New thinking, innateness and inherited representation.

2012·0.21 MB·English

by  Shea, Nicholas

Phil. Trans.R.Soc. B(2012) 367, 2234–2244
doi:10.1098/rstb.2012.0125
Review
New thinking, innateness and inherited
representation
Nicholas Shea*
Faculty of Philosophy, University of Oxford, 10 Merton Street, Oxford OX1 4JJ, UK
The NewThinking contained in this volume rejects anEvolutionaryPsychology thatiscommitted
to innate domain-specific psychological mechanisms: gene-based adaptations that are unlearnt,
developmentally fixed and culturally universal. But the New Thinking does not simply deny the
importance of innate psychological traits. The problem runs deeper: the concept of innateness is
not suited to distinguishing between the New Thinking and Evolutionary Psychology. That
points to a more serious problem with the concept of innateness as it is applied to human psycho-
logical phenotypes. This paper argues that the features of recent human evolution highlighted by
the New Thinking imply that the concept of inherited representation, set out here, is a better tool
for theorizing about human cognitive evolution.
Keywords: evolutionary psychology; innateness; inherited representation; genetic information;
cognitive evolution; cultural inheritance
1 INTRODUCTION of interdependent adaptive changes, but it is not
(a)NewThinkingversusEvolutionaryPsychology adapted to any one particular task or outcome. It
Itisstandardlyassumedthatinnatepsychologicalmech- appears to have been selected instead for its facility
anisms should be the central focus of an evolutionary as a generalist: to perform an open-ended range of
account of human cognition. One version of this tasks with great skill, where the concrete outcomes
approach isthe ‘Evolutionary Psychology’ (capitalized) that contribute to fitness vary widely.
ofToobyandCosmideswithitsseriesofspecial-purpose Two domain-general changes in hominin evolution
modules, each evolved to solve a particular problem in arecentralhere.Thefirstisamarkedincreaseinplas-
thePleistocene[1].EvenoutsidehighchurchEvolution- ticity: in how quickly human individuals and social
ary Psychology, forexampleamong thosewho point to groups adapt to different ways of life in different
more general-purpose evolved human faculties like the ecological niches [6]. These developmentally plastic
capacity for fine-grained visuomotor control, an evol- innovations depend for their development on rich
utionary approach to human cognition is usually allied support from the environment: information about
withacommitmenttoinnatecapacities. particular variable features of the developmental
A strong theme emerging from the papers in this environment, including resources and information
volume puts pressure on the idea that there are derived from the ecological niche, from parents and
innate psychological capacities at all. These are not from the wider culture [7].
blank slate empiricists or cultural determinists who The second change is a much greater reliance on
deny the importance of evolution for explaining why culturally transmitted information. Among the
we are as we are. If the New Thinking is on the right adaptively-significant information on which develop-
track,however,theninnatenessisnotausefulconcept ment depends, some concerns facts that individuals
for theorizing about recent human cognition. could not learn for themselves in their own lifetime.
Many papers in the volume emphasize domain- Inthesecircumstancesindividualsdonottestandcon-
general adaptations: improved physical/causal under- firm for themselves that the developmental path they
standing, increased visuomotor skill, and an extended are adopting is adaptive; they rely on the information
childhood coupled with increased parental investment in their culture being a reasonably reliable guide to
[2,3].Atarecentconferenceonthenewthinking,Eva adaptive outcomes. So human behavioural develop-
Jablonka argued that the hand is a much better meta- ment is extremely plastic, and some of that plasticity
phorthantheSwiss-armyknifefor thespecialfeatures involves sensitivity to culturally transmitted infor-
of recent human cognition [4,5]. The hand is exqui- mation [7,8]. More strongly still, it is plausible that
sitely complex and highly adapted, involving a suite humanshaveadaptations for theculturaltransmission
of information, including through teaching and learn-
ing[9].So,ifcognitionhasevolvedtobelikethehand,
*[email protected] itisahandthatwaspartlyshapedbyacultureofother
hands, and which in turn playsa role in shaping other
One contribution of 15 to a Theme Issue ‘New thinking: the
evolutionofhumancognition’. hands, being itself partly adapted to passing on the
2234 Thisjournalisq2012TheRoyalSociety
Review. New thinking and innateness N. Shea 2235
kindofculturalresourcesonwhichitdependedonfor (b) Deeper troubles with innateness
its own development. Inutilityisbadenough,butthetroubleswiththeinnate-
An evolutionaryapproachis often coupled with the ness concept run deeper. As philosophers, we want to
thoughtthatadaptivetraitsareinnate.Aninnatetraitis know what property the concept of innateness refers
prototypically1anadaptationthathasevolvedbygene- to.Thatis,wewantatheoreticalreconstructionofthe
based natural selection, and is thereby coded in the claim that a trait X is innate—we want to know what
genes. Its development does not depend on learning, itisaboutatraitthatdetermineswhethertheinnateness
anditwoulddevelopevenifexperiencewereimpover- claimistrueorfalse.Thisexerciseisnotmerelyofphi-
ished in certain ways. Correlatively, it is relatively losophical interest. It is continuous with the scientific
developmentally fixed in the face of environmental project of trying better to understand the properties
variations, and perhaps canalized against such vari- that are appealed to in a scientific theory. To do so,
ations. Indeed, on some views an innate trait is we can observe the pattern of inferences in which a
genetically determined from birth. And it is universal theoretical concept such as innateness is deployed, and
in the species, either in the sense that all members seektounderstandwhatthepropertyisthatunderpins
have it apart from non-typical cases in which there is those inferences: that makes them defeasible but
an explanation of their exceptional status or, if there reasonably reliable ways of reaching true conclusions
is typical variation in how the trait is manifested, then fromtruepremises.Thedeeperproblemwiththecon-
some shared basis for thatvariationis universal. cept of innateness is that there is no good account of
Evolutionary Psychology aims to account for the what the property of being innate could be, so as to
distinctive features of human life by appealing to underpin the ways that the concept is used in relation
special-purpose psychological capacities that have tohumanpsychologicalcapacities.
exactly those features: they are prototypically the Theconceptofinnatenessisusedtomakeinferences
result of gene-based natural selection, do not depend between various properties like those mentioned ear-
upon learning for their acquisition (and so admit of a lier: developmental fixity, lack of learning and genetic
poverty of the stimulus argument), are relatively coding—‘i-properties’ (adapting the terminology of
developmentally fixed and hence culturally universal. Mameli & Bateson [17]). For instance: X is an adap-
This is in sharp contrast to the central features of the tation!X is innate!X’s development does not
NewThinkingidentifiedearlier.Soitmightbethought dependuponlearning.Thatinferenceishighlyunreli-
that Evolutionary Psychology believes in, whereas the able because, as the New Thinking points out, many
New Thinking denies, the importance of innate recent human adaptations depend crucially on learn-
psychologicalcapacities;andthusthattheNewThink- ing. The problem is not just that the concept is
ing makes less appeal to evolutionary considerations sometimes used to make bad inferences. Rather, the
than high churchEvolutionaryPsychology. problem is that there is no good proposal about how
That is a misdiagnosis: the New Thinking is just theconceptcouldbeused tomakereliableinferences.
as motivated by evolutionary considerations. It relies What could the property of being innate amount to,
heavily on selectionist explanation, phylogenetic such that the conclusions about human psychological
methods and cross-species comparisons. It sees many capacitiesdrawn using the concept should be reliable?
domain-specific adaptations in human cognitive and If there were occasional exceptions, the concept
social life. Some arise from gene-based selection, would still be useful. If there were systematic excep-
others from culturally based inheritance processes. tions, they could be guarded against in deploying the
Some are universal and others vary in different eco- concept. Buttheconceptismuch moreproblematicif
logical and cultural contexts. Learning is involved in there is no property at all that would make inferences
the acquisition of many of these traits, even when among the i-properties generally reliable in relation to
individuals are not able to learn about the adaptive humanpsychologicaltraits.
significance of the trait for themselves. The New The New Thinking puts pressure on the possibility
Thinking does not sit well either with accepting or ofasatisfyingtheoreticalreconstructionoftheconcept
denying that the relevant capacities are innate. That of innateness in two ways, one familiar, and the other
is because the concept of innateness is fundamentally less so. Its richly interactive picture of development
unsuited to elucidating the contrast between the New undermines the tie, mediated by the innateness con-
Thinking and Evolutionary Psychology. cept, between adaptation and developing without
This paper will argue that the concept of inherited reliance on experience. There has been selection for
representationdoesamuchbetterjobofdrawingthedis- a variety of clever behavioural phenotypes, such as
tinction. Evolutionary Psychology is committed to the ability to process foodstuffs, or to make tools, or
gene-based selection as the main source of the adap- to cooperate with others in a hunt. Extended child-
tively relevant information that is responsible for hood and increased parental investment allow the
the adaptiveness of human psychological capacities; development of many human phenotypes to be more
hencesuchcapacitiesareunlearnt.TheNewThinking richly interactive with the environment for a longer
emphasizes the complementary roles of gene-based period.Thesechangesinlifewaysarethemselvesplau-
and of culturally-based selection processes of various sibly adaptations [19]. Evolution does not care about
kinds for generating the adaptively relevant infor- the devious and richly interactive route by which
mation on which human psychological capacities are some clever behavioural phenotype has developed. It
based; hence the importance of interactions with the sees the phenotype that is the end point of develop-
physical and social environment, and of learning, in ment, and selection acts on that. Plausibly, many are
developing those capacities. adaptations. That developmental process happens to
Phil.Trans.R.Soc.B(2012)
2236 N. Shea Review. New thinking and innateness
depend very heavily on learning and culture. When The other leading proposal for a theoretical recon-
development is so richly interactive, depending upon struction of the concept of innateness is that it just
all kinds of material and informational support from picks out the property of being unlearnt [14]. That
the structured ecological environment, and from would make sense of the way the innateness concept
other people,itisjustamistaketolinkbeinganadap- is used in relation to human psychological capacities
tation with developing relatively independently of if traits that depend on learning tend not to have the
experience. i-properties, i.e. tend not to be species typical, nor
Furthermore, the emphasis on plasticity severs any developmentally canalized, noradaptations. The diffi-
potential link between adaptation and developmental culty for that construal is that no way of making the
fixity. Typical human innovations develop in a way concept of learning more precise does the job, as will
that is open to environmental information, so as to be argued later. To preview: if learning is understood
fit in with variations in the environment. Even if narrowly then many species-variable, developmentally
such outcomes are robust against some forms of plastic traits will count as unlearnt, hence innate. On
environmental variation, it is misleading to think the other hand, if learning is understood widely to
ofphenotypeswhosedevelopmentdependsuponvari- include all the cases where an individual extracts
able features of the environment as being relatively information from the environment during develop-
fixed or canalized against environmental variation. In ment of a psychological trait, then many universal
short, in recent human evolution at least, there is no developmentally canalized traits, both adaptations
tie between being a phenotypic adaptation and devel- and non-adaptations, will count as learnt, hence not
opment’s unfolding in a certain way (i.e. being fixed innate. Neither way of delimiting what is to count as
and/or independent of experience). a learning process will make inferences among the
The interactive nature of development has long i-properties generally reliable in respect of human
been the basis of critiques of the innateness concept psychologicalcapacities.
[20].Standardcounterexamplessuggestthatevenuni- Inshort,eveniftheconceptofinnatenessworkswell
versal adaptations need not be particularly well- enough in other biological domains, it is problematic
insulatedagainstenvironmentalvariation.Forinstance with respect to the phenotypic traits that have arisen
normal skin structure in humans depends on dietary in recent hominin evolution. Nevertheless, the New
vitamin C. Nevertheless, it was still argued that, for Thinking is a thoroughly evolutionary approach.
gene-based adaptations, development is so organized What replaces claims about innateness? This paper
that genetic information plays a special role. Granted, argues that the innateness concept should be replaced
environmental factors give essential causal support to bytheconceptofgeneticrepresentation,anditsgener-
the development of an adaptive trait, just as they did alizationtoepigeneticandculturalinheritancesystems:
when it was selected, but the information that allows inherited representation. Inherited representation is
it to be adaptive derives from the genes, not from not a direct substitute for innateness, but allows us to
the environment. That is the proposal offered by articulate the way in which selection and adaptation
KonradLorenzinresponse tocritiquesoftheconcept arecentraltotheNewThinking.Itissuitedtoexplain-
of innateness [21]. His proposed theoretical recon- ing aspects of phenotypic development that the
struction of the innateness concept is that innate abandonment of the concept of innateness would
traitsarethosewhosedevelopmentdependsongenetic leave unexplained. Section 3 makes that case, and §2
information. That line is challenged by the second lays the foundations by setting out an account of the
feature of the New Thinking identified earlier: the semantic information carried by inheritance systems
fact that much adaptively-significant information is asit applies in other biological domains.
culturally transmitted.
Although some forms of cultural evolution remain
controversial, there are a variety of ways that broadly 2 GENETIC REPRESENTATION
cultural processes accumulate adaptively-relevant (a) Information generated by natural selection
information that is relied on for the development of on genes
adaptive phenotypes [22]. It has been argued that in The first step is to see how genetic representation
humans gene–culture coevolution accelerated this explains what is going on with innateness claims in
processthrough positive feedback [6], both increasing other cases, outside human cognition. Then §3
the number of phenotypes that depend in part on argues that the concept goes particularly badly awry
gene-based adaptation, and magnifying the extent to whenappliedtotheresultsofrecenthumanevolution.
which human cognitive development relies on cultu- The concept of innateness is applied on the basis
rally transmitted information. That undermines the of, and used to make inferences to, the various
possibilityofevenaroughdichotomybetweenindivid- i-properties mentioned earlier. We have seen that the
uals learning for themselves and relying on genetic robustnessoftheconnectionbetweenthese properties
information. Reliance on culturally transmitted and being a genetically-based adaptation has been
adaptively-significant information forms an important seriouslyquestioned.Nevertheless,evolutionproduces
thirdclass:neithergeneticallytransmittedandpresent exquisitely adaptive phenotypes, and one feature of
at birth, nor just a matter of individuals learning how they develop would be entirely mysterious but
for themselves about their environment—since cul- for the theory of natural selection.
tural processes allow development to be sensitive to Consider the close match between the bills of the
information beyond that available in the lifetime of purple-throated carib hummingbird Eulampis jugularis
an individual. andtheshapeoftheflowersonwhichtheyfeed.Males
Phil.Trans.R.Soc.B(2012)
Review. New thinking and innateness N. Shea 2237
develop bills that match the short straight flowers of verymanyfactorsthatmakeacausalimpactondevel-
Heliconia caribeae, and the smaller females develop opment, including the genome, carry correlational
longer curved bills that fit perfectly into the long information about phenotypic outcomes—had the
curved flowers of Heliconia bihai [23]. The humming- factor been different, the outcome would have been
bird somehow manages to develop a bill that will different.Mostofthesefactorsjustcarrycorrelational,
perfectly match a particular species of flower, but not notsemanticinformation.Theclaimthatthereisgen-
as a result of causally interacting with the flower etic representation is the claim that genetic material
during development. The shape of the bill carries also carries information of the stronger, semantic var-
information about the shape of the flowers on which iety.Roughly theidea isthatgenes donotjusthappen
they feed. That information does not derive from to carry correlational information, but they are sup-
interacting with the flower shape during development. posed to—in the unobjectionable naturalistic sense of
So where did it come from? How does the developing ‘supposed to’ underpinned by evolutionary functions
hummingbird‘know’whatshapeofbilltoproduce,to and natural selection.2
match available nectar sources in its local ecology? The case for genetic representation is founded on
There is evidence that the bills and flowers have coe- the fact that DNA does not just happen to be a
volved, so the same question can be asked about the means of transmitting phenotypes down the gener-
development of Heliconia flowers [24]. ations. There is good evidence that it has undergone
Thesamequestionisjustaspressingforbehavioural selection for the way it performs that function
traitssuchasthoseweareconsideringhere.Howdoes [26,27]. Thatisto say, in addition to the evolutionary
theanimalknowhowtobehavewithoutlearning[25]? functions acquired by individual genes, the whole
More carefully, how does the developing individual apparatus of DNA coding, translation, transcription
acquireadispositionforspecies-typicaladaptivebehav- andreplicationhasthemeta-levelfunctionoftransmit-
iourwhentheinformationthatenablesthebehaviourto ting phenotypes down the generations. When a new
match the environment has not been derived fromthe genetic variantarisesthatcorrelateswith aphenotypic
environmentduring development? Ofcourseweknow difference,naturalselectioncanact.Thosecorrelations
that natural selection explains how adaptations arise, can be very long range, with the phenotypic variants
but it is less remarked that it also provides an answer being the result of a long and interactive process of
to this informational question about development. development.Still,theresultisthattheselectedgenetic
The developing individual does not need informa- variantanditsassociatedphenotypicvariantincreasein
tion because there has been natural selection for a frequency,hencecarrycorrelationalinformationabout
developmental process that arrives at the appropriate the circumstances of selection. The DNA system is
phenotypereasonablyreliably.Atsomepointagenetic designed to transmit that phenotype to future gener-
variantorvariantsarosethatbiaseddevelopmentinthe ations (by transmitting the genetic variant with which
direction of a particular phenotype, which affected it correlates). Now consider the zygotic DNA in an
fitness and was selected. Neither at the stage of selec- individual in a subsequent generation. For the system
tion, nor subsequently, did development need to pick of DNA expression to perform its meta-function it
uponinformationabouttheenvironment.How,then, should react to the selected gene by producing, by
did the informational connection between adaptive the same long interactive process of development, the
phenotype and environmental feature arise? By virtue phenotype for which it was selected.
of selection—increasing the frequency of the variant Thosefactsmeanthatthegeneticsystemfallswithin
whichadaptivelymatchedtheenvironment.Thedevel- a simple model for characterizing the contents carried
opmental outcome of hummingbird bill development by low-level representing systems [28–30]. The
now carries information about flower shape by virtue elementsarea‘consumer’systemwiththeevolutionary
ofahistoryofselectioninapopulation,increasingthe function of responding to a variety of different rep-
frequencyofmatchingvariants. resentations by producing a variety of different
outcomes,andsomemethodbywhichrepresentations
areproducedbywhichtheycorrelatewithexternalfac-
(b) Genes carry semantic information tors in such a way that the outputs of the consumer
This account of how genetic information plays a systemareabletoperformtheirevolutionaryfunctions.
role in individual development can be made more Asimpleexampleisthehoneybeenectardance.‘Con-
preciseusingthephilosophicalconceptofgeneticrepre- sumer’ bees respond to the dance of an incoming
sentation. We can distinguish between the kind of worker by flying off a certain distance and direction
information studied by information theory, based on inrelationtothesun.Thatbehaviourperformsitsevol-
correlations between one thing and another, and utionary function on condition that there is nectar at
semantic information, in which a sign or representation that distance and in that direction. The behaviour of
supports a distinction between correctness and incor- the consumer bees manages to fulfil that evolutionary
rectness, or being satisfied versus not. Correlational function because incoming worker bees produce
information is absolutely ubiquitous. Semantic infor- dances that correlate in a corresponding way with the
mation is much rarer in nature, but very familiar in location of nectar. The correlation produced by the
ordinary life. It is the kind of content carried by incoming workers is an ‘exploitable relation’ [30],
sentences and thoughts. which outgoing consumer bees make use of, in the
Heritability at the time of selection means that at sense that the correlation is a reasonably reliable indi-
the time of selection genetic variants carry correla- cator that a condition on the behaviour they will
tional information about phenotypic variants. And perform being useful isin place.
Phil.Trans.R.Soc.B(2012)
2238 N. Shea Review. New thinking and innateness
In the genetic case the genes in the zygote are the parentaleffectsmediatedbyothermeans.Someepige-
representations and the whole long extended interac- netic effects may be transmitted with high fidelity
tive processes of development are the consumer, down many generations, in which case they can
eventuating in the phenotypic features exhibited by accumulateinformationbyselectiononepigeneticvar-
the organism. (DNA is also active within individual iants,aswesawwithgenes,asmentioned earlier[38].
cells during development, but for present purposes If some epigenetic system turns out to have been
that is a distraction, because it is not part of the selected in part because of its ability to transmit phe-
reason why zygotic DNA has the representational notypes down many generations, then it too will have
content it does.) Because DNA has the exotic meta- the earlier-mentioned meta-function. I call any mech-
function mentioned earlier, its job is to react to indi- anism with that meta-function an inheritance system.
vidual genes by producing, possibly at the end of a The selected phenotypes that are so-transmitted are
long and complex developmental process, the pheno- inherited representations (genetic representations are one
types for which they were selected. Development kind). Most importantly for our purposes, human
makes use of the correlation between genes and the evolution seems to have depended in part on one or
environments in which they were selected as being a more cultural inheritance systems. In that case, some
reasonably reliable indicator that the current environ- of the information on which human phenotypic adap-
ment is conducive to the phenotypes that result. tations depend for their development derives neither
If the environment has changed in relevant ways from the genome, nor from the individual detecting
since the time of selection, that condition will no correlationalinformationavailablewithinitsownindi-
longer be in place, so the content carried by the gen- vidual experience, but from the developing individual
etic representation will be false. For example, if the reacting to information in its culture. Cultural pro-
selected phenotype depended upon the environment cesses of selection and transmission account for the
beingwarm,andithassincebecomecold,thenthecon- way such cultural variants carry correlational infor-
dition that the selected gene is representing—a warm mation about adaptively-significant facts.
environment—is absent. The genetic material then
continues to represent (falsely) that the environment
is warm and DNA expression and development, in (c) Genetic representation underpins some
accordancewith its meta-function, faithfully produces connections between i-properties
a warm-appropriate phenotype as result. That pheno- Genetic representation explains why the properties
type is no longer good for the organism. This failure associated with the concept of innateness cluster
is explained by the fact that the content represented together in some cases. In organisms with little or no
bythezygoticDNAisnowfalse. culture there are two main sources of adaptively-
This account of genetic representation has been relevant information. First, there is information that
developed in detail elsewhere [31–34]. It is not individuals can detect in their developmental environ-
simply a philosophers’ gloss on well-known facts mentintheirlifetime.Second,thereistheinformation
about natural selection. It has various explanatory conveyed by genetic representations (and, if they turn
benefits, for example when natural selection makes out to be empirically significant, by epigenetic rep-
choices between relying on genetic or environmental resentations that are carried along with the genome).
information about some fitness-relevant factor Aphenotypethatdevelopsinrelianceonageneticrep-
[35,36]. For our purposes the key payoff is that it resentation may adaptively match its environment in
offers an explanation of the question we started with: respects that the individual has not detected in its
how the developmental process in an individual man- own lifetime. So there is a connection with poverty
ages to achieve an adaptive match to its environment of the stimulus arguments. The trait can develop in
withoutderivingfromthedevelopmentalenvironment the absence of certain kinds of information about the
all the adaptively-relevant information that is actually environmental feature it comes adaptively to match.
carried by the adaptive phenotype. For example, an For example, the female hummingbird E. jugularis
individual E. jugularis female ends up with a bill that develops its long curved bill even if deprived during
so carefully matches the shape of the H. bihai flowers its development of anything that provides information
that it will feed from because the zygote contains gen- about the shape of H. bihai flowers.
etic material that instructs the development of a long Thatexplains alink between genetic representation
curved bill, and indicates that the environment is andone ofthei-properties.Thereisalsolikelytobea
likely to contain flowers conducive to that shaped certain amount of developmental fixity in a trait that
bill. That is why the individual does not need to develops in reliance on a genetic representation.
interact with the shape of the flowers, or derive infor- To have been selected, the trait must have developed
mation about their shape in any other way from its reasonably robustly across the kinds of environmental
own experience. variations encountered when it was selected. To the
If DNA were the only inheritance system, then a extent that current environmental variations fall
developing individual would have access to only two within that range, it is likely that the trait will arise in
sources of adaptively-significant information: genetic developmentdespitesuchvariations.Ifundercontinu-
material, and the correlations that it could detect in ingselection pressure,thetraitmayhaveevenbecome
the course of its own development. In fact the sources developmentally canalized against such variations.
are much wider. Transgenerational epigenetic effects Adaptive polymorphisms have sometimes been
allow an individual to make use of information thought to be an exception to this principle. If we
detected by its parents in their lifetime [37], as do adopt the right grain of analysis, it becomes clear
Phil.Trans.R.Soc.B(2012)
Review. New thinking and innateness N. Shea 2239
thattheyarenot.Evolutionhasselectedforadevelop- human evolution that fatally undermines any utility
mental program that produces different outcomes of the innateness concept.
in different environments, with some cue detected in Sometimes when individuals get information from
developmentresponsibleforthefactthatanindividual social sources they still learn for themselves. For
ends up with the variant that adaptively matches its example, with stimulus enhancement the individual
particular environment. That developmental disposi- learnsforitself,throughtrialanderror,howtoperform
tion has a causal basis, which must itself develop. So some useful novel action on or with the stimulus.
it is that causal basis, rather than the various poly- Nevertheless, with many of these social sources of
morphic outcomes of development, that is likely to information, a component of the adaptively-relevant
be developmentally fixed, at least across the range of informationisbeingcontributedbyaculturalprocess.
environments in which it evolved. The adaptive Itconsistsofinformationtheindividualdidnotdetect
matchoftheeventualmaturephenotypetoitsenviron- for itself. In some cases, it gives the individual access
mentispartlyduetogeneticrepresentationandpartly to a source of information that they could not detect
due to adaptive reliance on environmental infor- for themselves.
mation. We will see later that such mixed models Learning by ‘overimitation’ offers a clean example
have particular importance in the human case. of this point. Both humans [39] and other primates
Althoughatraitthatdevelopsinrelianceonagenetic [40] are capable of copying the means by which a
representation is likely to be fixed or canalized against demonstrated action is performed. Both can appreci-
the kinds of environmental variations present when it ate the difference between means to an intended goal
was selected, there is no reason to expect its develop- andirrelevantactions[41–44](cf.[45]).Thereisevi-
ment to be causally independent of the environment. dence that human children, unlike other apes, have a
Gene-basednaturalselectionworksonthebasisofcor- tendency to copy all the actions demonstrated in
relations between some genetic variant and some obtaining some desirable outcome, not just those
phenotypic outcome, correlations that are reasonably that are causally necessary to obtaining the outcome.
reliable at the time of selection. Those correlations Chimpanzees, in contrast, cut out action steps that
can be very long range indeed. A genetic variant can seem to them causally unnecessary [41,46].
bias the production of one phenotype over another by This contrast is apparent in experiments using two
all sorts of devious means, and the pathway leading versions of a mechanical box. In the transparent ver-
fromzygoticDNAtophenotypicoutcomecanbeintri- sion the mechanism is visible so it is apparent when
cate and interactive without limit. The informational some actions performed by the demonstrator are cau-
contribution of a genetic representation is compatible sallyirrelevanttoachievingthegoalofgettingareward
withextremelyrichcausalcontributionsoftheenviron- out of the box. When the action sequence is demon-
menttothedevelopmentofthetrait. strated on an opaque box, it is not apparent that
Taking stock: recognizing the role of genetic rep- some of the action steps are unnecessary. Both chim-
resentations in development explains why, in the panzees and human children copy all the action steps
absence of cultural inheritance systems, the following performed on the opaque box in order to obtain the
properties should co-occur better than chance: being reward. With the transparent box, chimpanzees cut
a universal or species-typical adaptation, developing out the unnecessary action steps and go straight for
inawaythatisinvariantacrosssomerangeofenviron- the reward. Human children, in contrast, have a ten-
mental variations, and there being an aspect of the dency still to copy all the demonstrated actions, even
adaptive information carried by the trait about its those that, by their own lights, are unnecessary to
environment that is not the result of learning and obtain the outcome. (That the children understand
about which a poverty of the stimulus argument that some steps are unnecessary can be demonstrated
could be made. These connections admit of excep- in other experiments.)
tions, for a variety of reasons, but they co-occur There also seems to be a developmental trajectory.
often enough to make inferences between these prop- At the age of 12 and 18 months infants are focused
erties go right better than by chance. So when a on outcomes and tend not to copy specific demon-
non-human trait is labelled ‘innate’ and taken to strated actions [47]. Three-year-olds are capable of
have these properties, the inferences it underpins will copying specific actions [48] but less inclined to imi-
be reasonably reliable. tate unnecessary steps in the sequence. By the age of
fiveyears, childrenimitate obviously irrelevantactions
in a wide range of conditions [49–52] (but cf. [53]).
3 INHERITED REPRESENTATION Overimitation appears to be even more pronounced
(a) Adaptation through cultural transmission in adults [54].
We cannot be so sanguine about innateness claims in Thehumantendencytooverimitatecanseempara-
thedomain ofhuman cognition. Therichinteractivity doxical—would it not lead to the spread of useless or
of human behavioural development firmly severs any even detrimental behavioural variants?—until we rea-
connection of gene-based adaptations with genetic lize that making individuals’ tendency to imitate less
determinism, or with their development being less sensitive to individual learning can increase its fidelity
causally dependent on the environment. That is the as an information transmission system [55]. The fide-
morefamiliarpointaboutinnatenessthathumanevol- lityofculturaltransmissioncanhavealargeimpacton
ution raises in a particularly pressing way. It is the the power of cultural inheritance mechanisms [56].
NewThinking’sconclusionthatculturallytransmitted If humans have a tendency blindly to imitate the
information has been particularly important in recent demonstrated intentional actions even when, by their
Phil.Trans.R.Soc.B(2012)
2240 N. Shea Review. New thinking and innateness
own lights, various of the actions are causally otiose, information onwhichtheyare based wasgeneratedby
then this would allow the transmission of behaviours selection,theirdevelopmentwillbesusceptibletoapov-
the utility of which is not apparent to any individual. ertyofthestimulusargument:therelevantinformation
So the tendency to overimitate may itself have about an adaptively significant feature of the environ-
evolved—genetically, or perhaps even culturally [9]— ment may not be detected or detectable in the course
for the sake of transmitting behavioural phenotypes ofindividualdevelopment.Culturalselectiongivesrise
inareasonablyhigh-fidelitywaydownthegenerations. totraitsthatwouldcountasnon-innateonthe‘learning’
For example, the connection between food prep- definition,butforwhichthestandardinferencesdrawn
aration practices and nutritional outcomes may be so aboutnon-innatetraitswillgoawry.
stochasticandvariablethatitisrealisticallyimpossible Other forms of cultural transmission depend to a
for a single individual to keep track of which practices greater extent on individuals being able to learn for
have positive and negative impacts on fitness. But if themselves [57]. For example, the most successful
lineagesofindividualstendtoadoptthesamepractices strategies in the social learning strategies tournament
astheirparents,thenmorebeneficialvariantswillpro- would try out observed behaviours for themselves
liferate on average, because of their positive effects on and only add the behaviour to their repertoire if they
thefitnessoftheindividualswhohavethem.Similarly, themselves received a benefit for the behaviour [8].
theconnectionbetweenactionandoutcomemaybetoo Thatwasamoreefficientstrategythantryingoutabe-
long-term for an individual to detect it for themselves haviour at random, because behaviours at high
(cf. the link between smoking and cancer, which it frequency in the social environment were more likely
tookmodernepidemiologicalstatisticstodetect). to be beneficial than a strategy picked at random.
Thus, given the right evolutionary conditions, the That is to say, the frequency of behavioural variants
typeofbehaviouraldispositionsfoundinapopulation, in the social environment carries information about
which have been transmitted by overimitation, will whatisadaptivelyuseful.Howisthatinformationgen-
carry information about past environments. Those erated? In part through individual learning: high
thatwerebeneficialtofitnessinpastenvironmentswill frequency behaviours are those that other individuals
befoundathigherfrequency.Totheextentthatthecur- have tried out and found to be rewarding. But part
rent environment is relevantly similar to those past of the information is generated by selection, because
environments,thecurrentpatternofbehaviouraldispo- behavioural dispositions are inherited from parents
sitions carries adaptively-relevant information about and the payoffs obtained by an individual affect the
features of the environment. That information has not number of offspring they have.
beengeneratedbyindividuallearning,norbyanyindi- Thatmixischaracteristicoftheusefulnessofcultu-
vidual being able to detect the adaptively-relevant rallytransmittedinformation.Someisinformationthat
featureoftheenvironmentfor themselves.Rather,itis others have learnt for themselves, so the imitator is
selection over evolutionary time that has built up the short-cutting the need to learn something that they
information in the population. When a new individual could,ifnecessary,havelearntfromtheirenvironment.
adoptsabehaviouraldispositionbyoverimitationfrom Otheraspectsofculturallytransmittedinformationare
its parents, it is relying on adaptively-relevant infor- generatedinthetransmissionprocessitself,asaresult
mation that has been generated by selection. But of selection. In the latter sense individuals are doing
unlike genetic representation, the information carried more than ‘standing on the shoulders’ of the learning
in the overimitation-based cultural inheritance system done by earlier individuals; cultural transmission is
(ifthereisone)isnotpresentatbirth. giving rise to adaptively relevant information that is
Thatunderminesacleandistinctionbetweeninher- more than the sum of the individual learning of the
ited information that is present at birth and acts as individuals whomakeup the culture[22].
a constraint on the developmental trajectory (e.g.
genetic representations), andadaptively-relevantinfor-
mation that individuals learn for themselves in their (b) A central role for culturally transmitted
own lifetime. Culturally transmitted information is information
acquired by individuals in the course of theirown life- Theargumentthatculturaltransmissionplaysacentral
time, but individuals do not learn for themselves that roleingeneratingadaptively-relevantinformationfaces
it is adaptively-relevant; indeed, they may not be able twoobstacles.Thefirstisatensionbetweenindividuals
todoso.Onanyreasonableview,therichlyinteractive learning for themselves, and cultural transmission
processbywhichthesetraitsdevelopcountsaslearning. building up information through natural selection. As
So if innateness were simply equated with being noted earlier, the ability of individuals to learn for
unlearnt, as suggested earlier, these traits would not themselves and adopt the behaviour that seems best
count as innate. That would overlook the fact that tothem,fromtheirownlimitedexperience,introduces
theysharemanypropertieswithgene-basedadaptations noise into the system for cultural transmission of be-
that are paradigmatically innate. Their development is havioural phenotypes. As Godfrey-Smith argues [22],
likelytoberelativelyfixedinthefaceoftherangeofvari- the more transmission depends on the intelligence of
ation encountered in the developmental environments the individuals, the less ‘Darwinian’ the process will
inwhichtheywereselected.Indeed,theirdevelopment be. It will be less well-characterized by the models of
may be have become canalized against environmental quantitativegenetics, forexample.
variation by means of other culturally or genetically However, Godfrey-Smith argues that this at most
inherited processes (gene–culture coevolution in the undermines the applicability of Darwinian micro-
second case). And since the adaptively-relevant evolutionary models to the process. At the macro
Phil.Trans.R.Soc.B(2012)
Review. New thinking and innateness N. Shea 2241
level, phylogenies of cultural practices can be highly benefits outweigh within-group relative fitness costs).
illuminatingevenifthetransmissionprocessisnothing If there is strong group selection then the fitness
likehigh-fidelityindividual-to-individualcopying[58]. advantageofbeing inagroupthatsharesinformation,
Also,atanintermediatelevelthatGodfrey-Smithcalls vis-a`-vis individualsin lessinformationallyopen social
‘meso-evolution’, a Darwinian approach can explain groups,mayoutweightherelativefitnessbenefitstobe
theoriginofcomplexadaptivetraits, evenwith intelli- garnered by selfishly keeping useful information to
gent transmission, since it explains the spread of oneself. Although there is widespread scepticism
adaptivelybeneficialphenotypes,whosehighfrequency about the importance of group selection in other
in turn increases the chances of the next cumulative species, it may well have been a particularly powerful
step of adaptive progress being discovered by an indi- influence in recent hominin evolution [61,62]. There
vidual in the population. In short, the mix of isarchaeological,palaeontologicalandanthropological
individuallearningandinformationproducedbyselec- evidence thathumans livedintight-knitsocialgroups,
tion, identified above, is compatible with a broadly perhapsextendedkingroupsinthefirstinstance,with
Darwinianapproachtounderstandinghowindividuals importantvariationsbetweengroupsinculture,behav-
manage to produce phenotypes that adaptively match ioural practices and even way of life. Powerful group
theirenvironment. selection would make a tendency to transmit adap-
The second problem concerns the sharing of infor- tively relevant information throughout the social
mation. Developing individuals’ use of culturally group less puzzling.
transmitted adaptively-relevant information found In sum, the following factors have probably played
within their society is a form of information sharing. a role in recent human evolution: (i) high-fidelity
How can such information sharing practices be evolu- transmission processes that closely model genetic
tionarilystableinthefaceofthefitnessadvantagethat micro-evolution; (ii) more diffuse ways that cultural
could be garnered by an individual in defecting— transmissioncanbuildupadaptively-relevantinforma-
keeping a piece of useful information to themselves tion through selection; and (iii) cultural transmission
[2]? Natural selection runs off fitness differences; so as a means for disseminating and preserving useful
if keeping information to oneself has costs, both to information that individuals have learnt for them-
oneself and others, then the selfish strategy will be selves, allowing subsequent individual learning to
selected if the costs to others are higher than to one- ‘stand on their shoulders’. The relative importance of
self. For example, there are obvious advantages to these factors remains an open empirical question.
sharing information about good hygiene. If others
adopt a hygienic practice that I have discovered, that
will on average reduce the overall risk of disease in (c) No clustering of i-properties for human
the population, and so will indirectly reduce my own psychological phenotypes
disease risk. But suppose I can get some benefit to These considerations mean that human development
myself from the hygienic practice, and suppose also islikelytobeparticularlyreliantonaninterestingcate-
thatitispracticaltokeepittomyself.Then,althoughI goryofadaptively-significantinformation:information
woulddobetterinabsolutefitnessifIweretosharethe whoseutilityindividualsarenotwhollyresponsiblefor
practice,IwilldobetterinrelativefitnessifIkeepitto detecting for themselves, but information which,
myself. I gain a relative fitness advantage since the risk unlike more familiar forms of inherited representation
of disease, although higher for all, falls more on others (genetic, epigenetic), is picked up by the individual
thanitdoesonme. Whythenshareusefulinformation? duringherownlifetime.Thatdecisivelyseversanycon-
Part of the answer may lie in the inherent difficulty nectionbetweennaturalselectionbeingresponsiblefor
of keeping some of these forms of information secret. the adaptive match between phenotype and environ-
Behaviours are observable, and a behavioural pattern ment, and the adaptively-relevant information being
that must be performed repeatedly is likely to be present at birth (in the zygote) and unlearnt. When
observed by those in the same social group. A restric- the developing individual absorbs a culturally trans-
tiontoverticaltransmissionofbehaviouralphenotypes mitted practice, or is steered by some culturally
wouldalsohelp.Ifparentsareonlyinclinedtodemon- transmitted factor towards an adaptive phenotype,
strate useful behaviours to offspring and close kin, those are clearly cases of getting information from the
then they derive a direct fitness benefit from sharing environment. The information carried by these kinds
useful information. Interestingly, there is evidence of inherited representations is not information about
thatoverimitationismorecommonin‘naturalpedago- which a poverty of the stimulus argument would suc-
gical’ contexts [59,60]. So far natural pedagogy has ceed. It comes from (social) stimuli. The rough
mostly been studied as a modifier of when children clustering of the i-properties in other species was
are disposed to learn. If it also restricts when adults underpinned by the fact that, if an individual did not
areinclinedtoteach,thenitcould,intheevolutionary learn the adaptively-relevant information for them-
past,haveconstitutedasystemwherebylineagesofbe- selves, the information embodied in the adaptive
havioural phenotypes transmitted by overimitation match between a developmental outcome and the
roughly aligned with parent–offspring lineages, and environment is very likely to have been generated by
hence with biological fitness. natural selection on genes, and to have been trans-
A restriction to roughly vertical transmission will mitted to this individual genetically, in factors that
notcoveranythinglikeallofthecases.Amoreimpor- arepresentinthezygoteandactsoastobiasthecom-
tant part of the answer may lie in group selection (in plex interactive pathways of development towards the
the specific sense that between-group relative fitness adaptive phenotype. Even that rough clustering will
Phil.Trans.R.Soc.B(2012)
2242 N. Shea Review. New thinking and innateness
break down if the New Thinking is right about the that argues against those claims. But the difference
importance of cultural transmission. between the twoviewpoints cannot be usefully encap-
What about the universality or species-typicality of sulated using the concept of innateness. The concept
adaptive traits, that is presupposed by the concept of of inherited representation furnishes a clearer distinc-
innateness?Recallthat,whenfacedwithadaptivepoly- tion. Evolutionary Psychology holds that genetic
morphisms in other species, I argued that there is representation is by far the most important type of
plausibly a shared developmental program, designed inherited representation responsible for human cogni-
to be guided by environmental information so as to tive phenotypes. The New Thinking instead sees
producedifferentoutcomesindifferentenvironments, culturally transmitted inherited representations as an
each more adaptive in the environment in which it is important source of the adaptively-relevant infor-
produced. That does not carry over to the human mation presentin human psychological phenotypes.
case either. Or at least if it does, it is only in The characteristics that the New Thinking identifies
the most tenuous way. The New Thinking suggests as distinctive of human cognitive evolution also put
that the ‘developmental program’ underpinning a pressureon thecoherenceof theconceptof innateness,
huge swathe of polymorphic adaptive phenotypes is asitisappliedtohumanpsychology.Theproblemisnot
humans’ highly adapted capacity for plasticity itself. just that some of the inferences about innate traits that
That is a very domain-general ability. Of course, it would go through in other biological domains fail in
too carries adaptively-relevant information—that the the case of human psychology. It is that there is no
environment is likely to be very variable, that these good proposal for a theoretical reconstruction of the
are good ways to engage in learning—but what is concept of innateness that would explain why any sig-
absent is a more specific developmental program nificant portion of the inferences about human
designed by evolution for a particular problem or set psychological traits standardly made using the concept
of problems. In short, extreme plasticity in some of innateness should be reliable. If that is correct, then
cases drives the nature of the potentially universal one important mistake of Evolutionary Psychology,
adaptation to the very highest, most general level. which the New Thinking has transcended, is to make
In other cases cultural inheritance may form the use of the concept of innateness. In contrast the New
basis of universal, developmentally robust or even Thinking manages to be deeply evolutionary, while not
environmentally canalized human phenotypes. being dependent on the concept of innateness at all.
In sum, an implication of the direction taken by the That might be thought to leave a lacuna. How can
NewThinkinginthiscollectionisthatthegeneralization theNewThinkingaccountforthekindsofphenomena
that makesthe i-properties cluster together roughly for that innateness claims have been thought to explain?
non-human adaptations, giving the concept of innate- In this paper I have focused on one explanandum in
ness some utility there, is absent in the case of recent particular: the adaptive match between phenotype
human behavioural adaptations. There is no particular andenvironmentachievedastheendresultofdevelop-
reason to expect those human behavioural phenotypes ment.Whenthatmatchisnotwhollyexplainedbythe
thatareplausiblyadaptationstohavethefollowingfea- individualaccessingadaptively-relevantinformationin
tures: to be due to natural selection on genes; to be its own experience, having detected its adaptive rel-
unlearnt, or to develop such that aspects of the adap- evance itself, we are left with a puzzle: where did the
tively-relevant information on which they depend are informationcome from?
not found in the developmental environment (such Inherited representation provides an answer, point-
that a poverty of the stimulus argument could be ing to the multiple routes by which information
made); to have a developmental course that is particu- generatedbynaturalselectioncanconstrainindividual
larly invariant or canalized against environmental development towards an adaptive phenotypic out-
variations;or tobebasedonauniversaldevelopmental come, so that aspects of the adaptively-relevant
program, save the most domain-general capacities for information carried by the phenotype need not
plasticity,learningandculturalinheritanceitself. derive from individual experience. Genetic and epige-
netic mechanisms can play that role, giving rise to
adaptively-relevant information that is present at
4. THE NEW THINKING ARTICULATED IN TERMS birth;butwealsosawthreeculturalroutestoadaptive
OF INHERITED REPRESENTATION phenotypes,innoneofwhichistheadaptively-relevant
This paper has argued that recognizing the semantic information present at birth, but which involve learn-
information carried by genetic transmission down the ing (albeit that individuals do not detect the adaptive
generations—genetic representation—allows us to see significance of all the information for themselves).
whypropertiesconnectedtotheconceptofinnateness Althoughculturalprocesses,learningandinteractive
dotendtocluster togetherwhenappliedtomanybio- development are central, the New Thinking is also
logical adaptations. The utility of the innateness thoroughly evolutionary, relying strongly on natural
concept in those contexts makes it tempting to selection in explaining human psychology, unlike
import it into our account of human evolution. High blank slate empiricism and the standardsocial science
church Evolutionary Psychology does indeed claim model. Phylogenetic methods and cross-species
that human psychological capacities are innate in a comparisonsarereliedonheavily,aspartofthecareful
closely parallel sense: a suite of domain-specific adap- accumulation and assessment of all available evidence
tations produced by natural selection on genes, that about the hominin and primate past. The importance
are unlearnt, developmentally fixed and cross-cultu- that the New Thinking attaches to historical evidence
rally universal. The New Thinking points to evidence shows just how serious it is about evolutionary
Phil.Trans.R.Soc.B(2012)
Review. New thinking and innateness N. Shea 2243
explanation. However, using the concept of innateness 16 Griffiths,P.E.,Machery,E.&Linquist,S.2009Thever-
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