Expression unleashed

The evolutionary & cognitive foundations of human communication

VERSION September 2021

Christophe Heintz & Thom Scott-Phillips

Department of Cognitive Science, Central European University, Budapest, Hungary

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Abstract

Human expression is open-ended, versatile and diverse, ranging from ordinary language use

to painting, from exaggerated displays of affection to micro-movements that aid coordina-

tion. Here we present and defend the claim that this expressive diversity is united by an in-

terrelated suite of cognitive capacities, the evolved functions of which are the expression and

recognition of informative intentions. We describe how evolutionary dynamics leash com-

munication to narrow domains of statistical mutual benefit, and how they are unleashed in

humans. The relevant cognitive capacities are adaptations to living in a partner choice social

ecology; and they are, correspondingly, part of the ordinarily developing human cognitive

phenotype, emerging early and reliably in ontogeny. In other words, we identify distinctive

features of humans’ social ecology to explain how and why humans evolved the capacities

that, in turn, lead to massive diversity and open-endedness in means and modes of expres-

sion. We make cross-species comparisons, describe how the relevant capacities can evolve in

a gradual manner, and survey how unleashed expression facilitates not only language use but

novel behaviour in many other domains too, focusing on the examples of joint action, teach-

ing, punishment and art, all of which are ubiquitous in human societies but relatively rare in

other species. We aim to help reorient cognitive pragmatics, as a phenomenon that is not a

supplement to linguistic communication and on the periphery of language science, but

rather the foundation of the many of the most distinctive features of human behaviour, soci-

ety and culture.

Keywords: communication; language; pragmatics; social cognition; language evolution;

cognitive anthropology; comparative cognition; philosophy of language; evolutionary psy-

chology

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1. Introduction

Why is human expression so rich and multifaceted? Living things  communicate in a great

variety of ways, from the quorum sensing of bacteria, to songbirds, to the gestural and vocal

communication of primates, but humans are expressive in ways and to an extent that is

clearly distinctive. There is language use of course, but also points, nods, winks and other

behaviours that although not linguistic are still conventionalised; and also many ad hoc, im-

provised behaviours, such as a small hand gesture used to visually park a topic of ongoing

conversation, subtle body movements that connect dance partners, and the open-ended ex-

pressiveness of music, painting and other art forms (Green, 2007). It is likely that humans

are not naturally sensitive to many forms of interaction in non-humans, and as such there is

much still to discover from a comparative perspective; but at the same time, non-human ex-

pression is, on all available evidence, limited to finite domains. Bees, for instance, communi-

cate about the location of flowers and the quantity of their nectar but, apparently, little or

nothing else. The gestural communication of non-human great apes is more diverse and flex-

ible than most other cases (Cartmill & Byrne, 2010; Fröhlich & Hobaiter, 2018; Graham et

al., 2018), but its scope is still clearly limited relative to humans.

In explaining the open-endedness of human communication, many researchers em-

phasise the combinatorial and generative quality of natural language: the fact that individual

constituent parts can be recombined in many ways, making infinite use of finite means. An

emphasis on combinations and codes is, moreover, sometimes linked with particular as-

sumptions about evolution: that as complex systems of words, rules and combinations, nat-

ural languages are enrichments of the communication systems of other species (e.g. Progov-

ac, 2015; Sterelny, 2017; Leroux & Townsend, 2020). This picture is attractive because it de-

scribes human and non-human communication systems in terms that appear continuous.

However, as a broad explanation of human expression, this focus on the evolution of com-

binatorics faces a number of fundamental problems, of which we here highlight two. First, it

says very little about quasi- and non-linguistic means of communication and expression.

Second, the evolutionary story of complex combinations evolving from more simple ones

does not address the “central problem” (Maynard Smith & Harper, 2003, p.v) for the evolu-

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tion of communication, viz. stability in the face of incentives to deceive. This is not an incid-

ental critique. On the contrary, the problem of evolutionary stability is, we shall shortly ar-

gue, fundamental, for it holds expression on a leash, keeping it constrained to narrow do-

mains of statistical mutual benefit (§2). In consequence, explaining evolutionary stability

and explaining expressive versatility are deeply interlinked problems. One cannot be re-

solved without the other.

We present and develop an alternative explanation of the zoological distinctiveness

and open-ended richness of human expression, based not on combinatorics but cognitive

pragmatics. That is, we describe the evolution of distinctly human means of communica-

tion—sometimes called an “interactional engine” (Levinson, 2006, p.39)—as the evolution of

mechanisms of social cognition targeted at navigating distinctive features of the human so-

cial ecology; and we specify how these cognitive mechanisms in turn unleash expression.

More specifically, we argue that expression can be unleashed in partner choice social ecolo-

gies where it is simultaneously adaptive (i) for interpretative inferences to be predicated on

spontaneous prior assumptions that communicators are cooperative, and (ii) for expressive

behaviour to exploit this assumption. Natural languages, in all their combinatorial richness,

are a means by which we exploit unleashed expression, rather than being the source of un-

leashed expression. If we are right about this, then our account provides an overtly adapta-

tionist answer to the ‘Why humans?’ question about language origins, that is clearly different

to prominent biolinguistic approaches (e.g. Berwick & Chomsky, 2016; 2017).

These contributions substantially enrich previous insights that human communica-

tion is evolutionarily grounded in cooperative social ecologies, and that key cognitive pro-

cesses involved in communication derive, evolutionarily, from non-communicative aspects

of social cognition shared with other primate species (e.g. Sperber, 2000; Hurford, 2007;

Tomasello, 2008; Fitch et al., 2010; Frith & Frith, 2010; Arbib, 2012; Sterelny, 2012; Levin-

son & Holler, 2014; Scott-Phillips, 2015; Moore, 2017; Seyfarth & Cheney, 2018; Hrdy &

Burkart, 2020; Johansson, 2021). Going beyond this point of agreement, we provide an es-

pecially focused description of the relevant cognitive capacities, grounded in a precise theory

of cognitive pragmatics. In other words, we describe the computational tasks the interaction-

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al engine must perform. We also describe how these cognitive capacities are employed for

use in a wide range of domains, extending far beyond how communication is ordinarily con-

strued. In doing this we do not make recourse to notions such as second-person or ‘we’ in-

tentionality, which are, in our view, not descriptions of cognitive processes, but phenomena

that are themselves in need of explanation.

The structure of the paper is illustrated in Figure 1. As it suggests, we elaborate on the

problem of leashed expression in the next section, and we explain how it is resolved (§5)

after we have provided a taxonomy of ways in which individuals can affect the minds of oth-

ers (§3), and make adaptive inferences based on others’ behaviour (§4).

Figure 1: Structure of the article. We first elaborate on how the key problem of
evolutionary stability leashes expression (§2). We then describe a series of suc-
cessively embedded, graded subsets of the different ways in which individuals
might affect the minds of others (§3), and make adaptive inferences based on the
behaviour of others (§4). We bring these two sides together by describing how the
innermost subsets on each side of the equation combine to unleash expression
(§5). Three further sections elaborate and enrich this point in different ways. We
identify the social ecology in which the relevant cognitive mechanisms are mutu-
ally supportive and can hence gradually co-evolve (§6); we make relevant com-
parisons with other species, chimpanzees in particular (§7); and we describe how
the framework presented here unifies and provides new insights on otherwise di-
verse and disparate means of human expression (§8). As the figure suggests, the
arguments made in these later sections are mutually supportive but largely inde-
pendent of one another. To conclude, we summarise how this evolutionary and
cognitive perspective redefines the domain and goals of pragmatics as a scientific
discipline (§9).

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2. Expression leashed

We use ‘expression’ to describe any trait or behaviour whose function is to inform others.

This characterisation is solely functional in nature, and not mechanistic. In this way, it is

sufficiently broad to be inclusive of whatever means this function might be achieved, includ-

ing, for instance, emotional expression; but also sufficiently narrow so as not to include any

and all cases of information flow, regardless of function. In this section we describe how ex-

pression in this functional sense is leashed rather than freely open-ended. In later sections

we focus on one specific manifestation of expression, namely behaviours that result from in-

formative intentions.1

Crucially, an organism’s expressive range is limited by the fact that only a specific and

finite range of stimuli will actually generate a psychological reaction in other organisms. Af-

ter all, organisms attend only to a limited subset of other organisms’ behaviour, and will take

from this only a limited range of information. This makes the expression of, for instance, fu-

ture events, or the location of far away food sources, effectively impossible without mecha-

nisms of interpretation that complement mechanisms of production. Consider bee dance: it

would not—and could not—express the location of pollen if other bees had no mechanisms

specifically dedicated to exactly that, i.e. interpreting dances as indicators of the location of

pollen.

Communication is thus, in our terms, a subset of expression. Whereas expression is

the production of stimuli the function of which is to generate a psychological reaction, com-

munication involves the production of stimuli the function of which is to generate a reaction

by the particular means of stimulating complementary mechanisms of interpretation (Scott-

Phillips et al., 2012). By ‘complementary’, we mean that each mechanism can perform its

function only in conditions when the other mechanism is in place. Bee dance, for instance, is

communication because it generates a reaction by means of stimulating complementary

mechanisms of interpretation; and those mechanisms of interpretation can only perform

1 From the perspective of evolutionary theory we are defining ‘expression’ at the ultimate/functional
level of analysis, with the goal to focus, in later sections, on its proximate manifestation in humans,
and potentially other species (for elaboration of the ultimate-proximate distinction see Scott-Phillips
et al., 2011).

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their function (to learn about pollen) if bee dance actually takes place. In contrast, frighten-

ing behaviour can be expressive but not communicative: it can generate a reaction, but not

necessarily by stimulating complementary mechanisms of interpretation.

Correspondingly, we characterise communicative stimuli as those that generate a re-

action by triggering complementary mechanisms of interpretation; and we characterise non-

communicative stimuli as stimuli that generate a reaction by triggering other mechanisms,

with different functions. Expression thus involves producing either communicative or non-

communicative stimuli to change others’ psychological states, while communication involves

producing specifically communicative stimuli for the same function. As such, the evolution-

ary emergence of mechanisms of interpretation in audiences enrich expression because they

complement mechanisms of production, producing communication systems as a result. (This

is not the same as unleashing expression: see below.) This distinction between expression

and communication is useful and important because it frames the important questions in the

right way. First, it raises the question of whether or not an expressive function is met com-

municatively i.e. by means of triggering the audience’s dedicated interpretative capacities.

Second, it raises the questions of how and why expressive and interpretative capacities might

co-evolve.

Crucially, however, the evolutionary emergence of communication systems does not

usually unleash expression, because communication systems are tied to domains of statistic-

al mutual benefit (Maynard Smith & Harper, 2003; Searcy & Nowicki, 2005; inter alia). This

is because the interdependence of mechanisms of production and mechanisms of interpreta-

tion means that for communication to be evolutionarily stable, it must be beneficial, on aver-

age, to both communicator and audience. This does not imply that communication is always

of mutual benefit, or that deception never occurs. However, it does imply that communica-

tion must be sufficiently beneficial, sufficiently often, for both parties, otherwise it would col-

lapse, because on one side or the other there would be selection against the interaction. Ex-

plaining why this does not happen is the central theoretical issue in animal signalling theory

(ibid.).

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 In some cases this mutual benefit derives from genetic relatedness, such as with ant

pheromones or bee dance. In other cases it derives from direct fitness effects on communica-

tor and audience. For instance, the pattern on the wings of many poisonous butterfly species

communicates their non-palatability to potential predators, because this is beneficial to both

the butterfly and the potential predator. The function of the pattern is to inform the potential

predator, and the function of the predator’s reaction is to avoid feeding on such butterflies.

Clearly deception can occur: other species of butterfly can be and have been selected to mim-

ic the focal species, even if they (the other species) are not themselves poisonous. Missing

out on the possibility of preying on the mimic species is an opportunity cost for the predator,

but this is outweighed by the benefits of avoiding poisonous butterflies. If this were not so—

if, in other words, the same pattern is used by so many actually palatable organisms that the

predator’s opportunity costs outweigh the risks of eating unpalatable prey—then the com-

munication system would collapse. The predator will, under these circumstances, not evolve

any mechanism for attending to the signal in the first place; or, if they already have such a

mechanism, it will be selected against (Scott-Phillips et al., 2012). These and other dynamics,

such as those associated with the differential costs of signalling (see e.g. Lachmann et al.,

2001), leash communication to relatively narrow domains of statistical mutual benefit.

Human communication appears to be in flagrant violation of this limitation. Its range

is certainly not restricted to any particular topic: humans can communicate about potentially

anything. Moreover, we commonly and ordinarily take people at their word, even for state-

ments that can have immediate and serious fitness consequences, such as, for instance, a

doctor’s medical diagnosis and prescriptions. Moreover, humans frequently communicate

about phenomena for which no directly observable evidence could ever be provided, such as

statements about past or future events. Given this vast expressive range, audiences should be

massively vulnerable to misinformation and deception; and what should follow, on ordinary

evolutionary logic, is the collapse of the communication system itself (see above). Yet this

does not happen. Explaining why this is so is, we believe, not only necessary for explaining

the evolution and the expressive richness of human communication, but fundamental.

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 Let us summarise. Expression can be enriched when supplemented with complemen-

tary mechanisms for interpretation, generating a communication system. This does not how-

ever unleash expression: this does not make expression open-ended. On the contrary, stan-

dard evolutionary considerations tell us that communication systems are still expressively

limited because they are only evolutionarily stable when there is little gain (in the aggregate)

to deception. Yet as a truly open-ended means of expression, human communication does

not seem to be restricted in the same way. How is this paradox resolved? Why would it be

adaptive for humans to have the sort of interpretative mechanisms they do, given the central

evolutionary problem of stability? The assertion that human expressivity is enabled by com-

binatoriality offers no answer to this problem. We will provide an answer (§§3-7) by relating

the evolution of human communication to the evolution of cognitive mechanisms that specif-

ically function to allow humans to make the most informative use of social interaction. These

mechanisms are, we shall argue, both a consequence and a cause of a partner choice social

ecology.

The study of cognitive mechanisms for human expression is traditionally the domain

of cognitive pragmatics: the study of the capacity of mind that facilitates human communic-

ative competence. The relevant literature has its most important origins in the work of philo-

sopher Paul Grice (1957; 1975; 1989). Grice was particularly concerned with meaning, and

his key originality was to approach it as a primarily psychological phenomenon, and a lin-

guistic phenomenon only derivatively. In particular he developed the idea that intentions

might play a key role in determining meaning itself. This work provides the foundations on

which a cognitive theory of communication and expression can be built, and since Grice an

extensive literature has developed this approach in various ways, including from evolution-

ary and developmental perspectives (e.g. Bach & Harnish, 1979; Sperber & Wilson,

1986/1995; Clark, 1996; Levinson, 2000; Tomasello, 2008; Csibra & Gergely, 2009; Wilson

& Sperber, 2012; Scott-Phillips, 2015; Moore, 2017; inter alia). Specific approaches differ

from one another in some of the detail, but all agree that the expression, recognition and epi-

stemic evaluation of intentions together play a foundational role. In what follows we adopt

and enrich the post-Gricean approach commonly known as Relevance Theory (Sperber &

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Wilson, 1986/1995; Carston, 2002; Wilson & Sperber, 2012; Clark, 2013; Padilla Cruz, 2016;

see also the Relevance Theory Online Bibliographic Service), but our analysis could poten-

tially be adapted to fit with more classically Gricean or neo-Gricean approaches (for focused

comparisons see e.g. Carston, 2004; Sperber & Wilson, 2005).

3. Graded forms of manipulative intention

In a pair of seminal papers, Krebs and Dawkins characterised animal communication as an

arms race between means of affecting the minds and behaviours of other organisms—la-

belled ‘manipulation’—and means of reacting in an adaptive way to the behaviour of others

(1978; 1984; see also Guildford & Dawkins, 1991). Manipulation is a broad term, to include,

for instance, the handling of objects in a goal directed way. Krebs and Dawkins’ insight was

to think of informing others as a means of ‘manipulating’ their behaviour. 2 In this section we

relate this teleological characterisation to the specific, cognitive mechanisms by which it is

achieved in humans. In the next section we do the same for the audience side.

The specific mechanisms by which communication is achieved in the natural world

are many and varied. They might be, for instance, physiological, as in the case of, say, butter-

fly wing patterns; or chemical, as in the case of, say, quorum sensing (Diggle et al., 2007).

Our focus is on cognitive means, and more specifically the expression and recognition of in-

tentions. We distinguish three embedded categories of manipulative intention, elaborating

on each with examples (Figure 2). Each of the following subsections begins with a concise

statement of each category, followed by examples and elaboration.

2 Despite the pejorative tone of ‘manipulation’ in everyday language, here it is used just to describe
actions on others, regardless of whether those actions are pro- or anti-social.

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 Figure 2: Embedded subsets of manipulative intention. Aligning with our func-
tional characterisation of expression (§2), here we differentiate expression as ac-
tion based on informative intention, from ostensive communication, which is ac-
tion based on communicative intentions. This implies that, for instance, conspic-
uous consumption can express wealth, but does not communicate it.

3.1. Intentional action on others

The broadest set are behaviours that are intentional and manipulative. For instance, experi-

mental studies show how orangutan mothers will, if necessary, use their offspring as physical

tools (Völter et al., 2015). Because of their small size, infants can reach food in locations that

mothers cannot reach, so mothers can (and do) use them to reach the food, with the mother

then consuming the food herself.

3.2. Action based on informative intention

In the second set are behaviours that are intended to inform others, and which can do so

without overtly bringing attention to the informative intention itself. For instance, an indi-

vidual might dress in a smart and conservative way, as a means to suggest to others compet-

ence and professionalism, yet without bringing excessive attention to oneself. Conspicuous

consumption is intended to provide evidence of wealth and other markers of status, but

without necessarily advertising this intent in a formally overt way. In the presence of others

we might adopt a bodily posture that suggests, say, social ease and competence, and while

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this can be done in an overt or otherwise exaggerated way, it need not be. More generally,

impression management—individuals presenting themselves in ways intended, subcon-

sciously or otherwise, to generate and maintain a positive image in the eyes of others, but

without overtly bringing attention to this informative intent—is a common feature of human

social life.3

Such behaviour can generate a degree of shared knowledge about the actor’s informa-

tive intent. In other words, it may be salient that the actor has and is acting on an informa-

tive intention. That said, this need not necessarily be the case. In fact in some cases the actor

might have informative intentions but also have strategic motives to actively keep those in-

tentions hidden or at least deniable (called ‘hidden authorship’; see e.g. Grosse et al., 2013).

A criminal who plants misleading cues in a crime scene has an informative intention and is

acting on it, but simultaneously hiding that intention. A dinner guest who wishes to have

more wine but, recognizing it would be impolite to ask, might wait until her hosts’ attention

is elsewhere and then move her empty glass to a conspicuous location where it will, in due

course, be noticed. Many public acts of generosity fall within this category also: generous in-

dividuals want to be seen as generous, so they gain a positive social reputation, but often

they do not want their acts of generosity to be seen as simply attempts to gain a positive so-

cial reputation, because that would immediately undermine their purpose (Frank, 1988;

Berman et al., 2015; Hoffman et al., 2015; Karabegovic & Heintz, under review; see also §8.3

on punishment).

In all these cases, agents satisfy their informative intentions simply by means of

providing non-communicative, or ‘direct’, evidence for what they want to convey.4 If, for in-

3 A further example is ‘phatic communication’: pleasantries and other means of communication that
serve plainly social functions, and which mainly consist of expressing a willingness to engage and to
observe local conventions of politeness. Phatic communication often uses languages and other means
of communication from within §3.3, even though the informative intentions that are expressed in
phatic communication bear mostly on the willingness to engage, in and of itself, rather than on what is
linguistically expressed.
4 Precisely, we say that X is non-communicative evidence for Y if and only if X (might) generate, in the
audience, the inference that Y without the observer necessarily computing that the informer has the
informative intention that Y. Note also that this notion of non-communicative evidence is similar to
Grice’s ‘natural meaning’ (1957). Interestingly, many languages have grammatical evidential markers
for information acquired in this non-communicative (‘direct’) way.

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stance, Amy puts three apples on the table with the intention of informing Barry that there

are three apples, she is providing non-communicative evidence for the presence of the three

apples. She can, moreover, do this without any indication that she actually has an informat-

ive intention: she can just place the apples on the table, without drawing any particular at-

tention to the fact that she is doing this. Looking comparatively, we take it as plausible that

non-human primates, and possibly some other species, have informative intentions, and sat-

isfy them by providing non-communicative evidence (see e.g. Genty & Zuberbühler, 2014 for

a plausible example; Zuberbühler, 2018 for a review; and Bar-On, 2013 for discussion). The

key comparative questions are, in our view, whether any non-human species act in the ways

described in the next subsection, in which we consider cases where the communicator

provides evidence for the informative intention itself Such cases are sometimes called

overtly intentional (because they involve making intentions overt) or, more simply, ostens-

ive.5

3.3. Action based on communicative intention

In this third set are behaviours performed not only with an intention to inform an audience,

as above, but, more than this, to make the actor’s informative intention mutually known.

This is achieved, as we said above, by communicators providing non-communicative evi-

dence for their informative intention itself.

To see the difference between this set and the one above, consider two possible ways

in which Mary might satisfy her intent that Peter be informed that some berries are edible

(see also Sperber, 2000; Wharton, 2006; Scott-Phillips, 2015). One way Mary might do this

is to simply eat the berries in Peter’s company (without Mary bringing any particular atten-

tion to the fact that she is doing this). In this case Mary has an informative intention which

she acts on by providing some evidence that the berries are edible, but without giving any

5 Regarding ‘ostension’, the word is used in slightly more broad or more narrow ways in different lit-
eratures. Within pragmatics the word was first used narrowly, for the actions described in §3.3 i.e.
making an informative intention manifest (Sperber & Wilson, 1986/1995). Since then sizable literat-
ures have developed studying ostension from many perspectives including development, evolution
and comparative cognition, and here the term is sometimes used more broadly, corresponding
roughly to the actions described in §3.2 (see e.g. Gómez, 1996; Tomasello, 2008; Csibra, 2010; Moore,
2017; Sperber, 2019).

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overt evidence that she is acting on an informative intention. As such she relies on Peter

simply attending to her behaviour and drawing the inference that the berries are edible. In

this case, Mary’s behaviour belongs in the second embedded subset (§3.2). There is however

an alternative. Mary might not eat the berries at all, but instead mime eating them, perhaps

with exaggerated movements and while tapping her tummy. Here she has the same informa-

tive intention, but provides evidence only about the intention itself. She does not eat the

berries, after all. She provides only communicative evidence of their edibility.

The most salient and important special case of overtly intentional behaviour is, of

course, the use of conventional symbols, especially, but not only, in the context of language

use (§8.5). Grice’s work on meaning was focused on these cases, and his crucial insight was

that the sort of actions we are describing in this section—providing evidence about informat-

ive intentions—is what generates meaning. As he put it, “‘A meant something by x’ is

(roughly) equivalent to ‘A intended the utterance of x to produce some effect in an audience

by means of the recognition of this intention’” (1957, p.385, italics added; the intention re-

ferred to here is best understood, in our analysis, as an informative intention).

That said, overtly intentional behaviour is also viable in cases where no conventions

are used to communicate. After all, almost any behaviour that humans can perform, they can

perform in an overtly intentional way. Sometimes we eat food, and sometimes we eat food in

an overt, exaggerated or otherwise ostensive way, to express to others that the food is tasty,

revolting, generous, or fancy. Sometimes we blink, and sometimes we blink with microscopic

exaggeration, such as with a slight delay in re-opening the eyes, to express, say, ironic sur-

prise. Such deviations from otherwise non-communicative behaviour have been experimen-

tally isolated in a number of studies, in both production and comprehension (e.g. Newman-

Norland et al., 2009; Scott-Phillips et al., 2009; McEllin et al., 2018a; Royka et al., 2018).

Again, like language use, such behaviour is Gricean. It provides evidence about informative

intentions.

We have so far presented the distinction between the three subsets in this section as

categorical, but they are in fact graded and continuous. That is, different means of manipula-

tion can, we suggest, vary in the extent to which the actor makes her informative intention

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manifest. Grice’s characterisation of meaning, quoted above, describes one end of this con-

tinuum; the cases described in §3.1 represent the other end; and in between are many cases

where communicators make informative intentions partially manifest. We shall return to

this graded quality in §8, where we shall suggest that it helps to generate the massive diversi-

ty of human expression. (For related but different continua see Wharton, 2009; Sperber &

Wilson, 2015; Duranti, 2015, p.290.)

In any case, informing others via the expression of informative intentions—common-

ly called ‘communicating’—has a distinctive property that is not present in the other subsets

described above. Crucially, the communicator is freed from the constraint of providing non-

communicative evidence (see also §5). Returning to the example above, Mary makes eye con-

tact with Peter and mimes eating berries in an exaggerated way, with the informative intent

that Peter believes the berries are edible, and in doing so she provides only communicative—

and hence indirect—evidence that the berries are edible.

However, for this potential freedom of expression to be realised two related issues

must be addressed. First, freedom from the constraint of having to provide non-communic-

ative (‘direct’) evidence depends on audiences’ complementary abilities to infer informative

intentions, more-or-less accurately, on the basis just of whatever other, communicative evid-

ence communicators are able to provide (see §2 on the importance of complementary mech-

anisms). Second, expressive freedom also provides communicators with the opportunity to

deceive, leaving the system clearly prone to evolutionary instability (§2). To discuss how

these issues are resolved, we turn now to the audience side.

4. Graded forms of social vigilance

All animal species have evolved adaptive reactions to the presence and behaviour of others.

As with manipulation (§3), we define these adaptive reactions functionally, recognising that

the specific mechanisms can be many and varied.6 For example, a chameleon’s adaptive re-

6 Note that our terminology here differs from Krebs and Dawkins, who used ‘mindreading’ instead of
‘adaptive reaction’ (1984). We have deviated from this usage because ‘mindreading’ is also widely used
in the social cognition literature to describe mechanisms rather than function i.e. at the proximate
rather than the ultimate level.

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action to the presence of potential predators is (we assume) largely physiological. We focus

here on cognitive means of adaptive reaction targeted at the behaviour of conspecifics. We

call these cognitive means ‘social vigilance’ (Heintz et al., 2017). Again we distinguish three

graded and embedded subsets (Figure 3), and we describe their relationship to the various

categories of intentional manipulation described in the previous section.

Figure 3: Embedded subsets of social vigilance. Importantly, the inferences con-

tained in the innermost subset include interpretative inferences derived on the
basis of a presumption of relevance, and also inferences about the epistemic value
of what is communicated (i.e. inferences derived on the basis of epistemic vigil-
ance; see §5).

4.1. Inferences about others’ intentions

The first embedded subset includes inferences based on the capacity to anticipate and re-

spond adaptively to the intentional action of others. Humans do this routinely, of course.

Others’ intentions are relevant to us, such as when we decide whether to avoid or engage

with them as friends, rivals, or indeed any social relationship. The capacity to behave in ways

that take account of other individuals’ intentions has also been experimentally documented

in many studies with non-human great apes (Emery & Clayton, 2009; Call & Tomasello,

2010; Andrews, 2017; Bettle & Rosati, 2021). In one such experiment, chimpanzees are given

the opportunity to take a piece of food from a bucket, the location of which is either known

or not known by a dominant conspecific. The key finding is that the subordinate chimpanzee

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is more likely to take the food if the location is unknown to the dominant (Hare et al., 2011).

The details of exactly which intentions and other mental states great apes attribute to others

remains a topic of active study; but in any case many experiments show that chimpanzees

are able, in some contexts at least, to adaptively modulate their behaviour in view of what the

conspecific is most likely to do, and what effects this might have on the focal individual. Sim-

ilar modulations have been documented in many non-primate species. Grey squirrels, for

instance, have been shown to modulate their caching behavior as a function of the presence

of onlookers e.g. moving a cache when the onlooker leaves (Leaver et al., 2007); and ravens

have been shown to guard their caches against discovery, taking into account other ravens’

possible knowledge of the cache (Bugnyar et al., 2016).7

4.2. Inferences about others’ informative intentions

In the second embedded subset are inferences about others’ informative intentions. The pos-

sibility of such inferences, and their nature, depends to a significant extent on whether the

informative intentions in question have been made overt (§3.3) or not (§3.2) by the actor, or

communicator.

If an informative intention is not overt and audiences do not recognise it, then what

is perceived is simply instrumental action, which like any behaviour might or might not be

relevant to the observer. If, alternatively, an informative intention is not overt but is recog-

nised nevertheless, audiences might take account of this absence of overtness, and the possi-

ble reasons for it, in their interpretation. Suppose, for instance, Claire leaves Dwight’s keys

on the table, with the informative intention that by virtue of seeing them Dwight does not

forget the keys when he goes to work. Dwight might simply see the keys and thus remember

to take them, not ever recognising Claire’s informative intention, which was after all not

7
Throughout this article, we have avoided using the terms ‘mindreading’ and ‘theory of mind’, be-
cause they are so widely contested. Nevertheless, a reviewer asked us to state our view and we are
happy to do so: we are mindreading ‘deflationists’ (e.g. Jacob & Scott-Phillips, 2020). We believe that
these notions are best used in a minimal way, to refer just to the spontaneous recognition of mental
states; and as such we believe that mindreading is much more akin to, say, visual cognition than to
conscious ‘thinking’. We think that mindreading in this deflationary sense is likely to be phylogeneti-
cally widespread, but enriched in various specialised ways in different species. The various cognitive
processes we describe in this article are just such specialised versions of mindreading in this defla-
tionary sense, falling within the broad functional category of social vigilance.

page 17 of 51
overt. Alternatively, Dwight might recognise that Claire had an informative intention even

though she did not make this overt, and he might hence infer that she thinks he is absent

minded but she does not want to embarrass him by saying so explicitly. In any case, humans

commonly act with informative intentions that they do not make overt, but which are some-

times recognised nevertheless. §8.1 on coordination smoothers and §8.3 on expressive pun-

ishment discuss some specific examples.

Actors attempting to make an informative intention overt have a communicative in-

tention (by definition: see §3.3). If the informative intention is nevertheless not recognised,

such as when, for instance, a raised hand, intended as a request to ask a question, is inter-

preted as a mere stretch of the arm, then that is a simple failure of communication. If, in

contrast, an overt informative intention is recognised as being overt, such as when a raised

hand intended as a request to ask a question is indeed recognised as a request to ask a ques-

tion—if, in other words, the audience recognises that the actor has not only an informative

intention but a communicative intention—then the audience is warranted in making an in-

ferentially powerful presumption about the behaviour. Specifically, the audience is warrant-

ed in presuming that the behaviour is the most effective one the communicator could pro-

duce, given the communicators’ goals, abilities, and the constraints acting on them. This in-

sight is central to Relevance Theory (Sperber & Wilson, 1986/1995; 2002; 2005), where it is

called the communicative principle of relevance. In the next section we describe in more de-

tail the nature of the inferential process that is triggered, in audiences, by this recognition of

communicative intent.

4.3. Inferences about others’ communicative intentions

In §3.3 we described how communicators sometimes express their informative intentions

overtly, with the goal to make their informative intention mutually known. Here we discuss

how such stimuli are interpreted.

Language use is a paradigmatic example, and one of Grice’s pioneering insights was

that the interpretation of utterances is guided by prior expectations about the cooperative

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intent of communicators (mirroring his characterisation of linguistic meaning: see §3.3). 8

Further developments in cognitive pragmatics have specified and debated the nature of these

expectations in more detail. Relevance Theory, for instance, describes these expectations in

terms of a single assumption, that ostensively presented stimuli are optimally relevant for

the intended audience, given the speakers’ goals, abilities, and the constraints acting on

them. Or, in other words: audiences have a strong positive prior expectation that overtly in-

tentional behaviour is cooperative; and this prior expectation of cooperativeness in turn li-

censes a presumption that informative intentions are worth paying attention to i.e. are op-

timally relevant. Here and elsewhere, “optimally relevant” means, more precisely, that com-

municators strive to optimise the trade-off between cognitive effects and processing effort,

subject to goals, abilities and constraints.

Here is an example. Amy and Barry are drinking in a bar. Amy’s glass, which is visible

to both her and Barry, is empty. This fact is on its own unremarkable. Suppose now that Amy

picks up the glass and gently waves it in front of Barry. Why would she do this? What could it

possibly ‘mean’, and how could Barry know? The Relevance Theory answer is that Amy’s be-

haviour triggers in Barry a spontaneous process of interpretation, governed by a cooperative

assumption that Amy’s behaviour is the most optimally relevant behaviour she could per-

form given her goals and the circumstances. The key point here, which features in some form

in all Gricean approaches, is that only with this cooperative assumption can Barry converge

on the conclusion that Amy’s behaviour is a suggestion that they stay for another drink. Oth-

erwise, without this assumption, Amy’s behaviour is simply mysterious. Many experimental

studies have shown how prospective audiences interpret communicative behaviours under

presumptions of optimal relevance (e.g. van der Henst et al., 2002; Gibbs & Bryant, 2008;

inter alia).

8 Note that the notion of cooperation used in the Gricean framework is not the same as that used in
standard evolutionary theory. The evolutionary notion is about evolutionary function, and describes
any behaviour that has a positive effect on another organism’s inclusive fitness. The Gricean notion is
about the assumptions that audiences make about the intent of the communicator. While these two
notions can align with one another, and are sometimes conflated, they are different levels of analysis
and not the same.

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 This is very similar to how other specialised cognitive mechanisms ‘embody’ know-

ledge about the nature of objects, magnitudes, species, and other basic, fundamental features

of the human evolutionary ecology (Carey, 2009; Spelke & Kinzler, 2007). In all these cases,

items perceived as being of a particular type (an object, a magnitude, and so on) trigger spe-

cific assumptions about the nature of that item. For example, items perceived as physical ob-

jects trigger assumptions that the item is physically cohesive, bonded, rigid, and cannot be

acted on at a distance (Spelke, 1990). In the present case, behaviour perceived as ostensive

triggers an assumption that the behaviour is optimally relevant for the audience, given the

communicator’s goals, abilities, affordances and constraints. This allows a specialised, ‘satis-

ficing’ process of interpretation to then derive the communicator’s intended meaning (Sper-

ber & Wilson, 2002; see also e.g. Ferreira & Patson, 2007 on ‘good enough’ approaches to

linguistic comprehension). This process is, moreover, spontaneous and largely unconscious,

such that we cannot ‘choose’ not to perform it even if we wish to. Consider film spoilers, for

instance: our desire to not recover the meaning of what is said does not and cannot suspend

the interpretive process. Again, this is akin to the recognition of objects, magnitudes and so

on, all of which we recognise and process in spontaneous and unconscious ways. We cannot

‘un-see’ objects, just as we cannot ‘un-understand’ spoilers. All in all, spontaneous interpret-

ation of ostensive stimuli is a functionally specialised form of social vigilance, targeted at the

specific phenomenon of others’ ostensive behaviour.

As with the other side of the equation (§3), we have so far presented the distinction

between the subsets in this section as categorical, but they are in fact graded and continuous.

Specifically, there is on the audience side variation in the extent to which recognition of the

actor’s informative intention contributes to correct interpretation i.e. to satisfying the in-

formative intention. Again, we shall return to these graded aspects in §8.

5. Unleashing expression, together

We can now summarise how a system of communication predicated on the expression and

recognition of informative intentions can unleash expression.

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 Crucially, the metarepresentational structure of ostensive communication generates a

‘virtual’ domain generality. Communicators provide evidence of their informative intentions,

which can in turn be about anything at all (see also Mercier & Sperber, 2009 on virtual do-

main generality in cognition more broadly). Consider again the example of Mary, who makes

eye contact with Peter and mimes eating berries in an exaggerated way. By doing so, she

provides evidence of her informative intention, that Peter understands that the berries are

edible; and this intention is informative about the actual edibility of the berries only in turn.

This metarepresentational structure makes the expressive domain of ostensive communica-

tion effectively open-ended (unleashed), even though the actual domain of the relevant cog-

nitive capacities is narrow and specific: it is just the communicator’s informative intentions.

This metarepresentational structure has two important consequences, which together

generate a further important corollary.

First, a metarepresentational structure is how ostensive communication can be ex-

pressively open-ended while still conforming to the central evolutionary constraint, that

communication systems are tied to narrow domains of statistical mutual benefit. In §2 we

summarised why, from an evolutionary perspective, all evolved communication systems

should be tied to narrow domains of statistical mutual benefit, and we observed that human

communication appears to be in flagrant violation of this constraint. Now we can state how

the paradox can be resolved. The actual domain of the cognitive capacities that underpin os-

tensive communication is indeed still restricted to a narrow domain of statistical mutual

benefit, namely the communicator’s own informative intentions. At the same time, the

metarepresentational structure generates an expressive domain that is truly open-ended.

Second, the metarepresentational structure of ostensive communication generates a

distinction between comprehension and acceptance. Comprehension is targeted at the in-

formative intention itself: to comprehend is to recognise the informative intention that an

individual has towards another (‘She wants me to believe that the berries are edible’). Ac-

ceptance, in contrast, is targeted at what the informative intention is about i.e. what is ‘virtu-

al’. To accept is to actually update one’s own beliefs in light of what has been communicated

(‘The berries are indeed edible’).

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 Together, these two consequences imply that audiences cannot actually gain from

communication unless they extend a degree of trust towards the communicator. The distinc-

tion between comprehension and acceptance, and the massive open-endedness of human

communication, together mean that audiences who do not extend a degree of trust towards

ostensive communicators would comprehend what others want to do to their minds, but

would never then actually update their beliefs in light of that knowledge. They would never

allow themselves to gain information in communication! Peter would understand that Mary

wants him to believe that the berries are edible, but Peter unless he extends some trust to-

wards her, he will never believe that the berries are actually edible. Of course, this trust must

be tentative and provisional, lest audiences be misinformed, but it must be extended in some

way, just for audiences to gain from communication in the first place.

In consequence, cognitive capacities for expressing and recognising informative in-

tentions must be complemented by further capacities that allow audiences to trust what is

communicated but in a vigilant way, possibly questioning the competence or the benevolence

of the communicator. Commonly known as epistemic vigilance, these cognitive capacities are

a specialised form of social vigilance, targeted at the assessment of informative intentions

(Sperber et al., 2010; Mercier, 2017). They also allow audiences to identify misleading com-

municators, and hence adjust the attention and trust they are willing to grant in the future.

In sum, then, the open-ended richness of human communication is achieved virtual-

ly, by a combination of cognitive capacities for the expression of informative intentions, cog-

nitive capacities for the recognition of informative intentions, and cognitive capacities of

epistemic vigilance, all of which are functionally tied to one another. Thus, to properly ex-

plain the expressive openness of human communication, what must be described is: (i) how

these cognitive capacities could all gradually co-evolve and be mutually supportive of one

another, such that they form a communication system; and (ii) the ecological reasons why

they have actually done so in humans. The next section addresses these questions.

6. The co-evolutionary ecology of human communication

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Many authors have observed how human communication must have co-evolved in a highly

social ecology, one way or another (§1). Here we identify which specific and distinctive as-

pects of human social ecologies can generate the co-evolution of the cognitive capacities de-

scribed in §3 and §4, such that fully enriched ostensive communication can become uniform

and stable in the population. We also provide precise description of how this co-evolution

could occur in a gradual manner.

Arguably the most distinctive feature of the human cognitive niche is that it is highly

social. Humans tend to live in social groups that are loosely-defined but long-lasting, and

comprised of both kin and non-kin. To a degree that surpasses that of other great apes, this

social ecology generates many opportunities for win-win cooperation, and risks of exploita-

tion. More broadly, human social ecologies involve an especially delicate balance of coopera-

tion and competition, with substantial evolutionary pressure for behaviours that make the

most of this mix (Noë & Hammerstein, 1995; Ferriere et al., 2002; Tomasello et al., 2012).

Individuals acting in their own adaptive self-interest seek out others (‘friends’, ‘colleagues’)

with whom to engage in mutually beneficial enterprises, and they behave in ways that in-

crease their chances of being chosen as a partner for joint enterprise (Barclay, 2013; Krems

et al., 2021; inter alia). Humans have hence evolved a number of cognitive capacities adap-

ted for this ecology, including moral dispositions, mechanisms of social vigilance that identi-

fy potential partners and opportunities for mutually beneficial interaction, an awareness of

potential opportunities to exploit others, a strong sensitivity to changes in one’s reputation,

and so on (see e.g. Origgi, 2004; 2005; Barrett et al., 2010; Delton & Robinson, 2012; Sper-

ber & Baumard, 2012; Baumard et al., 2013; Heintz et al., 2017; Curry et al., 2019; Engel-

mann & Tomasello, 2019; inter alia). These factors collectively constitute a ‘partner choice’

social ecology; or, possibly, an ecology of ‘self-domestication’. This means, minimally, that it

is advantageous to be selected as a partner for some joint enterprise (other than mating), and

page 23 of 51
that the selection of partners for joint enterprise is based on information about past actions.

Reputations are thus especially critical.9

In this partner choice ecology, the cognitive means of manipulation described in §3.3,

and the cognitive means of social vigilance described in §4.3, are each adaptive. Particularly

important is the role of social commitments. By providing evidence of their informative in-

tention, informers make themselves accountable to their audience, putting their reputation

at stake; and audiences can hence effectively assume the relevance of overtly expressed in-

formative intentions. This evolutionary dynamic is, incidentally, similar to that described by

partner choice approaches to fairness (André & Baumard, 2011; Debove et al., 2015; Barclay,

2013; 2016). In both cases, the adaptive value of maintaining one’s reputation in a partner

choice ecology constitutes a crucial selection pressure for psychological traits, which in turn

generates prosocial behaviour. In the case of ostensive communicators, prosocial behaviour

means being relevant.

The following five paragraphs elaborate this argument in more detail. They hence

provide an existence proof of how and why the cognitive capacities described in previous sec-

tions could have evolved in a gradual manner (for similar but different approaches see e.g.

Wharton, 2006; Cornell & Wharton, in press). Indeed, ‘lineage explanations’, in which

changes in the phenotype result from incremental changes, are an important constraint on

theories of cognitive evolution, especially co-evolutionary theories (Calcott, 2009).

Consider a social ecology with many, varied opportunities for win-win cooperation.

Here, informing others can be adaptive, because it can facilitate win-win cooperation or even

create new win-win opportunities. In particular, informing others can generate common

ground and hence facilitate many joint enterprises (such as, say, animal hunting, building

9
Three different notions of reputation can be distinguished. (1) An individual A can have a ‘reputa-
tion’ for X in the sense that another person believes X about A (this is sometimes called an ‘image
score’). (2) An individual A can have a ‘reputation’ for X in the sense that a community of others all
believe X about A. (3) An individual A can have a reputation for X in the sense that a community of
others all believe that they all believe X about A i.e. the notion that A is X is common ground. Our ar-
gument in this section turns just on notion (1) and (2). Clearly, the emergence of ostensive communic-
ation, which enables open-ended communication and hence gossip, will in turn facilitate the emer-
gence of (2) and (3), and can hence support cooperative phenomena such as indirect reciprocity (see
e.g. Nowak & Sigmund, 2005).

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shelter, maintaining a fire, alloparenting). Informing others can also have fitness advantages

by allowing the social transmission of opaque skills to cooperative partners and kin.

These potential adaptive benefits in turn mean that it is adaptive to recognise and

attend to others’ attempts to gain attention (§4.2). That said, attention is limited and thus

should be modulated depending on whether others’ attempts to inform are likely to be worth

the attention indeed i.e. are revealing of relevant information. Individuals should be socially

vigilant towards others’ informative intentions, evaluating whether, or to what extent, the

intentions are indeed cooperative. It will hence be in informers’ own interests to actually be

relevant for their audience, because those who intentionally attract attention but fail to do so

in ways that are useful (relevant) will, in time, incur costs to their reputation and lose their

capacity to manipulate their conspecifics’ attention. In other words, there will be selection

for behaviours that intentionally attract others’ attention only when it is likely to be worth-

while for the audience to indeed pay attention (see also Dessalles, 1998).

At this point, expression has not yet ‘gone Gricean’ (we adopt this useful expression

from Bar-On, 2013), and as such humans are not yet ‘language ready’. This is because in-

formative intentions are not intentionally made overt, and so expression is not yet predic-

ated on the systematic exploitation of the audience’s recognition of informative intentions

(Grice’s “by means of” clause: see §3.3). Expression is based just on behaviours that inform-

ers expect will be relevant to others; and ‘comprehension’ is based just on tentative assump-

tions that others’ expressive behaviour is likely to be relevant for the same reason. These

tendencies and dispositions do, however, constitute a new social ecology, and it is here that

‘going Gricean’ is adaptive.

Crucially, in this new social ecology—in which audiences might expect others’ infor-

mative behaviour to be relevant, and in which a reputation for being a good cooperator can

be gained and lost—it is adaptive to make manifest informative intentions themselves i.e.

make overt and common knowledge the intentions you have towards your audiences’ mind.

This is adaptive because, by making informative intentions manifest, informers effectively

offer a credible commitment that the overtly presented behaviour will indeed be relevant for

the audience; which in turn increases the probability that the informative intention will in-

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deed be satisfied. In other words: since the overt expression of an informative intention

makes that informative intention common knowledge; and since in a partner choice social

ecology there is the risk of developing a reputation for irrelevance and hence of losing the

possibility of influencing others’ minds; then communicators are effectively committed to

their behaviour being useful (relevant) for the audience. This in turn makes it adaptive for

audiences to simply presume—even if just tentatively at first—that the behaviour is indeed

relevant, and to interpret the behaviour in light of this presumption of cooperativeness.

The social ecology is now one in which the overt expression of informative intentions

effectively commits informers to being relevant to their audience, and in which informers

indeed abide by that commitment (see also Scott-Phillips, 2010; Sperber, 2013). In this new

ecology, it is adaptive for audiences to evolve two kinds of specialised cognitive disposition.

(1) High prior expectations that others’ communicative behaviour will be relevant. These will

eventually become the spontaneous presumptions of relevance described in §4.3. (2) Forms

of social vigilance that assess how, and to what extent, beliefs should be updated in view of

others’ informative intentions. These will eventually become the epistemic vigilance de-

scribed in §5. Now audiences have these two kinds of specialised cognitive disposition, indi-

viduals can inform simply “by means of” (as Grice put it) making their informative intentions

manifest. This is Gricean communication proper.

Let us summarise. What we are arguing is that outside a partner choice social ecolo-

gy, communication and expression are highly prone to irrelevance, deception and instability;

but within a partner choice social ecology there is selective advantage for behaviour that is

cooperative (statistically speaking at least), which in the context of communication means

relevant. Within this social ecology a gradual, cognitive co-evolution of specialised capacities

for ostensive communication is hence possible. As with other aspects of core cognition, these

capacities, which provide the foundations of human communication (§5), should become

part of the ordinarily developing cognitive phenotype, emerging at reliable and predictable

stages of ontogeny.

There is, correspondingly, abundant empirical evidence of this reliable and predict-

able cognitive development in humans, both in language use and communication more

page 26 of 51
broadly (e.g. Bates, 1979; Bloom, 2002; Goldin-Meadow, 2005; Tomasello, 2008; Bohn &

Frank, 2019; inter alia). In the next section we consider whether, or in what ways and to

what extent, the same cognitive capacities might be present in non-human primates.

7. Cross-species comparisons

Do any other species communicate ostensively, or in proto- or quasi-ostensive ways?

How would we know? These questions are worth asking because non-human species, great

apes and dogs in particular, sometimes appear to understand some human ostensive be-

haviour, at least in some specific contexts. Here we outline how this question can be ad-

dressed experimentally, we present an ecology-based explanation of key differences between

human and non-human great ape communication, and we reinterpret some key findings in

the comparative cognition of communication.

The key methodological challenge in studying ostensive communication from a com-

parative perspective is that ostension is ultimately a psychological construct, i.e. ostension is

not any particular behaviour, but rather any behaviour motivated by a particular cognitive

phenomenon, namely informative intentions (§3.3). This makes it impossible to fully isolate

behavioural characteristics. As such it will always be theoretically possible to reinterpret any

behavioural differences between experimental conditions in a non-mentalistic (‘killjoy’) way,

so the relevant cognitive capacities are not ascribed to the individual animal participants, or

species. One response to this methodological challenge has been to, effectively, abandon use

of the Gricean framework in the study of animal communication (e.g. Townsend et al., 2017).

In contrast, we suggest that experiments revealing the relevant intentions and interpretative

processes are still possible.

In particular, the hypothesis that non-human primates can be sensitive to ostension

qua ostension (i.e. sensitive to the expression of an informative intention) can be tested by

contrasting two scenarios in which the exact same ostensive behaviour prompts spontaneous

identification of different informative intentions, and hence different behavioural responses,

depending only on what is in the common ground. Real-world human communication is re-

plete with examples. Ordinary utterances such as, say, “It’s raining” can, even if produced in

page 27 of 51
exactly the same way in each case, be interpreted in wildly different ways depending only on

the present common ground (‘Take an umbrella’, ‘Even the whether can’t lighten my mood’,

‘Your parents won’t be visiting after all’). The same can be true of non-verbal means of com-

munication, such as points and nods, not to mention spontaneous and ad hoc gestures, and

indeed all human ostensive stimuli.10

We predict, tentatively, that if any non-human primates do reliably pass tasks of this

sort, it will be individuals with extensive experience of altruistic human caregivers, and not

those living the natural social ecologies of non-human primates. Non-human primate social

ecologies involve fewer and less frequent opportunities for interactions of mutual benefit.

That is not to say such opportunities are absent, but they are much less prevalent relative to

the human case, and in consequence the relevant selection pressures are not (as) present. At

the same time, non-human primates living in captive conditions, with human caregivers who

are more-or-less uniformly cooperative, could—perhaps—develop the relevant dispositions

and expectations ontogenetically (see e.g. Call, 2011 for discussion of how rearing conditions

can affect chimpanzee communication). In other words, a ‘proto’ presumption of relevance

could result from non-standard life history in non-human apes. If so, then we should expect

some recognition and interpretation of human ostensive behaviours qua ostensive in at least

some individuals, albeit in imperfect and happenstance ways. This prediction is of course

ultimately a matter for future empirical research, but it aligns with existing findings that

humans differ from other great apes in dispositions of trust and cooperation (see e.g. Moll &

Tomasello, 2007; Jaeggi et al., 2010; inter alia).

Here is an analogy that helps to articulate this difference between specialised compe-

tences that are part of the ordinarily developing phenotype (as ostensive communication is

in humans), and latent competencies that might be refined in the right ecology (as might be

the case for ostensive communication in chimpanzees and some other non-human

primates). Consider humans swinging from trees. Human bodies are not especially well-suit-

10Note that the spontaneity of responses is especially important for future experimental design, be-
cause for a compelling test the differential interpretations of a specific stimulus should not be learned
by training on that same specific stimulus. Infants have been shown to pass a third-person version of
such tasks i.e. a version in which the infant observes interaction between two other agents (Tauzin &
Gergely, 2018).

page 28 of 51
ed to this task. We lack the specialised biological apparatus of other primates and we do not

develop the relevant dispositions as an ordinary part of ontogeny. At the same time, there is

no absolute barrier. Some humans can swing from trees in some limited ways and to some

extent, and this basic ability can be refined and enhanced with training: in other words, in

the right ecology. What we are suggesting, tentatively, is that ostensive communication in

other primates, living in the right sort of social ecologies, might be similar: not impossible

and not wholly absent, but still unspecialised, disfluent, not a regular part of the environ-

ment, and not part of the ordinarily developing phenotype.

This perspective on non-human primate cognition can help make sense of otherwise

puzzling findings in the comparative psychology of communication. We highlight two exam-

ples in particular: performance in the ‘object choice task’, and the phenomenon of overimita-

tion.

First, in the object choice task a desirable object is shown to the participant, and then

placed in one of two boxes, or bins. The participant does not know which of the two boxes

contains the desirable object. The two boxes are placed either side of the experimenter, who

then points to the box with the desirable object. The participant is then free to open the

boxes. Many non-human primates ‘fail’ this task: in many studies non-human primates do

not choose the indicated box at levels greater than chance (see Clark et al., 2019 for a recent

review). We suggest that this occurs simply because the relevant cognitive processes em-

ployed by the audience are, in non-human primates, not ordinarily predicated on a presump-

tion of cooperation, which in the context of pointing means communicative relevance. Dogs,

in contrast, perform far better at the object choice task (see below), as do human infants.

Second, we consider ‘overimitation’, in which individuals copy actions demonstrated

to them, including in particular those that are perceivably causally irrelevant (e.g. tapping a

box before opening it, even when the tapping makes no difference to whether or how the box

is opened). Intriguingly, overimitation is only consistently observed in humans, including

children, and not in chimpanzees or bonobos (Horner & Whiten, 2005; Lyons et al., 2007;

Johnson et al., 2016; Clay & Tennie, 2018; Hoehl et al., 2019). This finding has prompted

speculation that overimitation derives from a cognitive adaptation for acquiring generic, cul-

page 29 of 51
tural knowledge (e.g. Nielsen & Tomasello, 2010; Chudek & Henrich, 2011; Gergely, 2013;

Legare & Nielsen, 2015). We suggest, in contrast, that overimitation is best explained as a by-

product of audience presumptions of relevance. Overimitation reliably occurs only when the

copied behaviour has been performed in an overtly intentional (i.e. ostensive) way (see e.g.

Király et al., 2013 for experimental demonstration). This triggers in the audience a spontan-

eous process of interpretation predicated on a presumption of optimal relevance (§4.3),

hence delivering the (incorrect) conclusion that the demonstrated actions are useful, even if

that utility is currently opaque to the audience. We are suggesting, in other words, that the

reason only humans reliably demonstrate overimitation is that only humans reliably inter-

pret ostensive behaviour in terms of optimal relevance (see also Morin, 2016, p.244-245).

We note, consistent with this interpretation, that overimitation emerges in development very

soon after the emergence of ostensive communication.

Finally, dogs are also an informative point of comparison. Dogs have been subject to

a long period of domestication in which humans are often prosocial towards them. In this

ecology it can be adaptive for dogs to simply presume that when humans attempt to gain

their attention, it is indeed worthwhile to actually pay attention. Correspondingly, dogs seem

to be sensitive to some of the most salient human ostensive behaviours (see e.g. Topál et al.,

2014; Wynne, 2016 for reviews). This sensitivity, moreover, emerges early in development

and is highly heritable (Bray et al., 2021). Presumably, dogs do not make the same interpre-

tative inferences as humans, but they do show how, in the right evolutionary ecologies, it can

be adaptive to spontaneously presume that others—in this case, human owners—are being

cooperative when they attempt to gain attention.

8. Diversity in human expression

In this section and the next, we suggest how the cognitive mechanisms described in previous

sections underpin many otherwise diverse means of human expression. We focus in particu-

lar on the examples of coordination smoothers (§8.1), teaching (§8.2), punishment (§8.3),

art (§8.4) and the emergence of symbols and grammar (§9). (This list of expressive domains

is of course not exhaustive.) In each case, we summarise how these different means of ex-

page 30 of 51
pression each employ the common, unified set of cognitive capacities described in previous

sections, but in different ways in each case. If we are right about this, then the evolutionary

emergence of specialised, cognitive means of manipulation (§3) and social vigilance (§4),

which together unleash expression (§5), is a root cause of many of the most distinctive as-

pects of human behaviour and societies.

We will highlight in particular the importance of graded aspects of human expres-

sion. In §3.3 we mentioned how there is a graded quality on the production side: different

means of expression can vary in the extent to which the actor makes her informative inten-

tion manifest. In §4.3 we mentioned how there is, in turn, variation on the audience side:

recognition of the actor’s informative intention can contribute to satisfying the informative

intention to different degrees in each case. In the subsections below, we describe how differ-

ent forms of human expression make use of these graded aspects in a range of different

ways; and we suggest that in general people aim to make their informative intentions mani-

fest just to the extent that the informative intention is likely to be satisfied, but no more so.

8.1. Coordination smoothers

To coordinate with one another in joint actions, such as dancing with a partner, maintaining

a tempo, moving large objects together, and many others, individuals must be informed

about each other’s behaviour and likely future behaviour, often on a moment-by-moment

basis (Sebanz et al., 2006). This can occur passively, but individuals also behave in ways that

actively facilitate the flow of information for joint action. For instance, two people may have

a goal to lift and move a large table. In lifting their end of the table, each person might move

in slightly exaggerated ways, in order to be predictable. Such behaviours are called ‘coordina-

tion smoothers’: they enable predictability for coordination (Vesper et al., 2010; 2017). Some

cases of coordination smoothing are clearly communicative, such as road signs and forms of

language use targeted at easing the flow of conversation (‘discourse markers’, ‘procedural

meaning’: see e.g. Blakemore, 2002; Gibbs & Bryant, 2008). However, cases in which in-

formative intentions are less overt have only recently been explicitly analysed in terms of

communication and expression (e.g. Pezzulo et al., 2019).

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 Consider two people, Jane and Paul, walking towards one another on a relatively nar-

row street. Jane makes a clear movement towards one side in order to make her action pre-

dictable to Paul. This informs Paul that Jane intends to proceed on the right, and it can do so

even if Paul does not recognise that Jane has this informative intention. Alternatively, Jane

might exaggerate her movement to one side. This makes her informative intention more

manifest, and Paul can hence infer that she has an informative intention that he believes she

will proceed on the right, and trust in it. This raises the question: when, why and to what ex-

tent do individuals make their informative intentions manifest? When should Jane not just

clearly move, but exaggerate her movement to one side? And to what extent? Corresponding-

ly, on the audience side: to what extent does recognition of the actor’s informative intention

contribute to successful coordination smoothing? These are all empirical questions whose

answers depend on how individuals take into account the constraints and the affordances of

the situation, and how they navigate graded dimensions of human expression.

Many results in the experimental study of joint action suggest that people indeed ex-

aggerate or otherwise adjust their actions to the extent that it is useful to do so for the pur-

poses of informing others; or, in other words, they make the least costly difference that is

large enough to make a difference (e.g. McEllin et al., 2018a,b; Schmitz et al., 2018; Curioni

et al., 2019). Observers, in turn, attribute to others commitments to behaving in the predict-

ed way just to the extent that those others are perceived as acting on a communicative inten-

tion (see e.g. Gibbs & Bryant, 2008; Bonalumi et al., 2020; 2021). In sum then, we are sug-

gesting that coordination smoothing is a form of human expression, in which people navi-

gate graded aspects of human expression in a competent way.

8.2. Teaching

Human teaching is richly diverse. Ethnographies of teaching reveal practices that span the

full range of human expression, ranging from tolerated observation, in which a skilled indi-

vidual just allows others to observe her in practice, to, at the other extreme, direct verbal

statements from the teacher, which the learner is expected to internalise (e.g. Lave &

Wenger, 1991; Marchand, 2010; Sugiyama, in press). Most actual instances of teaching lie

page 32 of 51
between these extremes and also include, for instance, repeated demonstration, perfor-

mance, exaggeration, role-play and countless other forms of expression (see e.g. Kline, 2015

for one recent review). We suggest that, as with coordination smoothers, the diversity of

teaching can be organised in terms of graded distinctions in human expression. Different

means of teaching vary in the extent to which the teacher makes her informative intention

manifest; and also the extent to which recognition of the teacher’s informative intention con-

tributes to successful interpretation, and hence learning.

Consider, for instance, a dance teacher (see e.g. Downey, 2008 for cognitively-in-

formed ethnography of dance teaching). At one extreme, she might simply repeat a dance

step multiple times, possibly from a range of different angles, allowing learners to observe,

without any further guidance about which aspects of the movement to attend to. Here the

teacher has an informative intention, and this intention is not hidden at all, but the teacher

does not make the intention manifest, and the learners employ means-end relations to learn.

Alternatively, at the other extreme, the teacher might openly exaggerate some of her move-

ments and hence highlight especially relevant aspects, which would otherwise remain unno-

ticed. By doing this the teacher makes her informative intention manifest. This triggers in

learners spontaneous presumptions of relevance (§4.3), which allows them to differentiate

exaggeration from the actual target behaviour, and hence identify what is especially relevant

about the teacher’s movements. In between the extremes are cases where the teacher makes

her informative intention somewhat manifest, such as when she slows her movement but in

a slight way only. In these ways and others, teaching is a phenomenon that, in its diversity,

spans the two graded dimensions of human expression we highlighted above.

When should different modes of teaching be employed? Sometimes it is suitable to

simply perform the target behaviour as usual and just allow observation, sometimes it is

suitable to make an informative intention somewhat manifest—and sometimes it is neces-

sary that the teacher’s informative intention be made wholly manifest. In particular, by trig-

gering learners’ presumption of relevance, teachers can teach things even though what

makes those things relevant is opaque to the learners (Gergely & Csibra, 2006). A real-world

example is teaching the counting routine to children, who learn by presuming that the rou-

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tine is relevant even though they do not yet understand its real utility. This mode of teaching

is arguably crucial for conceptual change (Heintz, 2011).

This perspective on teaching, as a subclass of human expressive behaviours, can in

turn help to explain why teaching is prevalent in human societies but relatively rare in other

species (see e.g. Hoppitt et al., 2008; Thornton & Raihani, 2008 for comparative perspec-

tives). Human teaching involves the dynamic use of unleashed expression. Analogous behav-

iours are observable in other species, but not with the same dynamic, open-ended and flexi-

ble range of behaviours that are afforded by truly unleashed expression, and readily exploit-

ed by human teachers and learners.

8.3. Punishment

Punishment is not always or intuitively thought of as a means of expression. Particularly

within cognitive science and cognate fields such as behavioural economics, punishment has

traditionally been modelled simply as retribution or incentive, such that it discourages or

deters specific behaviours. The actual delivery of punishment is then necessary only to main-

tain the integrity of the incentive structure. Yet the rewards and punishment that humans

tend to produce are actually inefficient for this goal. That is, they do not incentivise the target

behaviour well, contrary to the intuitive model (Ho et al., 2019; Cushman et al., in press).

The way people punish is, rather, best explained in terms of expression (ibid.). More pre-

cisely, we suggest that punishment is used to inform others that future exploitative beha-

viour will result in future costs (see also e.g. Sripada, 2005).

But why, then, is this expressive function not patently apparent? In other domains

(linguistic communication, teaching, art) expression and communication are utterly plain to

see. We suggest that punishment is usually most effective when its communicative aspects

are somewhat hidden, even though its informative aspects are present. In short, punishment

commonly involves hidden authorship (see §3.2).

The crucial point is that in ordinary social relations, punishment is credible only if

the incentives behind it are perceived to be stable; but in fact, in ordinary social relations the

incentives to inform are unstable. Specifically, they are dependent on the possibilities of fu-

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ture collaboration: if we are unlikely to interact in the future, I have no substantive incentive

to inform you that your behaviour was unacceptable. So if the real incentives that motivate

punishment were actually made manifest, they would be revealed as unstable and would

hence undermine the credibility of punishment as a means of informing. In this respect pun-

ishment is akin to generosity. We mentioned in §3.2 that there is a slight paradox to generos-

ity, in the sense that while it can be motivated by an intention to advertise oneself as proso-

cial, this intention should not itself be too manifest, lest the act of generosity be seen as in-

sincere. We are suggesting that a similar dynamic plays out on the punishment side: pun-

ishment does not credibly demonstrate a willingness to retaliate against anti-social beha-

viour if it is perceived as being motivated only by an intention to demonstrate a willingness

to retaliate against anti-social behaviour.

Institutional forms of punishment, in particular the legal punishments of nation

states, provide a revealing contrast. In jurisprudence the communicative aspects of legal

punishment have been long recognised: punishment is described as, for instance, “a conven-

tional device for the expression of attitudes of resentment and indignation, and of judgments

of disapproval and reprobation” (Feinburg, 1965, p.400; more recently see e.g. Primoratz,

1989; Hampton, 1992; Duff, 2001). The expressive dimension of punishment is straightfor-

wardly recognised in the legal domain because, we suggest, there is in this domain no real

doubt that the punisher (the nation state) has a long term, stable incentive to inform the au-

dience (citizens) about what is unacceptable. Nation states hence have no real need to hide

the informative dimensions of their punishments.

The expressive dimensions of punishment can be investigated in much more detail.

In particular, we know of no studies focused on whether, or to what extent, people interpret

or understand punishment as communicative; nor on the effectiveness of punishment as a

means of expression. Such studies would form an important bridge between cognitive prag-

matics and other fields, such as the newly emerging area of experimental jurisprudence (e.g.

Sommers, 2021). We also know of no existing research examining the expressive dimensions

of legal punishment from a historical perspective.

page 35 of 51
8.4. Art

Art is clearly expressive in some way, and audiences interpret artistic outputs in open-ended

ways. Modern audiences in particular are granted a great deal of autonomy in how they en-

gage with art, and are encouraged to develop and seek out their own, often highly personal

interpretations. We suggest that, like the other examples above, the open-endedness of artis-

tic expression derives from the natural character of human expression more broadly; and

hence that the interpretations that audiences derive from artwork, which are often highly

personal, are nevertheless often prompted by and derive from the same set of cognitive pro-

cesses that govern more ordinary forms of communication.

Crucially, what differentiates artistic expression from ordinary behaviour is not any

fundamental aesthetic quality as such, but rather the overt presentation of an object as an

aesthetic experience worthy of attention. This is the conclusion of many lines of research and

argument from at least four different fields: traditional art theory (e.g. Danto, 1964; Dickie,

1987), cognitive pragmatics (e.g. Pignocchi, 2019; McCallum et al., 2020), philosophy of

mind (e.g. Fodor, 2012), and social anthropology (e.g. Dissanayake, 1988; 2003). The overt

presentation of objects as worthy of attention is respectively labelled ‘Artworld’, ‘ostensive’,

‘Gricean’, or ‘making special’, in each of these four literatures. This consensus in turn sug-

gests that while proclivities towards aesthetic experience might be observable in other spe-

cies, the evolutionary emergence of cognitive capacities for unleashed expression, as a core

part of the human cognitive phenotype (§6), allows those proclivities to be expressed in

overtly intentional ways, thus differentiating art from other forms of aesthetic experience.

The emergence of art institutions can further reinforce these effects . Contemporary

art galleries in particular use white walls, open spaces and other features of curation to

present artworks as items to be considered and appreciated, hence triggering, we claim, the

audience presumptions of relevance that are an indelible part of ordinary communication.

These and other institutional effects, which can generate highly personal, even idiosyncratic

interpretations, are well established in art theory. What cognitive pragmatic perspectives

help to provide is description of how these effects exploit and otherwise build on the way au-

page 36 of 51
diences spontaneously interpret ostensive behaviour in more ordinary forms of social inter-

action (Pignocchi, 2019; McCallum et al., 2020).

8.5. Languages

Language use is quintessentially ostensive. Unlike some of the other means of expression

described above, language use involves making informative intentions wholly manifest. We

are arguing, in other words, that cognitive capacities for ostensive communication are

foundational to language use: there could not be any languages or linguistic communication

without the prior existence of cognitive capacities for ostensive communication (see also e.g.

Levinson, 2006; Tomasello, 2008; Scott-Phillips, 2015). As we put it in §1, natural languages

exploit unleashed expression: not the other way around.

Languages have, of course, their own particular features that collectively distinguish

them from other means of communication. In particular, linguistic communication makes

ubiquitous and multi-layered use of communicative conventions: phonemes, morphemes,

words, grammars and so on, which function to associate particular behaviours (speech, ges-

ture) with particular inferences that the communicator intends to trigger in the audience.

This ‘pragmatics-first’ perspective on the nature of languages aligns with usage- and con-

struction-based approaches to grammar, which emphasise how linguistic constructions are

used as a means to provide evidence of speaker meaning (e.g. Tomasello, 2003; Goldberg &

Suttle, 2010, 2003; Bybee & Beckner, 2010; Schmid, 2020; Hartmann & Player, 2021).

Where do communicative conventions come from? In recent decades the emergence

and stabilization of communicative conventions has been documented in several real world

case studies (e.g. Kegl et al., 1999; Meir et al., 2010; Brentari & Goldin-Meadow, 2017; inter

alia), and studied experimentally in the laboratory (e.g. Fay et al., 2013; Schouwstra & de

Swart, 2014; Granito et al., 2019; Motamedi et al., 2019; Raviv et al., 2019; inter alia). What

is commonly observed in these literatures is how behaviour that is sufficiently similar to pre-

viously successfully informative behaviour tends to be interpreted by audiences as a further

token of the same type as previously used, even after just one interaction; and also how fur-

page 37 of 51
ther repetition causes the focal behaviours to become increasingly conventional. How and

why does this happen?

In answering this question, what is not often recognised is that the very same cogni-

tive capacities that make ostensive communication possible in the first place, also play a piv-

otal role here. In particular, without audience presumptions of optimal relevance (§4.3), be-

haviour that resembles past communicative behaviour is mysterious and strange. Why re-

peat a past behaviour, and bring attention to it? Such behaviour is worth doing only if the

attention grabbing repetition of a past behaviour triggers in audiences the interpretation that

the behaviour is being used for the same or similar purposes as previously. So past events

have a role in communication not simply because of statistical learning of associations, but

because alluding to past events is typically the most efficient way to trigger, in the target au-

dience, interpretative inferences that are the same or similar to those triggered previously.

Furthermore, this allusion to past events will often mean that a given behaviour can

afford to be slightly less complex or less elaborate than the previous version, so long as the

allusion is still made. Communicative success becomes increasingly governed by the simple

fact that echoing how behaviours have been successfully used in the past is the most efficient

means of prompting the intended inferences in the audience. Repeated many times over, this

leads to gradual simplification of the stimuli; and in time shapes many of the features that

are characteristic of natural languages, such as displaced reference, compositionality, low

levels of resemblance between form and use (‘symbolism’), statistical relationships between

word length and frequency of use (e.g. ‘Zipf’s law’), and so on. These processes can, we be-

lieve, be fruitfully analysed within an epidemiological framework (e.g. Enfield, 2003; 2014;

Scott-Phillips et al., 2018).

Finally, we note that linguistic stimuli are processed by some dedicated cognitive ca-

pacities (see e.g. Hagoort, 2017 for one recent summary). Crucially, these capacities appear

to work in parallel with cognitive processes of ostensive communication more broadly. On

the comprehension side in particular, inference of what is said and inference of what is

meant are not serial, with one following the other, but instead seem to impact on each other

in a dynamic process of parallel ‘mutual adjustment’ (see e.g. Carston, 2002; Wilson, 2004;

page 38 of 51
Sperber & Wilson, 2005 for post-Gricean description of this process; and e.g. Spotorno et al.,

2012; Vanlangendonck et al., 2018; Paunov et al., 2019 for neuroscientific evidence). Further

and deeper integration of findings in psycholinguistics and cognitive pragmatics are import-

ant future research goals (Noveck, 2018); but in any case, the evolutionary emergence of

these dedicated capacities must have followed, rather than preceded, the evolutionary emer-

gence of ostensive communication described in §6.

9. Conclusion: Rethinking pragmatics

Historically speaking, pragmatics has been situated on the periphery of the language sci-

ences, appearing in textbooks usually only as a fringe topic. We have argued, in contrast, that

cognitive capacities for ostensive communication are foundational, because they unleash ex-

pression on a grand scale. This in turn redefines the domain of pragmatics itself. Rather than

being narrowly conceived as the study of how context influences the interpretation of utter-

ances—which is how it is conceived in many approaches—pragmatics should be charac-

terised as the study of how, and the many means by which, informative intentions are sat-

isfied. The core questions for pragmatics are how informative intentions are made manifest,

and to what effect. Language use and other conventionalised means of expression are the

most salient specific instances, and are clearly central to human sociality, but there are many

others too.

Here we have grounded this rethinking of pragmatics in a broad evolutionary and

cognitive context. Human communication has long posed a challenge to evolutionary biology

and signalling theory, due to its versatility, diversity, and clear proneness to deception (§2).

We have described how this problem is resolved in humans, allowing truly unleashed expres-

sion (§5); described key graded differences between the specialised capacity of mind that

drives human communication and other means of social cognition (§3 & §4), some of which

are shared with other great apes (§7); and provided a crucial proof of evolvability, by relating

these graded differences to specific and distinctive aspects of the human social ecology (§6).

We also described how several different means of human expression each employ the relev-

ant cognitive capacities in interestingly different ways (§8). If we are right about this, then

page 39 of 51
the evolutionary emergence of specialised, cognitive means of manipulation and social vigil-

ance is a root cause of many of the most distinctive aspects of human behaviour and societ-

ies.

An essential future goal is to establish a formal foundation for this reconceived prag-

matics. An important first step in the development of scientific understanding is informed

description of the phenomenon of interest, recognising its full complexity while simultan-

eously providing some provisional order and generality. We hope to have contributed to that

goal here, which is too often neglected in contemporary cognitive psychology and cognate

disciplines (Rozin, 2001; Rai & Fiske, 2010; Doliński, 2018; inter alia). Formal models build

on these descriptive foundations. In this respect Bayesian models of interaction, in which

production and comprehension are modelled as inter-connected planning problems over

others’ mental states, are a particularly promising direction (e.g. Shafto et al., 2014; Good-

man & Frank, 2016; Ho et al., 2019). Different specific models differ in the detail and we do

not subscribe to all of the assumptions made. 11 Moreover, we do not believe that any existing

model is sufficiently general to cover the full range of prototypical cases of communication

(pointing, gesture, inter alia), let alone the broader diversity we highlighted in §8. Nonethe-

less, we do believe that modelling human interaction in these terms is an important and

promising research direction. We hope that our description of the evolutionary and cognitive

foundations of human expression, in all its diversity, will aid this agenda.

More broadly, we agree with recent arguments that the human evolutionary sciences

are presently too ‘cognition blind’ (e.g. Morin, 2016; Heyes, 2019), and that greater theoret-

ical unity will be achieved by deeper consideration of the cognitive processes that underpin

otherwise diverse human behaviours. The case of communication and expression is, we be-

lieve, a clear case in point. Quite commonly, particular types of expressive behaviour (lan-

guage use, teaching, punishment, art, inter alia) are considered each in isolation, without

11Consider, for instance, the Rational Speech Act framework (Goodman & Frank, 2016). Here, com-
municative behaviours are assumed to have ‘literal’ meanings, independent of use, from which speak-
ers might deviate. We do not share this assumption; that is, we do not believe that there is any such
thing as ‘literal meaning’ so conceived. Indeed, this is one of the key points of difference between neo-
Gricean approaches and the post-Gricean approach we have adopted in this article (Sperber & Wilson,
2005; see also citations in §8.5, on the process of parallel mutual adjustment, which undermines the
notion of literal meaning).

page 40 of 51
much consideration of the possibility that they might in fact derive from the same underlying

biological trait: as if the evolution of running was considered in isolation from the evolution

of walking, when in fact both are derived sub-functions of the unified capacity for bipedal

locomotion. We have argued that the same applies here. Different types of human ‘expressive

act’ are each derived sub-functions of an underlying unity.

Acknowledgements

For comments on previous drafts, we thank Josep Call, György Gergely, Günther Knoblich,

Dan Sperber, Cordula Vesper, and members of the Smash, Oscar and Aces research groups

at Central European University. For extensive discussions on the cognitive and evolutionary

basis of human expression, we thank the ‘Relevance Researchers Network’, and all members

of the Somics project, ‘Constructing social minds: Coordination, communication, & cultural

transmission’.

Conflicts of interest

None

page 41 of 51
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