechanism. Participants responded to the

decider’s choice and the outcome by either applying punishment or engaging in partner

choice. In both studies, punishment and partner choice were applied differently: Partner

choice decisions were less sensitive to outcomes than were punishment decisions.

Instead, partner choice was conditioned principally on how the partner intended to play.

In other words, in response to the same inputs, punishment and partner choice are

applied differently, potentially suggesting that punishment and partner choice are

subserved by distinct underlying processes.

The distinctiveness of partner choice has been supported by recent

developmental evidence as well. If partner choice and punishment are supported by

distinct underlying processes, we might expect them to emerge in development

differently. In particular, given the importance of information about intentions in making

partner choice decisions, we might expect sensitivity to intentions to emerge first in

partner choice. Indeed, recent work demonstrates that when children are asked to

make judgments about punishment and partner choice while actors’ intentions and
 24
Partner choice

outcomes caused are varied, partner choice decisions demonstrate sensitivity to

intentions at an earlier age than punishment (Martin, Leddy & McAuliffe, in prep).

Thus, we find that, in both adults and children, punishment and partner choice

decisions are distinct. Partner choice decisions are conditioned more on agents’

intentions than the outcomes they cause relative to punishment decisions, and this

sensitivity to intentions emerges earlier for partner choice decisions than punishment

decisions. Combined, these results support the idea that the psychological processes

supporting punishment and partner choice are distinct in important ways.

Market influences on personal characteristics

Market characteristics can have an important influence on how sensitive we are

to personal characteristics (Bednarik, Fehl, & Semmann, 2014; Debove, Andre, &

Baumard, 2015; Fehrler & Przepiorka, 2016; Lammers et al., 2017; Rockenbach &

Milinski, 2011). For instance, when many outside individuals (“proposers”) compete to

partner with a solitary chooser, the chooser can weight information about generosity

heavily, whereas this is less true when there are equal numbers of choosers and

proposers (Debove et al., 2015). When generosity varies within a market (indicated by

donations to charity), participants choose partners who are more generous, even

though more generous individuals are not necessarily better partners in this context

(Fehrler & Przepiorka, 2016). In a market where partner choice is more costly,

individuals are more sensitive to previous cooperativeness, relative to markets in which

partner choice is less costly or free (Bednarik et al., 2014). And, a partner’s past
 25
Partner choice

behavior matters less for partner choice in markets in which we know that individuals

behave inconsistently relative to markets in which we know that individuals behave

consistently across partners (Lammers et al., 2017). Finally, in a market where

punishment is possible, relative to one where punishment is not possible, those who

punish are not preferentially chosen as partners (Rockenbach & Milinski, 2011), even

those we do prefer those who punish in other contexts (Barclay, 2006; Jordan, Hoffman,

Bloom, et al., 2016). Thus, the social context or market in which we find ourselves

influences the features of partners that we care about, across a number of features and

a number of market manipulations.

Market characteristics also impact our consideration of a potential partner’s

ability to generate benefits (Eisenbruch & Roney, 2017; Raihani & Barclay, 2016). First,

when wealth is stable – we know a partner will start with the same pot of money across

trials – we weight information about a partner’s past fair behavior more heavily,

choosing a partner who is poor but fair over a partner who is rich but selfish (though

only by a small margin) (Raihani & Barclay, 2016). This preference is drastically

increased when wealth is unstable, suggesting that in variable contexts – relative to

stable contexts - we care more about how a partner intends to treat us (Raihani &

Barclay, 2016). Second, our preferences for partners who are better at generating

wealth are sensitive to whether the market is one in which skill vs. luck is highlighted

(Eisenbruch & Roney, 2017). When success in a market is described as dependent on

skill, we choose partners who can generate more benefits than when market success is

described as dependent on luck (Eisenbruch & Roney, 2017). However, it remains the

case that, in a market where individuals vary on how generous they are and how
 26
Partner choice

effective they are at generating benefits, information about generosity is prioritized in

selecting partners (Eisenbruch & Roney, 2017).

Neural correlates of partner choice

In the final sections of this review, we broaden our focus, first by discussing work

on the neural correlates of partner choice. While these neural correlates remain poorly

understood, related work has investigated how individuals choose interaction partners in

the Trust Game (TG) (for a review see Tzieropoulos, 2013; Delgado, Frank, & Phelps,

2005; King-Casas et al., 2005; Krueger et al., 2007). The investor in the TG faces a

type of partner choice decision: They can either keep their endowment – in effect

rejecting an interaction with the potential partner – or decide to enter into the interaction.

Work on the neural basis of these decisions has yielded two main findings. First,

activity in the striatum tracks with the trustworthiness of the potential partner, with higher

activity for less-trusted individuals (Tzieropoulos, 2013). Second, paracingulate and

cingulate cortices support the mentalizing necessary to anticipate a partner’s behavior

(Tzieropoulos, 2013). Other related work largely corroborates these results. In one

study, participants made partner choice decisions in two different domains (Yokoyama,

Sugiura, Yamamoto, Nejad, & Kawashima, 2017). Across both, choice behavior was

associated with activity in posterior cingulate cortex and the right temporoparietal

junction. In another study, participants chose between potential partners for an

economic game on each trial (Hackel et al., 2015). Partners varied in demonstrated

cooperative disposition and the amount of reward they could deliver. Activity in
 27
Partner choice

ventromedial prefrontal cortex coded for the difference in generosity between chosen

versus non-chosen partners (Hackel et al., 2015). And, participants who weighted

generosity higher in their choices had larger differential activity in ventromedial

prefrontal cortex (vmPFC). Given the role of vmPFC in representing decision-related

value signals, this suggests that vmPFC combines information about reward value and

generosity when making a partner choice decision. Indeed, similar vmPFC activation

was found in other work comparing social vs. non-social partner choice (Heijne, Rossi,

& Sanfey, 2017). Here, participants chose one of four possible partners or slot

machines that varied in expected value provided on each trial. Activity in vmPFC was

highest for staying with the previously chosen partner in the social condition relative to

the non-social condition. In total, though limited research on the neural basis of partner

choice decisions exists, the results implicate a number of regions – including the

striatum, paracingulate and cingulate cortices, ventromedial prefrontal cortex and

temporoparietal junction - in the core processes underlying partner choice decisions.

Partner choice has deep developmental and evolutionary roots

Recent work suggests that partner choice has deep roots in human psychology,

by probing both the breadth of partner choice across nonhuman animal species and the

development of partner choice in children.

Phylogenetic roots of partner choice

 28
Partner choice

Work on partner choice in animals has approached it from both functional (Bull &

Rice, 1991) and proximate mechanism (Schino & Aureli, 2015) perspectives. Partner

choice (in a potentially cooperative context) has been observed in ravens (Asakawa-

Haas, Schiestl, Bugnyar, & Massen, 2016), a variety of primate species - including

chimpanzees (Engelmann & Herrmann, 2016; Engelmann, Herrmann, & Tomasello,

2015; Melis, Hare, & Tomasello, 2006; Molesti & Majolo, 2016; Suchak, Eppley,

Campbell, & de Waal, 2014; Suchak et al., 2016), capuchin monkeys (Sabbatini, De

Bortoli Vizioli, Visalberghi, & Schino, 2012) and vervet monkeys (Fruteau, Lemoine,

Hellard, van Damme, & Noë, 2011; reviewed in Schino & Aureli, 2015) – wasps

(Grinsted & Field, 2017) and fish (reviewed in Dugatkin & Wilson, 1993; Bshary &

Grutter, 2002; Bshary & Schäffer, 2002; Vail, Manica, & Bshary, 2014). For instance,

when chimpanzees are presented with an adapted version of the trust game, they

preferentially choose to cooperate with those who have previously been cooperative

(Engelmann et al., 2015). This taxonomic breadth appears comparable to punishment

(Clutton-Brock & Parker, 1995; Riehl & Frederickson, 2016), which has been found in

chimpanzees (Jensen, Call, & Tomasello, 2007; Koyama, Caws, & Aureli, 2006; Suchak

et al., 2016), rhesus macaques (Hauser, 1992) and fish (Bshary & Grutter, 2002;

Raihani, Grutter, & Bshary, 2010). In primates, relationship quality is a large

determinant of partner choice decisions (Asakawa-Haas et al., 2016; Engelmann &

Herrmann, 2016; Sabbatini et al., 2012; Suchak et al., 2014). However, when

demonstrated cooperative tendency has been manipulated, partner choice is also

sensitive to these signals (Engelmann et al., 2015; Suchak et al., 2014; but see

(Sabbatini et al., 2012). The comparative breadth of partner choice and punishment is
 29
Partner choice

interesting, given the potential difference in costs borne by those engaging in these

behaviors: Whereas punishment invariably/commonly involves a cost to the one

enacting it, partner choice may, by contrast, carry an immediate benefit. For instance, if

the current partner is not as productive, skillful or cooperative as potential other

partners, interacting with a new partner / engaging in partner choice may be

immediately beneficial. Given this asymmetry in associated costs, we may expect

partner choice to be more phylogenetically widespread than punishment, and future

work should explore this question.

While comparative work demonstrates the widespread existence of partner

choice in the animal kingdom, it does not provide strong clues as to the mechanisms

supporting these decisions. For this, we can turn to evidence from the development of

partner choice in children.

Ontogenetic roots of partner choice

Signatures of partner choice appear early in development. Eight to ten-month

olds choose those who accidentally cause harm over those who intentionally cause

harm (Hamlin, 2013; Woo et al., 2017) and prefer those who act positively toward

prosocial individuals and act negatively toward antisocial individuals (Hamlin et al.,

2011). Children also appear to incorporate information about fairness in partner choice

decisions in relatively sophisticated ways. Children at 13 months of age can already

use information about distributions a third party makes to others to inform their social

evaluations: After observing a third party distribute resources either equally or

 30
Partner choice

unequally, infants prefer to interact with (e.g. accept a toy from) someone who

distributes fairly (Lucca et al., 2018). And, infants are willing to give up resources to

interact with someone who helps others rather than harms others (Tasimi & Wynn,

2016). However, there are limits to these generalizations: Infants do prefer those who

are consistently prosocial over those who are consistently anti-social, but evince no

preference when individuals are not completely consistent (Steckler, Woo, & Hamlin,

2017). By around 2 years of age, children prefer to help those who have helped others

over those who have harmed others (A. Dahl et al., 2013), though this partiality

competes with preferences for equality (Kenward & Dahl, 2011). By the age of 6,

children’s preferences for fairness must be reconciled with preferences for favoritism

toward themselves, and this preference for favoritism is increased in competitive

contexts (Shaw et al., 2012).

Recent work has begun to probe the development of partner choice decisions

more directly (Martin, Leddy, & McAuliffe, in prep). Here, children were presented with

vignettes in which a protagonist’s intentions and the outcome they caused were varied.

That is, the protagonist could either intend harm or not, and could cause harm or not.

For half of the experiment, children were asked whether engaging in punishment was

appropriate and for the other half they were asked whether engaging in partner choice

was appropriate. When assessing punishment, results mirrored prior work: Decisions

were sensitive to outcomes early in life and only demonstrated sensitivity to intentions

with development (Cushman et al., 2013; Hebble, 1971; Imamoglu, 1975; Nobes et al.,

2016; Nobes, Panagiotaki, & Pawson, 2009; Zelazo, Helwig, & Lau, 1996). When

assessing partner choice, a different pattern emerged. By 5 years of age, children were
 31
Partner choice

sensitive to whether a protagonist intended harm. Thus, we find that punishment and

partner choice emerge in development differently: Partner choice decisions are

sensitive to intentions at an earlier age than punishment decisions.

These results relate to past work on the development of children’s moral

judgments. For instance, prior work has found that preschoolers’ judgments of moral

character (a judgment similar to partner choice) pattern with their punishment judgments

(Van de Vondervoort & Hamlin, 2017). However, children in this study were presented

with cases in which intentions and outcomes were matched: A protagonist either both

intended and produced a bad outcome or intended and produced a good outcome.

Consistent with this, we find that punishment and partner choice judgments are similar

in the case of intentional harm. This pattern of results nicely illustrates the point that

punishment and partner choice decisions are often similar, and only distinct in cases in

which the factors they are sensitive to differ. Thus, future work should continue probing

these responses, focusing in particular on cases in which these responses may be

teased apart.

Thus, in adults, children and animals (at least non-human primates), partner

choice decisions are affected by a partner’s demonstrated cooperative tendencies. This

is not to say that this is expressed similarly in all cases: Whereas children and adults

are sensitive to a partner’s adherence to fairness norms, this has not been

demonstrated in animals. However, this behavioral consistency across species may

suggest that at least some of the psychology underlying partner choice may also be

conserved across species.

 32
Partner choice

Consequences of partner choice

Engagement in partner choice is just one side of the equation – partner choice

also has consequences for those who are rejected. Exploring these consequences

yields insights into the functional purpose of partner choice. One such functional

purpose is to promote cooperation. Being rejected can lead individuals to become more

cooperative with subsequent partners (Coricelli et al., 2004). This is especially true

when communication and gossip are possible (Feinberg et al., 2014; Strømland, Tjøtta,

& Torsvik, 2016) and is less likely when punishment is also possible (Pleasant &

Barclay, 2018). Further, the mere option of engaging in partner choice (even when it

only occurs infrequently) leads group members to be more cooperative (Bednarik et al.,

2014; Wu, Balliet, & Van Lange, 2016), although partner choice appears less effective

at encouraging cooperation when it can occur unilaterally rather than by mutual decision

(Hauk & Nagel, 2000). Being chosen for an interaction by a partner (versus being

randomly assigned) leads one to be more willing to accept less advantageous offers

from that partner (Falco, Albinet, Rattat, Paul, & Fabre, 2019). Finally, being excluded

and then re-included by a group leads to greater risk-taking on behalf of that group (E.

Dahl, Niedbala, & Hohman, 2019). However, there is also conflicting evidence. Being

excluded can lead individuals to avoid those who are excluding them (e.g. in an

extended version of cyberball; Driscoll, Barclay, & Fenske, 2017). In other words, we

return a potential partner’s rejection with our own rejection. And, there are some

examples where partner choice does not lead to higher levels of cooperation (Barclay &
 33
Partner choice

Raihani, 2016). How do we resolve these conflicting findings? One potential mediator

of the influence of partner choice is the number of partners involved: Partner choice

appears to most effectively encourage cooperation in contexts of multiple partners

(Bednarik et al., 2014; E. Dahl et al., 2019; Feinberg et al., 2014; Strømland et al., 2016;

Wu et al., 2016) and work less well in a dyadic context (Barclay & Raihani, 2016). This

is consistent with the idea that partner choice is most effective in the presence of

outside options, or other (potentially more cooperative) partners (Baumard et al., 2013;

Debove et al., 2015). Another mediator of the influence of partner choice appears to be

future oriented thinking: Those who are more future oriented become more cooperative

following rejection, whereas those who are less future oriented become less cooperative

(Balliet & Ferris, 2013). In sum, future research should explore the influence of partner

choice on the one not chosen, especially focusing on delineating contexts in which

rejection encourages versus discourages cooperation.

Outstanding questions

A large number of questions remain about the psychology underlying partner

choice. Here, we highlight a few of the most pressing and interesting avenues for future

research. First, what is the role of reasoning in partner choice? Are partner choice

decisions mainly supported by implicit or explicit processes? Second, are partner

choice and punishment subserved by distinct neural computations? We have reviewed

research exploring the neural correlates of partner choice, but it is unclear to what

extent these are shared with other kinds of responses or unique to partner choice.
 34
Partner choice

Third, to what extent is partner choice influenced by group psychology? Are we

particularly harsh when making a partner choice decision about in-group transgressors,

or are we instead more lenient? Fourth, are there individual differences in the tendency

to engage in partner choice relative to other responses (e.g. punishment)? Do

individuals have a preferred method for dealing with transgressors and if so, how strong

is this preference? Fifth, what is the scope of the functional consequences partner

choice has? For instance, do individuals learn from partner choice, both when

personally rejected and when observing others being rejected? Does this tendency to

gather information from partner choice decisions vary across individuals? Sixth, how

does partner choice vary cross-culturally? Much work suggests that the use of

punishment varies across cultures (Barrett et al., 2016; Henrich et al., 2010, 2006;

Herrmann, Thöni, & Gächter, 2008) – does the use of partner choice similarly vary?

Seventh, how did partner choice first evolve in humans? As a mechanism for mate

selection and then subsequently for selection of cooperative partners, or vice versa? Or

did these capacities evolve relatively separately, as some have suggested (Hrdy, 2011).

And, do distinct psychological mechanisms support partner choice in distinct contexts

(e.g. when selecting a mate versus a cooperative partner)? Eighth, what is the

relationship between the development of partner choice and the development of theory-

of-mind capacities? That is, to the extent that partner choice is particularly sensitive to

intentions, we might expect these capacities to be tightly coupled. However, no

research has explored this question.

Conclusions
 35
Partner choice

Potential partners exist in a biological market; in this market, we evaluate our

options and decide who to partner with and who to avoid. Our partner choice decisions

have important consequences, yet, until recently, the psychology supporting them has

been poorly understood. Here, we reviewed a recent surge of evidence on the

proximate mechanisms involved in partner choice decisions. Drawing on research from

a range of fields, we showed that these partner choice decisions are supported by a

suite of identifiable mechanisms focused on individuals’ social normative behavior.

Partner choice is sensitive to signals of partners’ generosity and fairness, their

cooperative disposition, their moral decision-making, and their intentions. We then

reviewed evidence that sensitivity to these factors may be modulated by social context

and highlighted some initial evidence that the process underlying partner choice may be

unique. Finally, we provided evidence that partner choice is observed across a range of

animal species, emerges early in human development and has important consequences

for those who are rejected as partners. By aggregating and summarizing existing work

on the psychology underlying partner choice, we hope to spur new research on

remaining questions on this emerging topic. A better understanding of the psychology

of partner choice will deeply inform theoretical accounts of the forces governing

cooperation and social behavior in humans.

Acknowledgements

We thank Nichola Raihani for feedback on earlier versions of this manuscript.

 36
Partner choice

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