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Laland et al 2010 - How Culture Shaped the Human Genome

Laland et al 2010 - How Culture Shaped the Human Genome

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Published by: Jerry Zee on Apr 08, 2011
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02/06/2013

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Accts f ha elti feqetl asse thatthe selectie eets that shaped s wee chages i theexteal eiet, steig f eets bedha ctl. F istace, theies f the icepti f 
Homo
species ephasize a glbal ted twads cle,die cliates, which pshed a abeal ape t f c-tactig fests it saaah
1
. Likewise, heat stess ithe pe is a plasible hpthesis f the elti f bipedalit, hailess ski ad sweatig
2–4
. B ctast, lit-tle csideati has bee gie t the pssibilit thatcltal pactices ight hae tasfed the selectipesses actig  has. This a be becase it hasl ecetl bee shw that atal selecti ca bigabt sbstatial chages i gees that ae detectablee thsads f eas, as eealed b easeets f the tpical ates f espse t atal selecti agaials i the wild
5
ad statistical aalses that detectedecet apid adaptati i the ha gee
6–11
.This taditial ccepti f ha elti isw beig challeged b ecet athplgical stdiesthat shw that ha cltal pactices hae difiedeietal cditis, tiggeig chages i allelefeqecies
12,13
. I additi, aalses f data f theha gee hae eealed es gees that haeexpeieced ecet psitie selecti, a f whichexhibit fctis that ipl that the ae espses tha cltal pactices
6–11,14
. F istace, seeal liesf eidece shw that dai faig ceated the selec-tie eiet that faed the spead f alleles fadlt lactse tleace
12,13,15,16
. Estiates f the bef ha gees that hae bee sbject t ecet apidelti age f a few hded t tw thsad:Willias
et al.
14
cclde that p t 10% f the hagee a be affected b likage t tagets f psi-tie selecti. Althgh i the ajit f cases it is tkw what phetpe was the taget f the ifeedselecti,  which eietal cditis faedsch phetpes, ha cltal pactices eai stgcadidates, ad geeticists ae iceasigl csideigclte as a sce f selecti  has
17,18
.Sch data ae csistet with tw baches f atheatical eltia aalsis: gee–clte c-eltia the, which exples hw geetic adcltal pcesses iteact e eltia tie
19–23
,ad iche-cstcti the 
24–30
, which iestigatesthe eltia ipact f the dificati f ei-ets b gaiss. The dels pide hptheses f, el isights it, the elti f leaig, clte,lagage, itelligece, cpeati, sex diffeeces adatig sstes. Aalses f these dels hae c-fied that gees ad clte cld plasibl c-ele,fte eealig pattes ad ates f chage that aechaacteistic f e taditial pplati geeticthe 
22,31–34
. Gee–clte daics ae tpicall faste,stge ad peate e a bade age f cditistha cetial eltia daics, leadig sepactities t age that gee–clte c-elticld be the diat de f ha elti
32–34
.
*School of Biology, Universityof St Andrews, Bute Building,Westburn Lane, St Andrews,Fife KY16 9TS, UK.
School of Anthropology,University of Oxford, 51/53Banbury Road, Oxford OX2 6PE, UK.
§
Institute for GenomicDiversity, Cornell University,Ithaca, New York14853‑2703, USA.
||
Biology Department,Acadia University, Wolfville,Nova Scotia B4P 2R6,Canada.Correspondence to K.N.L.e‑mail:knl1@st‑andrews.ac.uk
doi:10.1038/nrg2734
How culture shaped the humangenome: bringing genetics andthe human sciences together
Kevin N. Laland*, John Odling‑Smee
and Sean Myles
§ ||
Abstract | Researchers from diverse backgrounds are converging on the view that humanevolution has been shaped by gene–culture interactions. Theoretical biologists haveused population genetic models to demonstrate that cultural processes can have aprofound effect on human evolution, and anthropologists are investigating culturalpractices that modify current selection. These findings are supported by recent analysesof human genetic variation, which reveal that hundreds of genes have been subject torecent positive selection, often in response to human activities. Here, we collate thesedata, highlighting the considerable potential for cross-disciplinary exchange to providenovel insights into how culture has shaped the human genome.
REVIEWS
nATurE rEvIEWS
|
 
Genetics 
voLumE 11
|
FEBruAry 2010
|
 
137
© 20 Macmillan Publishers Limited. All rights reserved10
 
The peise f this aticle is that cltal pacticeshae shaped the ha gee. We fthe sggest thata gee–clte c-eltia pespectie pidespptities t itegate fidigs f ha geet-ics ad eltia the with athplgical adachaelgical data, geeatig el hptheses adltiatel esltig i a e cpehesie de-stadig f ha elti. Piet eseacheshae called f a itedisciplia pject alg theselies
17,18
. This pespectie als ffes el hptheses,ethds ad explaat echaiss with whicht destad ha geetic aiati ad aspectsf ha iqeess, ad helps t explai secflictig fidigs.We begi b descibig theetical dels f gee–clte c-elti, tliig the isights that thesehae geeated. This is fllwed b a descipti f theathplgical eidece f gee–clte c-elti.We the discss geetic stdies that hae idetifiedlci that hae bee sbject t ecet selecti, addescibe hw the cclsi eached b geeticists, thata f these selectie sweeps a be i espse tha actiities, fits with bth the fal the adathplgical eidece. Fiall, we cside hwthese diese appaches ca be bette itegated, adexple the iplicatis f eseaches iestigatigthe ha gee.
Models of gene–culture co‑evolution
The aget that gees ad clte c-ele waschapied e 30 eas ag b piees f the field f ‘gee–clte c-elti’, a bach f theetical pp-lati geetics
19–23,31,33–35
. These eseaches iew geesad clte as tw iteactig fs f iheitace, withffspig acqiig bth a geetic ad a cltal legac f thei acests. Geetic ppesities, expessedthght deelpet, iflece what cltalgaiss lea. Cltall tasitted ifati,expessed i behai ad atefacts, speads thghpplatis, difig selecti pesses that act back  pplatis (see
BOX 1
 
f a defiiti f clte).Gee–clte c-eltia aalses tpicall bild  cetial pplati geetic the. Iadditi t tackig hw allele  getpe feqe-cies chage i espse t eltia pcesses, schas selecti ad dift, the aalses icpate cltaltasissi. A wked exaple f the delligappach is shw i
BOX 2
, which addesses the apidspead i East Asia f a ai-acid-alteig aiati ectdsplasi A ecept
 
(
), a gee iled ihai phlg. Gee–clte aalses exple hwleaed behai c-eles with alleles that affectthe expessi ad/ acqisiti f the behai whse fitess is affected b the cltal eiet.The appach has bee depled t exple the adaptieadatages f eliace  leaig ad clte
21–23,33,36
, tiestigate the iheitace f behaial ad pesal-it taits
37–39
, ad t iestigate specific tpics i haelti, sch as lagage  cpeati
(TABLE 1)
.These aalses hae bee ail cdcted withta specific kwledge f the ha gees iled;athe, hpthetical gees hae bee ppsed withassed fctis, ad the daics f thei c-eltiwith cltal taits hae bee expled. I spite f this,thee exists a spisig celati betwee the tp-ics addessed i these aalses ad the gees that aew kw t hae bee sbject t ecet selecti(discssed belw). I
BOX 3
, we shw that gee–cltec-eltia the pides pssible explaatisf wh gees expessed i the exteall isible phe-tpe (f exaple, hai ad ee cl) ight be likel tagets f selecti. These exaples pide attactiehptheses f fthe iestigati becase the ela-tiship betwee gee ad phetpe is cpaatiel well established.
Effects of culture on selection.
Gee–clte c-eltiists iew clte as a daic pcess that cashape the ateial wld
21,22,33,34
. Thei dels hae estab-lished that cltal pcesses ca daaticall affect theate f chage f allele feqecies i espse t selec-ti, seties speedig it p ad seties slwigit dw. recet estiates f the cefficiet f selectiassciated with selected ha gees expsed t clt-all dified selecti pesses eeal extadiail stg selecti. The lactse-tleace allele has speadf lw t high feqecies i less tha 9,000 eas sicethe icepti f faig, with a estiated selecti cef-ficiet f 0.09–0.19 f a Scadiaia pplati
40
. Sch
Box 1
|
What
 
is culture?
To the layperson, the term ‘culture’ typically evokes images of fine art and fashion,but historically anthropologists have characterized culture as the complex of beliefs, values, behaviour and traditions associated with a particular population.
 
Neither notion is particularly conducive to scientific analysis.
 
Human culture hasproven a difficult concept to pin down, and there exists little definitional consensuswithin the social sciences
12
.
 
In this vacuum, geneticists and biologicalanthropologists, eager to explore how cultural phenomena interact with genes,have taken a pragmatic line to studying culture.
 
For these researchers, culture isinformation that is capable of affecting individuals’ behaviour, which they acquirefrom other individuals through teaching, imitation and other forms of sociallearning
33
. Here, ‘information’ includes knowledge, beliefs, values and skills. Culturalchange can then be modelled as a Darwinian process comprising the selectiveretention of favourable culturally transmitted variants, as well as variousnon-selective processes, such as drift
20,21
. Rather than attempting to describe theentire culture of a society, culture is broken down into specific traits (for instance,milk users or non-users, or consumption of a starch-rich or starch-poor diet), whichallows their frequencies to be tracked mathematically.This broad characterization opens up the possibility of culture in other animals,and indeed traditions for exploiting prey or food sites, tool-use and vocalizationshave been reported in a variety of animals, including fish, birds, cetaceans andnon-human primates
93
. These traditions exhibit several properties of interest tobiologists
93
. Perhaps the most obvious is that culture is a source of adaptivebehaviour; individuals can efficiently acquire solutions to problems, such as ‘what toeat?’ and ‘with whom to mate?’, by copying others. But a variety of studies, rangingfrom investigations of fish mating sites to human foraging traditions, haveestablished a capability of culture to propagate behaviour in a manner that is tosome degree independent of the ecological environment. Culture can also generatepatterns of phenotypic variation in space: clines in behavioural characteristics havebeen reported for orang-utan behaviour, birdsong and whale vocalizations.Moreover, these traditions modify the action of selection. For instance, modelssuggest that song learning in birds affects the selection of alleles influencing songacquisition and preference, can facilitate speciation and can lead to the evolutionof brood parasitism
94–96
.
REVIEWS
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© 20 Macmillan Publishers Limited. All rights reserved10
 
bseatis, cbied with the shallw tie-depthassciated with a ecetl selected ha gees
6
,aises the qesti: cld cltall deied selectipesses be stge tha -cltal es?The aswe is es, f tw eass. Fist, cltalpcesses cc b ite f acqied kwledge ca-ied i ha bais that is fte eliabl tasittedbetwee idiidals. Althgh thei cstac aiesf tait t tait, thee is eidece that cltall d-ified selectie eiets ae capable f pdcigsall stg atal selecti that is highl csist-et i diectialit e tie
40
. Secd, a gees aefaed as a eslt f c-eltia eets tiggeedb phetpic chages i the species,  i espse tthe gee-feqec chages i thei gees. Whechages i e geetic tait die chages i a secd,the ate f espse i the latte depeds i pat  theate f chage i the fe, which, as a le, is t fast.I cpais, ew cltal pactices tpicall speade qickl tha a geetic tati, sipl becasecltal leaig peates at faste ates tha bilgi-cal elti
22
. If a cltal pactice difies selecti ha gees, the lage the be f idiidalsexhibitig the cltal tait, the geate is the itesit f selecti  the gee. A apid spead f the cltalpactice leads e qickl t axial itesit f selec-ti  the adatages geetic aiat(s). The effectf these facts has bee epeatedl destated b gee–clte c-eltia dels, which csist-etl ept e apid espses t selecti tha c-etial pplati geetic dels
21,22,31,41,42
. This a help t explai the aget that clte has ‘apedp’ ha elti
17
, althgh the facts ae alslikel t hae a le, sch as the iceased be f ew tatis i the lage pplatis that hae beefacilitated b agiclte
17,43
.Hwee, eqall iptat is the bseati thatcltal selecti pesses a feqetl aise adcease t exist faste tha the tie eqied f the fixa-ti f the assciated beeficial allele(s). I this case,clte a die alleles l t iteediate feqec,geeatig a abdace f patial selectie sweeps
44
.Cp
et al.
44
sggest that cplete selectie sweeps atsigle lci i has a be c ad that adap-tatis e the past 70,000 eas a be piail theeslt f patial selectie sweeps at a lci.
Niche‑construction theory
niche-cstcti the is a bach f eltia bilg that ephasizes the capacit f gaiss tdif atal selecti i thei eiet adtheeb act as c-diects f thei w ad the spe-cies’ elti
24,45
. Exaples iclde aials afac-tig ests, bws ad webs ad plats difig
Box 2 |
Constructing
 
a mathematical model of gene–culture co‑evolution
To illustrate gene–culture co-evolutionary theory, we present a worked example inspired by the observation that afunctional non-synonymous substitution in the ectodysplasin A receptor
 
(
EDAR
) gene that is associated with thickerhair (as well as with changes in the skin, teeth and sweat glands) has experienced strong recent positive selection inEast Asia
9,97
. How could cultural practices explain this selection? One possibility is a gene–culture co-evolutionaryversion of sexual selection
(BOX 3)
. Here, we show how it is possible to quantify the changes in frequencies andinteractions between culturally transmitted mating preferences and genetic variants of 
EDAR
using gene–cultureco-evolutionary theory.For simplicity, we consider two culturally learned preferences, labelled
h
and
H
:
h
individuals mate at random withrespect to hair thickness (and other phenotypic effects of 
EDAR
), whereas
H
individuals preferentially choose as matespartners with thicker hair. We also consider a single diallelic locus that has alleles
A
1
and
A
2
:
A
1
A
1
individuals possessthin hair,
A
2
A
2
individuals possess thicker hair and the hair of 
A
1
A
2
individuals is a function of the dominance parameter
 d
(0 <
d
< 1). That means there are six possible combinations of genotype and mating preference, known asphenogenotypes:
A
1
A
1
h
individuals (who have thin hair and no preference for hair thickness in their partners), andsimilarly
A
1
A
1
H
,
A
1
A
2
h
,
A
1
A
2
H
,
A
2
A
2
h
and
A
2
A
2
H
individuals, to which we allocate frequencies
x
1
–x
6
. We assume that
H
 individuals choose
A
1
A
1
,
A
1
A
2
and
A
2
A
2
mates with frequencies (
x
1
+
x
2
)/
z
, (1 +
α
h
)(
x
3
+
x
4
)/
z
and (1 +
α
)(
x
5
+
x
6
)/
z
, in which
z
= 1 +
 
α
h
(
x
3
 
+
x
4
) +
α
(
x
5
+
x
6
). If mating preferences are handed down from parent to offspring, we can specify fourclasses of matings, namely
h
mothers with
h
fathers (or
h
×
h
), and likewise
h
×
H
,
H
×
h
and
H
×
H
matings, which give rise toproportions
b
3
,
b
2
,
b
1
and
b
0
offspring of type
h
(and 1 –
b
3
, 1
b
2
, 1
b
1
and 1 –
b
0
of type
H
). It is apparent that a mating of two
A
1
A
1
h
individuals would occur with frequency
x
12
and give rise to
A
1
A
1
offspring, proportion
b
3
of which would be
h
.We could similarly calculate the expected proportions of offspring in each phenogenotype category for each of the36 possible phenogenotype matings. Weighting these proportions by mating frequencies, summing across matingsand multiplying by phenogenotype fitness would give us the expected proportion of each phenogenotype in theoffspring generation. This leads to a system of six recursive equations, which specify the frequencies of eachphenogenotype in the offspring as a function of their frequencies in the parents, as well as the parameters
d
,
b
3
b
0
 and
α
. This system can then be analysed using conventional mathematical methods to investigate whether theappearance of a cultural preference for
H
could explain the rise in frequency of the
EDAR
allele, and to specify theconditions under which this occurs. Methods are available to incorporate assortative mating, learning from non-kin, avariety of cultural transmission biases (for example, conformity) and demographic effects.Note the above logic applies irrespective of whether it is hair thickness or some other character (for example,
REF. 98
)that is the focus of sexual selection, and similar models can be devised for naturally selected traits. The importantgeneral point here is that, because of its dynamic interactive properties, culture typically cannot adequately betreated as a background condition for selection, and must be incorporated into a dynamic model if the analysis is tobe accurate. For a more detailed introduction, see
REFS 20,21,41
.
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