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SIDCC !HC HC!IIC UC!dIS O !HCII DOSSIDIIIV SDdCCS dIC DOHHU !O DC
UICICHI. P VCIV IHDOItdD! CXdHDC O IHCSC IDVdIIdH! DIODCI!ICS IS
IHC CXIS!CHCC dHU UISIIIDHIIOH O SDCCId OI ICHdIKdDC DOIHIS |OI SCIS
O SHCH DOID!S} CdCU sinjulariris. HIS IS !HC COHCCDI !Hd! WC HCCU IO
HdKC !HC ICHdIKS IH !HC DICVIOHS DdId0IdDH CSS HC!dDHOIICd. IHC
DOSSIDII!ICS WIIH IHC HI0HCS! DIODdDII!V O OCCHIIID0 dIC IODOO0ICd
SIH0HdDIICS dC!IH0 dS dI!IdCIOIS.
LC!S HOW dDDV IHIS IHC O !HOH0H! !O Hd!HCHdIICd HOUCS. O
CCd!C d Hd!HCHd!ICd HOUC !HC DIS! SICD b IO CHHHCIdIC d !HC
ICCVdHI WdVS ID WHICH IHC DIOCCSS !O DC HOUCCU IS ICC !O CHdH0C.
1NL b1KM N 1NL LMU1LK
LCIS IHd0IHC IHdI WC dIC HOUCIH0 d SIHDC DHVSICd DIOCCSS CHdIdC
ICIIZCU DV IWO CHdH0IH0 DIODCI!ICS, SHCH dS ICHDCIdIHIC dHU DICSSHIC.
HCSC dIC CdCU IIS UC0ICCS O ICCUOH. HC DIOCCSS HdV, O COHISC,
dSO CHdH0C IH dH IHDDIIC HHHDCI O IIICCVdHI WdVS, IHC dI! O HdIH
CHdIICd HOUCIH0 DCIH0 DdSCU ID DdII OH IHC dDIIIV IO |HU0C WHdI
CHdH0CS UO, dHU WHdI CHdH0CS UO HOI, HdKC d UICICHCC. LHCC IHC
ICCVdHI UC0ICCS O ICCUOH O d DHVSICd DIOCCSS HdVC DCCH IUCHIIDCU
IHC HOUC CdH DC 0IVCD d SDdIId OIH DV dSSI0HID0 CdCH O IHCH IO
d UIHCHSIOH O d IODOO0ICd SDdCC. LdCH DOIHI IH IHIS SDdCC WI DC d
COHDIHdIIOH O VdHCS O ICHDCIdIHIC dHU DICSSHIC ICDICSCHIIH0 dH
IHSIdHIdHCOHS S!d!C O !HC DIOCCSS DCIH0 HOUCCU, WHIC IHC SCI O
DOIDIS dS d WHOC ICDICSCHIS IHC SDdCC O d DOSSIDC SIdICS OI IHC
DIOCCSS. OI IHISICdSOH !HC dDSIIdCI SDdCC IS ICCIICU IO dS srarsrac
|OI DHdSC SDdCC}. IHdV, SIHCC d 0IVCH DIOCCSS CHdH0IH0 ID IIHC
OOWS d SCCHCHCC O S!dICS IIS DCHdVIOI dDDCdIS IH SId!C SDdCC dS d
SCIICS O DOIHIS, !HdI IS, dS d CHIVC OI IIdjCC!OIV. I WdS DV ODSCIVIH0
IHC ICHUCHCV O HdHV O IHCSC IIdjCC!OIICS IO COHVCI0C OH SDCCIDC
dICdS O SId!C SDdCC, IO COHVCI0C OH SID0HdIIIICS, IHdI IHC CXISICHCC O
dSVHDIO!IC SIdDIIIV WdS DISI CSIdDISHCU. '`
1SIH0 !HCSC IUCdS IHC CXDdHdIIOH OI !HC HDICdSOHdDC CCCIIVC
HCSS O HdIHCHdIICS CdH DC DHIdSCU IKC IHIS. d HdIHCHdIICd HOUC
CdD CdDIHIC !HC DCHdVIOI O d Hd!CIId DIOCCSS DCCdHSC !HC SDdCC O
DOSSIDC SOH!IOHS cvrlars !HC DOSSIDIIIV SDdCC dSSOCIdICU WIIH IHC
HdICIId DIOCCSS. HC IWO DOSSIDIIIV SDdCCS HCCU HO! DC IUCHIICd DHI
HCICV OVCIdDDID0 DCCdHSC HOS! HdIHCHdIICd HOUCS HIHIC !HC
DCHdVI0I O d DIOCCSS OHV WI!HID d CCI!dID IdD0C O VdHCS O IHCII
COH!IO DdIdHC!CIS. P SHDCICHI OVCIdD CdH HCVCI!HCCSS CXIS! DCCdHSC
!HC SIH0HdIIIICS !Hd! S!IHC!HIC DO!H SDdCCS dIC IDUCDCHUCH! O DOIH
CdHSd HCCHdHISHS IH d DIOCCSS dDU OIHd ICd!IOHS ID dH CGHd!IOH.
HIS CXDdHd!IOH IHDICS dD OHIOO0ICd COHHI!HCHI IO !HC dHIOHO
HOHS CXIS!CHCC O !ODOO0ICd SIH0HdII!ICS, OI HOIC 0CHCIdV, !O IHC
S!IHC!HIC O DOSSIDII!V SDdCCS. LHCC IHIS OHIOO0ICd COHHI!HCHI HdS
DCCH HdUC !HC ICIH SIH0HdIIIV CCdSCS IO DC d DHICV Hd!HCHd!ICd
COHCCDI dDU DCCOHCS d DIODCIV DHIOSODHICd OHC. U DdI!ICHdI, OHCC
SID0HdIIIICS dIC !dKCH IO DC dS ICd dHU CDCICH! dS CdHSCS !HC HdIHIC
O IHCII ICdIIV DCCOHCS d DIODCH OI DHIOSODHCIS. O IHCV CXIS!, OI
CXdHDC, dS rranscndnrCH!I!ICS IH d WOIU DCVOHU IHd! O Hd!ICI dHU
CHCI0V: LI dIC IHCV immannr!O IHC Hd!CIId WOIU: d !HC Hd!!CI
IV
Z
NLbN N bMUL1N
and cncryo Ihc unvcrsc ccascd Io cxsI, vouId snuIarIcs aIso
dsappcar |mmancnI)orvouIdIhcyconInucIocxsI |IransccndcnI)?
AIIhouhIhcscqucsIons arcnoImaIhcmaIcaI buIphIosophcaIIhc
pracIccomaIhcmaIcans cansIII provdcnshIsnIoIhc ansvcrs.
I snuIarIcs arc mmancnI Ihcy musI bc boIh rrcducbIc I any
parIcuIarmaIcraIproccssvhIcaIIhcsamcImcrcqurnIhaIsomc
proccssor anoIhcr acIuaIIy cxsIs. Thcsc Ivo condIons arc rc!IccIcd
nIhc vay snuIarIcs arc sIudcd. TopoIosIs, or cxampIc, do noI
sIudyIhcsnuIarIcssIrucIurnIhcpossbIIyspaccomodcIrcc
Io chancnIsIcmpcraIurc andIsprcssurc, buIoallmcdwirhrwc
dorsc] ]dcm vhaIcvcr Ihcsc may bc.1 canbcprovcd, orcxam-
pIc,IhaInaIvo-dmcnsonaIspacconIyccrIanKndsosnuIarIcs
cxsI: our dcrcnI Iypcs o ponI snuIarIcs dsInushcd rom
cach oIhcr by Ihc orm o Ihc !Iov o ncarby IrajccIorcs |nodcs,
saddIcponIs, oc, and ccnIcrs) as vcII as oncIypc opcrodcsnu
IarIy. '" U Ihrcc-dmcnsonaI spaccs, Ihc our ponI snuIarIcs
arc sIII parI o Ihc rcpcrIorc buI nov pcrodc snuIarIcs comc
n IhrccdcrcnIorms: sIabIc, unsIabIc, andsaddIc-shapcdIoops.U
addIon, ancvIypcosnuIarIybccomcsavaIabIc, oncIhaIcanbc
pcIurcd as a Ioop IhaI has bccn rcpcaIcdIy sIrcIccd and oIdcd |a
so-caIIcd chaoIc aIIracIor).'' Ths mpIcsIhaI IopoIocaI acIs abouI
possbIIyspaccscanbcdscovcrcdvIhouIrccrcnccIoIhcnaIurco
Ihcdcrccso rccdm, onIyIo Ihcrnumbcr, andvIhouIrccrcncc
Io Ihc naIurc o Ihc radcnI |IhcrmaI, ravIaIonaI, mcchancaI,
hcmcaI) onIy Io Is cxsIcncc. uI Ihc acI IhaI Ihc xrnc o a
radcnI, any radcnI, sncccssary conhrms Ihc mmancnI sIaIus o
snuIarIcs.
SnuIarIcsarc,Ihcrcorc,pcrccIIyacccpIabIccnIIcsnamaIcr-
aIsIphIosophy.Thc man probIcm conronIn us novP Ihc cxIcnI
Io vhch vc can cncraIzc romIhcsc dcas. SIaIcspacc s onIyonc
Knd o possbIIy spacc, a spacc uscuI Io sIudy Icndcnccs buI noI
capacIcs. CapacIcs nvoIvc a much Iarcr scI o possbIIcs Ihan
IcndcnccsbccausccnIIcs cancxcrcscIhcr capacIcsnnIcracIon
vIh a poIcnIaIIy nnumcrabIc varcIy o oIhcr cnIIcs. Thc morc
compIc possbIIy spaccs assocaIcd vIh capacIcs, and Ihc naIurc
oIhc snuIarIcs IhaI sIrucIurcIhcm, arc noIncarIy asvcII undcr
sIoodasIhosc oIcndcnccs.On Ihc oIhcr hand, compuIcrs cansup
pIyIhcmcans IocxpIorcIhcscoIhcr possbIIy spaccs n a rorous
1NL b1KM N 1NL LMU1LK
vaybccauscIhcnIcracIonsnvhccapacIcs arccxcrcscdcanbc
sIacdn a smuIaIon andvarcd nmuIIpIc vaysunIIIhcsnuIar
caIurcso Ihc possbIIyspacc arc madcvsbIc. Eacho IhcoIIov-
n chapIcrs vII cxpIorc hov sIan a dcrcnI Iypc o smuIaIcd
nIcracIon |chcmcaI, boIocaI, socaI) can Icasc ouI Ihc snuIar
sIrucIurco Ihcr possbIIyspacs. Thc hrsI sIcp nIhscxpIoraIon,
hovcvcr, vI! noI addrcss Ihc qucsIon o Ihc rcIaIon bcIvccn
modcIsandrcaIIy. WchrsI nccdIo cIaryIhcconccpIocmcrcncc
n Ihc casc o smuIaIons. Ths s vhaI Ihc oIIovn chapIcr vII
aIIcmpIIodo.
ZI
ZZ
LHPJLJW
O||J|d| J|O0d|d d0 |d||O|5 O ||OW
Nuch IKc Ihc smpIicIy ol IcmpcraIurc and prcssurc maKc Ihcm
an dca!sIarIn poinI lorphiIosophca! rcLccIon oncmcrcncc, Ihc
qucsuon ol vhaI s an cmcrcnI propcrIy n compuIcr smu!aIons
can bc bcsI approachcd sIarIin vIh Ihc !casI comp!cx cascs. Thc
sm!csIol a!! compuIn machncs,]nirsraraurcmara. can pcrlorm
a compuIaIon by chanin lrom onc sIaIc Io anoIhcr n a vc!!-
dchncd scqucncc vIhouI havn Io sIorc nIcrmcdaIc rcsuIIs. IcIs
manc a popuIauon ol Ihcsc smpIc machincs cmbodicd physca!!y.
OncvayolIcIunIhcm nIcracI s Io crcaIc a!arc hshn ncImadc
olc!ccIrca!!yconducIn maIcraI and Io p!acc ahniIc sIaIc auIoma-
Ion vhcrcvcr Ivo vrcs cross. Thc inIcracIons vou!d IaKc p!acc
Ihrouhanc!ccIrcaIcurrcnI!!ovn IhrouhIhc vrcsaI!ovnIhc
sIaIc ol a vcn auIomaIon aI any onc Imc Io bc dcIcrmncd by
IhcsIaIcsolnchborin auIomaIaaccordnIo somc ruIc. Thcsamc
inIcracIin popu!aIon canbc crcaIcd vIh solIvarc nsIcad ol hard
varc l an absIracI Ivo-dmcnsonaI spacc s subdivdcd nIo "cc!!s
andasmu!aIcdhnIcsIaIcauIomaIonspIaccdincachcc!!. Thccc!!s
maybcIranu!ar, rccIanu!ar, hcxaona!, or anyshapc IhaIcxacI!y
I!csIhcp!ancsoIhaIIhccc!!sarcconnccIcdIooncanoIhcrbyshar-
incdcsandcorncrs.oIhIhcphyscaIrca!zauonolIhcpopu!aIon
ol auIomaIa and Is smu!aIon in solIvarc arc rclcrrcd Io as cllular
aurcmara. Thc qucsIonsvhcIhcrIhc samcsIraIcyuscd vIh Icm-
pcraIurcandprcssurcv!!su!!vorKvhcnvcrcp!accapopu!aIonol
mo!ccuIcsvIhoncolcompuInmachincs.ThaIs,vhcIhcrvccan
dcnulycmcrcnIpropcrIcs, Icndcnccs,andcapacIcs, andvhcIhcr
LLLLULK U1M1 N 11LKNb LW
vc can shov IhaI cmcrcnI cnuIcs aI onc sca!c can bc uscd Io
composccmcrcnIcnIIcsaI a !arcrsca!c.
Thc bcsI Knovn cxamp!c ol a cc!Iu!ar auIomaIon s Ihc so-ca!!cd
Camc]Lt].Thccc!!shousinhnIcsIaIcauIomaIa inIlc arcrccIan-
u!ar n shapc and Ihc auIomaIa arc capab!c ol bcn n on!y Ivo
sIaIcs, "on and "oll. Ths s usua!!y cxprcsscd as l Ihc Ivo sIaIcs
bcIoncdIoIhccc!!sIhcmsc!vcs vhcharcsadIobc"a!vcor"dcad,
rcspccIvcIy. Ilc s noI a rcu!ar compuIcr amc sincc Ihcrc P no
conunuousnIcracIonbcIvccnIhc uscrandIhccompuIcr: Ihc uscr
mcrc!y scIs Ihc sIaIc ol a lcv cc!!s |IhaI s, maKcs somc ol Ihcm
"a!vc ) andIhcn passvcIy vaIchcsasIhccc!!sallccIandarcallccIcd
by Ihcr nchbors. Thc nIcracIon ruIcs ol Ilc arc dcccpIivc!y
smpIc: laIa vcnmomcnIacc!!hasIvonchborsIhaIarc a!vc I
vi!IsIaynIhcsamcsIaIcIvasinIhc prcvousnsIanI,IhaIs, lIhc
ccI!vasaIivcIv!!sIayaIvcandl IvasdcadIvi!Ircmandcad: l
Ihccc!!sdcad andIhrccolIsnchborsarcaIivc, Ivi!I comc aIvc:
hna!!y, l Ihc cc!! s aIivc and onIy onc nchbor s aIivc, or l morc
IhanIhrcc arc, Iv!!dc. Thc ru!cs arc app!cdIo Ihc cnIrc popu!a
Ion smu!Iancous!y an rcpcaIcd!y lor as Ion as Ihc smu!aIon s
aI!ovcd Io !asI.Thc lacI IhaI Ihc nIcracuons arc rd!yspcOhcd by
ru!cs mpIics IhaI Ihcy arc ncr mrjnr. On Ihc oIhcr hand, as Ihc
popuIaIonnIcracIspaIIcrnsolsIaIcsnnchborinccI!sappcarand
IhcscarcndccdcmcrcnIsinccIhcpaIIcnshavcpropcrucs,Icndcn-
ccs, and capacIcsIhaI arcnoIprcscnIn Ihc indvdua! auIomaIa.
Thc mosI basc cmcrcnI Icndcnccs ol paIIcrns ol sIaIcs arc Ihc
samcasIhosclound nrca!physcaIproccsscs:IhcIcndcncyIorcmain
in a sIcady sIaIc and Ihc Icndcncy Io osc!!aIc bcIvccn sIaIcs. hcn
p!aynvIh Ilc onc qucK!y dscovcrs IhaI mosI nIaI paIIcrns dis-
appcar alIcr a lcvcncraIon.SoIhcIcndcncyIomanIain Ihcsamc
shapc, cvcn Ihouh hardIy a lorm ol bchavor duc Io Ihc abscncc

ol
chanc, snoIordnary.NanypaIIcnsin Ilcdisp!ayIhsIcndcncy,
IhcsmpIcsI oncbcncomposcdollournchbornccI!ssimu!Ianc-
ous!y aIvc, a paIIcrn ca!!cd a "b!ocK. Thcsc sIaIc buI cndurn paI-
Icrns arc caI!cd "su!! Iilcs. A sccond Uass ol paIIcrns, co!!ccIvcIy
Knovn as "osc!!aIors, dspIays a Icndcncy Io cyc!c Ihrouh a scrcs
ol conhuraIons ovcr and ovcr aan. Thc smpIcsI onc, ca!!cd a
"bIinKcr, conssIs ol Ihrcc nchborn cc!!s in a rov IhaI bccomcs
a coIumn in Ihc ncxI cncraIon and Ihcn bccomcs a rov aan.
ZJ
Z+
N LbN N b MUL1N
Thcb!nKcrsmp!ysvIchcsbacKandlor|hbcIvccn Ivophascs buI
oIhcrosc!!aIorsundcroanumbcrolchancsbclorcrcIunnIoIhc
nIa!phasc andrcpcaIn Ihcscrcs aan. AIhrdc!ass olpaIIcns,
ca!!cd "spaccshps, adds Io Ihs rhyIhmc bchavor a Icndcncy Io
movcnspacc. Thcsmp!csI onc smadcolhvc!vccc!!sarrancd n
a N shapc IhaI cyc!c Ihrouh lour phascs, Ihc !asI ol vhch s !Kc
Ihc nIa! onc buI dsp!accd onc cc!! daona!!y. On a compuIcr Ihc
paIIcrnsccmsIo!dcacrossIhcscrccnsoIsrclcrrcdIoasa "!dcr.'
G!dcrs andoIhcr spaccshps provdc Ihcc!carcsI cxamp!c ol cmcr
cncc n cc!!u!ar auIomaIa: vh!c Ihc auIomaIa Ihcmsc!vcs rcman
hxcd nIhcrcc!!sa cohcrcnI paIIcnolsIaIcsmovn aoss Ihcm s
c!car!yancvcnIIyIhaIscas!ydsInushab!clrom Ihcm.
SI!! !lcs, osc!!aIors, spaccshps, and oIhcr cmcrcnI paIIcns can
nIcracIandnIhcsc nIcracIonsIhcyrcvca!Ihcr capacIcs. As vIh
mo!ccu!ar popu!aIons Ihc nIcracIons arc smp!c co!!sons buI Ihc
rcsu!Is ol a co!!dn cvcnI arc so varcd IhaI Ihcy musI bc sIudcd
cmprca!!y.Two!dcrs, lorcxamp!c, can co!!dcvIh cach oIhcr n
73 dllccnIvays,IhcouIcomcdcpcndnonIhcrposIonsandIhc
phascolIhcrosc!!aIonaIIhcImcolIhccvcnI:28olIhcscco!!sons
rcsu!In IhcmuIua!annh!aIon ol boIh !dcrs: sx producc a b!ocK
asIhcr ouIcomc: Ihrcc rcsu!I n a b!inKcr: Ivo !cavc a sn!c !dcr
bchnd: and Ihc rcsI producc a varcIy ol sI!! !lcs and osc!!aIors.
Thcpossb!c ouIcomcs bccomc morc numcrous as vcncrcasc Ihc
numbcr ol !dcrs and vh!c cllorIs Io caIa!ouc Ihcm cxhausIvc!y
conInuc a morc promsn rouIc Io rcvca! Ihc capacIcs ol mu!I-
!dcrco!!sons s Io !ooK lor spcchccombnaIons IhaI synIhcszc
nIcrcsInobjccIs.Thcrcsa co!!sonol I 3 !dcrs, lor nsIancc, IhaI
synIhcszcsapaIIcrnmorccomp!cxIhanIhc!dcrIsc!l: ajlidrjun.
an osc!!aIn paIIcrn IhaI produccs a conInuous sIrcam ol !dcrs.
Thcrc sa!soan nIcrcsIncapacIydsp!aycdbysomc sI!! !lcs: Ihcy
arccapab!colco!!dnvIhanoIhcrpaIIcrndcsIroynIvh!caIIhc
samcImcrcconsIIuInIhcmsc!vcs. ThscapacIy "IocaI oIhcrpaI-
Icns a!so dcpcnds on Ihc capacIy ol Ihc paIIcns Iobc caIcn: a sI!!
!ilcshapcd !Kc 0hshhooK, lorcxamp!c, can caImosI paIIcrnsbuIs
noIcapab!coldcsInab!ocK.`
Knov!cdcolIhcsccmcrcnIIcndcnccsandcapacIcscanbcuscd
n Ihc consIruc|on ol "cnnccrcd Ilc paIIcns. !ocKs, b!nKcrs,
and !dcrs arc rclcrrcd Io as "naIura! paIIcns bccausc vhcn onc
LLLLULK U1M1 N LKNb LW
sIar|s a Ilcscsson vIh a random popu!aIon ol !vc anddcadcc!!s
Ihcsc Ihrcc paIIcrns |and oIhcrsmp!concs) cmcrc sponIancous!y.
Ths sponIancIy s cxp!ancd boIh by Ihc lacI IhaI Ihc paIIcns arc
sma!!-paIIcnsmadcouI ola lcv!vccc!!shavca hhcrprobab!Iy
ol occurrn Ihan !arc oncs-andbyIhc lac| IhaIIhcycan arsc lo!-
!ovn scvcra! scqucnccs ol prcdcccssorpaIIcrns: Ihc morc numcr
ous Ihc scqucnccsconvcrnon a paIIcrnIhchhcrIhc!Kc!hood
IhaI onc ol Ihcm W occurbychancc. NosI !arc andcomp!cx paI
Icrns, on IhcoIhcrhand,musIbc handcralIcd. A!dcrunsaood
cxamp!c ol an cnnccrcd paIIcrn. Thc hrsI un Io bc crcaIcd vas
bascd on a "shuII!c,a paIIcrnIhaImovcs!Kc a spaccshpbuIpcr
odca!!y chancs drcc|on. Thc shuII!c sa smp!cIranu!ar paIIcrn
IhaI may occur naIura!!y buI I s Iypca!!y shorI-!vcd bccausc as I
chancsdrccIon Ico!!dcsdcsIrucIvc!yaansIIsovndcbrs.uIl
b!ocKs acIn ascaIcrsarcposIoncdaIancxacIp!accIhcyW con-
sumc Ihc dcbrs sIab!zn Ihc shuII!c. Oncc sIab!zcd Ivo shuII!cs
can bc madc Io co!!dc vIh onc anoIhcr and l Ihcr posIon and
Imn s jusI rhIIhcy v!! proucc a !dcr cvcry 30 cncraIons.'
Ennccrcd paIIcrns arc mpor|anI bccausc Ihcy ho!dIhc KcyIoIhc
oIhcr IcsI ol cmcrcncc: Ihc ab!Iy ol cmcrcnI cnIIcs aIonc sca!c
Io bccomc componcnI par|s ol !arcr cnIIcs vIh Ihcr ovn cmcr-
cnI propcrIcs. G!dcr uns arc a!rcady an cxamp!c ol such !arcr
sca!c cmcrcncc buI lor Ihc IcsI Iobc snhcanI vc vou!d vanI Io
dcIcrmnc l usn Ihc paIIcrns IhaI cmcrc n a popu!aIon ol Ihc
smp!csIauIomaIavccanbu!danoIhcrauIomaIonvIhmuchhhcr
ccmruraricnalcaracin.
Thc spacc ol a!! possb!c auIomaIa W bc dcscrbcd n dcIa! n
ChapIcr I0 buI aI Ihs ponI I can bc characIcrzcd as sIruc|urcd
by snu!arIcs n compuIaIona! capacIy: a hnIc sIaIc auIomaIon
rcprcscnIs a capacIy mnmum vh!c a so-ca!!cd 1urinj machin
rcprcscnIsa capacIymaxmum. Thcman lacIor conIrbuIn IoIhs
rcaIcr capacIy s acccss Io mcmory, a dcrcc ol acccss IhaI varcs
lromzcronIhccascolhnIcsIaIcauIomaIaIoabso!IcnIhccascol
a Turn machnc and Is nnIc mcmory "Iapc. Thc ap bcIvccn
Ihc mnmum and maxmum ol compuIn capacIy can bc brdcd
bccauscIhcdllcrcnIauIomaIasIandnarc!aIonolparI-Io-vho!cIo
onc anoIhcr. A Tn machnc, lor cxamp!c, nccds a hnIc sIaIc
auIomaIon asa componcnI parI IoconIro! Ihc "hcad I uscsIorcad
Zb
Z
NLbN N bMUL1N
Ihc conIcnIsolIsinhniIcIapcandIo vrIconI.OnIhcoIhcrhand,
vhaI vc nccd lor our sccondIcsIol cmcrcncc snoIavayIo bui!d
a Turn machinc ouI olhnIc sIaIc auIomaIabuI amcans Io buId I
ouI ol cmcrcnI paIIcrns ol auIomaIon sIaIcs IKc !dcrs and !idcr
uns. ThcrcarcIvodllcrcnIvaysolapproachin IhschaI!cnc. Onc
s Io acIuaI!y bu!d a vorKn Turn machnc, vIh Is rcadlvrIc
hcad and Is mcmory Iapc, andmaKc I carry ouI a caIcu!aIon IhaI
couId noI bc pcrlormcd by a hniIc sIaIc auIomaIon. Thc oIhcr s Io
shovIhaIinrrincirlsuchaconsIrucIoncou!dbcpcrlormcdvIhouI
acIuaI!ycarryinIouI.
oIh approachcs rc!y on Ihc lacI IhaIcompuIin machincs can bc
bui!IouI ol basc diIa! circuIscaIIcd lcjicaljars. ADADdaIc, lor
cxamp!c, hasIvoinpuIsandoncouIpuI. l Ihcva!ucolboIhinpuIss
Ihcnumbcr " ! " IhcouIpuIv!!a!sobc a " ! ": anyoIhcrcombinaIon
yc!ds 0asan ouIpuI. ANoI aIcIaKcsasin!cinpuIandouIpuIs Is
opposIc. a 0lIhc npuI s " ! " and vcc vcrsa. Usn Ihcsc c!cmcn-
Iary crcuIs as componcnIs !arcr oncs can bc crcaIcd, such as a
IIp-BopcrcuIIhaIacIs asIhcsimpIcsIIypcolmcmory.ADyolIhcsc
c!cmcnIary circuIs canbc crcaIcd ouI ol IranssIors and mIhaI lorm
IhcycxsImIhc chpsIhaIpovcrmosIpcrsonaIcompuIcrs, buIIhcy
can aIso bc crcaIcd ouI ol cmcrcnI Ilc paIIcns. AD And aIc, lor
cxamp!c, may bc bui!I ouI ol a !dcr un and an caIcr. Thc !dcr
un produccs a conIinuous sIrcam ol !dcrs IhaI dsappcars oncc I
rcachcs Ihc caIcr. Thc inpuIs Io Ihc aIc arc Ivo addIonaI !dcr
sIrcams, produccd c!scvhcrc and aIrcady codcd vIh inlormaIon.
Ihcprcscnccola!dcrinIhcsIrcamrcprcscnIsa" ! " vhi!cIsabscncc
rcprcscnIs a 0. Thcsc Ivo inpuIsIrcamscoIIidc dcsIrucIvc!y vIh
IhcmansIrcamaI0dcrccsbclorcIrcachcsIhccaIcr.IlboIhinpuI
sIrcamshavca !dcr nIhc samcposIion |lboIh inpuIs havc avaIuc
ol" ! " ) IhchrsIinpuIW co!!dcandc!mnaIc a!dcr lromIhcman
sIrcamcrcaIinahoIc IhrouhvhchIhccorrcspondn!dcrin Ihc
sccond inpuI can ass. A!I oIhcr combinaIons do noI !cI any !dcr
Ihrouh. Thus, Ihc asscmb!y acIs !Kc an ADd aIc, Ihc inpuI !dcrs
IhaI manac o maKc iI Ihrouh Ihc main sIrcam consIIuIin Is
ouIpuI.Thc "inprincp!c" approachsIopshcrcbccauscvIhIhsAnd
aIc and a simp!cr Io bui!d NoI aIc d Ihc crcuIs nccdcd lor a luI!
compuIcr couIdbc bui!Ilvc had cnouh paIcncc.
LLLLULK U1M1 N 11LKNb LW
Thc sccond approach, on Ihc oIhcr hand, docs noI aim aI simpIy
provinIhaIIhc consIrucIons lcasb!c buIaIvorKnouIdIhcdl
hcuIIdcIai!s. AcurrcnIimp!cmcnIaIoncan scrvc asan iIIusIraIon. a
simp!c Turin machnc capabIc ol bcin in IhJcc dillcrcnI sIaIcs and
usinIhJcc symbo!sIovrIcinIsmcmoryIapc.Thismp!cmcnIaIion
is casicr Io dcscrbc sIarIin aI Ihc Iop vIh Ihc lu!! auIomaIon and
vornourvaydovnvards. Thc!arcsIsca!c componcnIsIhaImusI
bcbuIIarcahniIcsIaIcmachncconIro!InIhcrcad-vrIchcad,Ihc
mcmoryIapc,andasna!dcIccIorconnccIinIhcIvoIocIhcr.Each
olIhcIhrccmancomponcnIscanbc asscmb!cd ouIolsimp!croncs.
a hniIc sIaIcmachinc can bc imp!cmcnIcd vIh nincmcmorycc!!s:
a Iapc vIh Ivo sIacKs: and Ihc snaI dcIccIor can bc madc ouI ol
coIIidn!dcrsIrcamsandolunIsIhaIdup!caIccachsIrcam,scnd-
inIhccopcsIoIhcsIacKsandIhchnIcsIaIcmachinc.EacholIhcsc
sub-componcnIs can, n Iun, bc bui!I ouI ol cvcn simp!cr oncs.
A mcmory cc!!, lor cxamp!c, can bc madc ouI ol a !dcr un and a
spcca!IypcolshuII!c|caI!cda"quccnbcc")IhaIacIsasarcBccIorlor
!dcrs, and can bc acccsscd by Ivo sIrcams ol spaccshps cach pro-
duccdbyIsovn un. WhcnIhcsc andoIhcrcomponcnIsarcasscm-
bIcdIocIhcrIhcrcsu!Isavor!ODTurinmachincIhaIcanacIua!!y
carry ouI compuIaIions. Thc compuIaIons arc Ioo simp!c and Ihc
proccss is Ioo s!ov lor Ihc cmcrcnI auIomaIon Iobcol any pracIica!
usc, buI vaIchin Ihc Turin machinc in opcraIon nduccs in Ihc
vcvcr Ihc samc !ODdol avc crcaIcd byIhc cmcrcnIIhundcrsIorm
dscusscd in Ihc prcvous chapIcr. U Ihis casc, hovcvcr, vc arc noI
IcmpIcd Io vcv Ihc rcsu!I as "mracu!ous" bccausc Ihc mcchanisms
|usIdcscrbcdarc aIrcadyparIolIhc cxp!anaIon, ancxp!anaIonIhaI
may bc consdcrcd Io bc causaldcspIc Ihc lacI IhaI !dcr coIIisions
and oIhcr cvcnIs arc noI physca!. uI Ihs sIiII !cavcs par| ol Ihc
mysIcryuncxp!ancd Ihcmcchanism-indcpcndcnIparI.
WhaIpossbi!Iyspacc sinvo!vcdinIhisoIhcrparI ol Ihc cxp!ana-
Iion ivcn IhaI Ihc inIcracIons bcIvccn Ihc hnIc sIaIc auIomaIa
nhabIin Ihc ccI!s ol Ilc arc dchncdbyru!cs vhaImusI bc dcIcr-
mincd s Ihcsinu!arIcssIrucIurinrhsracc]allrcssiblruls. ADd
vhaI Ihcsc sinuIarIcsmusI accounI lor s Ihc cxsIcncc ol naIura!
paIIcrns!Kc b!ocKs and !dcrssncc a!!Ihc cninccrcdoncsuscd Io
bui!d Ihc cmcrcnITurinmachincdcpcndon Ihcm. SIrcI!yspcaKin
Z
Zd
NLbN N bMUL1N
vc nccd Ivo spaccs, Ihc spacc ol possb!c ru!cs and Ihc spacc ol
possb!cvaysolI!naspacc, andnoIjusIIvo-dmcnsona!spccsas
n IlcbuI spaccs ol any numbcr ol dmcnsons. Thc basc nshI,
hovcvcr, can bc i!!usIraIcd by sIudyn cc!!u!ar auIomaIa opcraIn
n a onc-dmcnsona!spacc. U Ihs casc Ihccc!!sbccomc ponIs n a
!nca!!ovnusIonorcIhc comcIry olIhcI!csandIoconccnIraIc
on Ihc ru!cs. U Ihcprcvous chapIcrvcdscusscd a possb!Iy spacc
|sIaIcspacc) IhaIcnIcrsnIoIhccxp!anaIonolIcndcnccs. Thcspacc
ol possb!cru!cs, on Ihc oIhcr hand, cnIcrs nIo Ihc cxp!anaIon ol
capacIcs bccausc Ihcru!cs spcclyIhc vays n vhch auIomaIa can
lcc| and bc allccIcdby onc anoIhcr. WhaIcmcrcslrom IhcacIa!
cxcrcsc ol Ihcsc capacIcs, onc !cvc! hhcr n Ihc par|-Io-vho!c
rc!aIon, s aan Icndcnccs: Ihc Icndcncy ol ccrIan Ilc objccIs Io
cmcrc sponIancous!y.
ThcszcolIhsspacc svcnbyIhcnumbcrolpossb!cru!cs. II can
bcca!cu!aIcd lromIhc numbcrol combnaIons ol Ivovarab!cs: Ihc
numbcrol sIaIcsnvhch a cc!!canbcandIhcnumbcrolnchbors
|nc!udin Isc!l) vIh vhch I nIcracIs. Thc numbcr ol sIaIcs rascd
Io Ihcnumbcrolnchbors yc!ds Ihc combnaIon ol possb!c sIaIcs
nvhch cachnchborhoodcanbc. Eachsuchpossb!cconhuraIon
olsIaIcs lor a nchborhood dcIcrmncs invhaIsIaIc a cc!!canbc aI
Ihc ncxImomcnI. Sncc Ihs s a!! a ru!c docs, dchnnIhcIransIon
lromoncsIaIcIoIhcncxI, IhcIoIa!numbcrolru!cs sIhcnumbcrol
sIaIcs rascd Io Ihc numbcr ol possb!c conhuraIon ol sIaIcs lor a
nchborhood. P onc-dmcnsona!cc!!u!arauIomaIon invhchcach
cc!! can bc n Ivo sIaIcs and inIcracIs on!y vIh Is Ivo mmcdaIc
nchbors has a IoIa!ol 256possb!c ru!cs. BuI in acc!!u!arauIoma
Ion!KcIlc,vIhIvosIaIcsandnncnchbors,Ihcnumbcrol ru!cs
s, rouh!y, Ihcnumbcr I0 lo!!ovcdby I 54zcros. Thscnormousszc
cxp!ansvhyIhchrsIscrousnvcsIaIonolIhcpossb!Iyspacclor
nIcracIon ru!cs vas pcrlormcd n Ihc onc-dmcnsona! casc. BuI
hovdocsoncscarch lorIhcsnu!arIcs nsuch a spacc?
WcsavIhaIIop!ayIhcamcolIlca lcvolIhccc!!sarcscIbyIhc
uscrIo Ihc !vsIaIc and Ihcn Ihc paIIcrn ol !vccc!!ssa!!ovcdIo
cvo!vc on Is ovn. Thc samc proccdurc can bc uscd hcrc: vc can
sc!ccIoncru!clromIhc256possbi!Ics,vcIasIarIinpaIIcrn, and
Ihcnlo!!ovIs cvo!uIon. U mosI cascs as Ihccvo!uIonprocccdsIhc
numbcr ol conhuraIons IhcsIaIcscan lormdccrcascs approachin
LLLLULK U1M1 N LKNb LW
n Ihc !on run a limirinj rarrm. Iour dllcrcnI !mIn paIIcrns,
assocaIcd vIh lourdllcrcnI c!asscs ol ru!cs, havcbccn dscovcrcd:
IhchrsIc!assolru!cs!cadsIohxcd, homocnouspaIIcrns:Ihcsccond
c!ass vcs rsc Io pcrodc paIIcns: IhcIhrdc!ass !cads Io rc!aIvc!y
random conhuraIons: and hna!!y, Ihc lourIh c!ass |or "c!ass I")
produccs Iruc comp!cxIy, inc!udn !oca!zcd sIrucIurcs capab!c ol
moIon, !icIhc!dcrsandoIhcr spaccships olIlc.'Thchucpopu-
!aIon ol ru!cs nhabIn Ihc possb!Iy spacc ol Ivo-dmcnsona!
cc!!u!ar auIomaIa cannoI, ol coursc, bc cxp!ord onc aI a Imc buI
I can bc IrcaIcd jusI !Kc anyoIhcr !arc popu!aIon: sIaIsIca!!y. U
oIh

rvords, Ihcpopu!aIon canbcsamp!cd n a sysIcmaIc vayand

Ihc !mIn paI|crns dscovcrcd by usn rcprcscnIaIvc ru!cs lrom
cach samp!cd rcon. Whcn Ihs sIaIsIca! ana!yss has bccn carrcd
ouIa!!lourc!asscshavcbccn rcdscovcrcd.
Thc lourc!asscs ol ru!cs arcIhcsinu!arIcs sIruc|urn Ihc poss-
b!Iyspacc, vIhc!assIinIcrac|onsrcprcscnIinamaxmumcapac-
Iy Io !cad Io comp!cx cmcrcnI paIIcrns vh!c c!ass rcprcscnI a
mnmum capacIy. BuI as n Ihc casc ol sIaIc spacc I s imporIanI
Io Knov noI on!y IhaI Ihc snu!arIcs cxisI buI a!so hov Ihcy arc
dsIrbuIcd. Ths s comp!caIcd in Ihc prcscnI casc bccausc un!Kc
sIaIcspaccIhaIsaccnrinucusrcrclcjicalsracvIhvc!!-dchncdspaIa!
connccIvIy Ihc spaccolpossb!cru!cs s a discrrccmbinarcrialsrac
posscssn no nIrnscspaIa! sIrucIurc: a!! Ihc possb!c ru!cs smp!y
!cncxIIocachoIhcr vIhouI lormnnchborhoodsoroIhcrspaIa!
arrancmcnIs. Wccan, on IhcoIhcrhand, imposc a par|cu!arordcr
on Ihc ru!cs Io rcvca! Ihc dsIrbuIon ol snu!arIcs. Onc sIraIcy
invo!vcsarrannIhcru!csnsuchavayIhaIIhoscbc!onnIoIhc
samc c!ass cnd up as ncihbors. P proccdurc IhaI yc!ds Ihs rcsu!I
sIarIsbydchnnIvo cxIrcmcs, IhcmosI homocnous andIhcmosI
hcIcrocncous ol ru!cs, andIhcn rcconsIrucIs Ihc lu!!spaccby sIarI-
naI Ihclu!!y homocnous cndands!ov!y ncrcasn Ihcdcrcc ol
hcIcrocncIy unI! Ihc oIhcr cnd s rcachcd." Thc sIrucIurc ol Ihc
rcsu!Inpossb!IyspaccsucsIsIhaIIhclourru!cc!asscsarcndccd
dsIrbuIcd n a parIcu!ar ordcr, vIh c!ass I sandvchcd bcIvccn
c!ass H and c!ass and occupyn a zonc much sma!!cr Ihan Ihc
zoncs on cIhcr sdc ol I.' This mcans IhaI c!ass I ru!cs arc noI
on!y snu!arby rcprcscnIn acapacIymaxmumbuI a!sobybcn
rc!aIvc!y rarc.
ZV
J
NLbN N bMUL1N
ccausc Ihc inIcracIion ruIcsor Iic bcIon Io cIass 1 Ihc cxis-
Icncc and disIribuIion o Ihcsc sinuIariIics ivcs us Ihc mcchanism-
indcpcndcnI componcnI o Ihc cxpIanaIion or Ihc Icndcncy o
naIuraIpaIIcnsIiKcIidcrs Io cmcrciniI. GIidcrs arc aprccondiIion
or Ihc cmcrcncc o a Turin machinc buI Ihc IaIIcr aIso nccds a
vhoIc varicIy o cninccrcdpaIIcrnsIo scrvc as iIscomponcnIparIs,
and Ihcsc oIhcr componcnIs do noI cmcrc sponIancousIy buI arc
Ihc rcsuII o counIIcsshours o sIIIcd human Iabor. In oIhcr vords,
human capaciIics arc aIso nccdcd or IhaI cmcrcncc. ConccivabIy
vc couIdrcpIacc humansI1IsbyanimpcrsonaIcvoIuIionaryproccss
Io discovcr and asscmbIc Ihc cninccrcd paIIcrnsbuI IhaIvouIdsIiII
noI |usIiy Ihc asscrIion IhaI cIass 1 ruIcs by IhcmscIvcs impIy Ihc
cmcrcncc o compIcx auIomaIa. This crror, causcd by a conusion
bcIvccn mcchanisms and mcchanism-indcpcndcnI sIrucIurc, is
compoundcdbyIhc acIIhaInoI onIycana Turin machinc bc buiII
ouI o Iidcrs, Iidcr uns, and oIhcr paIIcrns buI so can a univrsal
Turinmachinc. ThcTurinmachincsdiscusscdso ar arc spcciaIpur-
posc auIomaIa buiII Io carry ouI a spccihc compuIaIionaI IasK sIorcd
as a proram in Ihcir Iapcs. uI a univcrsaI Turin machinc is an
auIomaIonIhaIcan simuIaIc any spcciaIpurposc Turin machinc, iIs
mcmory Iapc incIudin amon oIhcr Ihins an cxccuIabIc symboIic
dcscripIion o Ihc auIomaIon iI is supposcd Io simuIaIc. Today's
pcrsonaI compuIcrs arc an approximaIion Io a univcrsaI Jurin
machinc, Ihc chcapcr and morc abundanI Ihcir mcmory Ihc cIoscr
Ihc approximaIion.
Thc rcason vhy Ihis ampIihcs Ihc phiIosophicaI crror o noIKccp-
inIhcIvo componcnso an cxpIanaIion scparaIc is IhaIcompuIcr
simuIaIionsIiKc IhaIo IhcIhundcrsIorm, asvcIIasaIIIhc oIhcrsycI
Iobcdcscribcd, arccarricdouIonIhcscapproximaIionsIoaunivcrsaI
Turinmachinc. U oIhcr vords, compuIaIionaIunivcrsaIiIy is a prc-
condiIion or Ihc simuIaIion o physicaI, chcmicaI, bioIoicaI, and
sociaIproccsscs. This can Icad Io Ihc misIaKcn idca IhaI Ihc maIcriaI
univcrsc iIscI is a ianI ccIIuIar auIomaIon usin cIass 1 ruIcs in
vhich aII acIua

proccsscs arc Ihc cquivaIcnI o simuIaIions. ' ' Thcrc

is indccd a connccIion bcIvccn ccIIuIar auIomaIa andphysicaI pro-
ccsscsbuIIhisIinKnccdsIobccxpIorcdmorccarcuIIy. IhysicisIsdid
noIIaKc ccIIuIarauIomaIa scriousIy unIiIiI vasshovnIhaIapopuIa-
Iion o hniIc sIaIc auIomaIa couIdmimic Ihc bchavior o imporIanI
LLLLULK U1M1 N LKNb LW
dicrcnIiaIcquaIions, cquaIionssimiIarIo Ihosc uscd u IhcIhundcr-
sIorm simuIaIion cxccpIIhaI Ihcy aIso incIudcIhc cccIs o viscosiIy
on Buid Bov. This dicrcnI Iypc o ccIIuIar auIomaIa arc rccrrcd Io
as lanic-jas auIomaIa. WhiIc iI is rcIaIivcIy uninormaIivc Io say
IhaI aunivcrsaITurinmachincbuiII viIhinaccIIuIarauIomaIoncan
simuIaIc physicaI proccsscs, Io say IhaI a IaIIicc-as auIomaIon can
dirccIIysimuIaIcpaIIcrnso Bov docs cncraIc ancvinsihI. iIadds
IhcisomorphicbchavioropaIIcnsoauIomaIonsIaIcsIoIhcaIrcady
kovnisomorphismbcIvccnIhcbchavior osoIuIionsIodicrcnIiaI
cquaIions and Ihc bchavior o physicaI proccsscs. And Ihis addcd
isomorphism nccdsIo bc cxpIaincd.
corcdcscribinIaIIicc-asauIomaIa vc nccd Io discuss in morc
dcIaiI Ihc physicaI proccsscs IhaI arc IhcIarcIo Ihc simuIaIion as
vcIIasIhcmodcIinsIraIcicsIhaIhavcprovcdsucccssuIinIhcpasI
andIhaImusIbcuscdin con|uncIionviIhIhcsimuIaIion.Wcarucd
u Ihc prcvious chapIcr IhaI scicnIihc modcIs IaIc advanIac o Ihc
dccomposabiIiIyorcaIiIymadcpossibIcbycmcrcnIpropcrIics. Onc
scicnIihc hcId, or cxampIc, canIaIc or ranIcd noI onIy Ihc cxis
IcnccoIcmpcraIurc andprcssurcbuIaIso IhcIcndcncyo radicnIs
o Ihcsc Ivo propcrIics Io dissipaIc, as Ion as iI ocs on Io cxpIain
Iarcr proccsscs in vhich Ihosc radicnIs arc componcnI parIs, such
asIhc inIcrnaImcchanism o asIcamcninc. AnoIhcrhcId can Ihcn
cxpIain hov Ihosc macro-propcrIics cmcrc rom Ihc inIcracIion
amon micro-componcnIs andIhcir propcrIics |moIccuIcs andIhcir
KincIic cncry and momcnIum) and vhy radicnIs posscss Ihc Icn-
dcncicsIhcydo.ThisoIhcrhcId, o coursc, IaKcsorranIcdIhcprop-
crIicsomoIccuIcsIhcmscIvcs, IrcaIinIhcmassoIidsphcrcscndovcd
viIh Ihc capaciIy IocoIIidc and rcdisIribuIc Ihcircncry, so anoIhcr
scicnIihc hcIdnccdsIo accounIor Ihc cmcrcncc oIhcscpropcrIics
u Icrms o Ihcir cvcn smaIIcr componcnIs |cIccIrons, proIons, ncu-
Irons) . Thcsc Ihrcc hcIds arc, rcspccIivcIy, cIassicaIIhcrmodynamics,
sIaIisIicaImcchanics, andquanIum mcchanics.
IaIIicc-asauIomaIa opcraIc aIIhc samcIcvcIo scaIc IhansIaIisIi-
caImcchanics andborrovssomcoiIsmcIhods. U parIicuIar, sIaIisIi-
caImcchanicsmusIdcaIviIhcnIircmoIccuIar popuIaIionsinordcrIo
accounI orproccsscsIiIcradicnI dcradaIion soiI cannoI dcaIviIh
moIcuIcsindividuaIIy. cvcnapcrccIIyriidsphcrchassixdcrccso
rccdom bcin capabIc o chanin iIs posiIion and momcnIum u
JI
JZ
NLbN N bMUL1N
Ihrcc dicrcnI dirccIions. U oIhcrvords,IhcsIaIcspaccora sinIc
moIccuIc is aIrcady six-dimcnsionaI, Ihc spacc o possibiIiIics or Ihc
vhoIc popuIaIion bcinIiIcraIIyunsurvcyabIc.Torcducc Ihcnumbcr
o dcrccs o rccdom modcIs in sIaIisIicaI mcchanics musI carcuIIy
choosc an inIcrmcdiaIc spaIiaI scaIc IhaI is Iarc rcIaIivc Io Ihc sizc
o Ihc moIccuIcs buI smaII rcIaIivc Io Ihc physicaI proccss bcin
cxpIaincd, andIhcn usc avcracs aI IhaI inIcrmcdiaIc scaIc as iIs cIc-
mcnIarycomponcnIs. 'ThissIraIcyorbridinIhcmicroandmacro
scaIcsis aIso uscd in IaIIiccas auIomaIa: choosin an inIcrmcdiaI
scaIcIhaIis Iarc cnouhrcIaIivc Io Ihc sizc o asinIc ccII buIsmaII
rcIaIivcIovhaIcmcrcsrom IhcinIcracIionsbcIvccn ccIIs. Thcusc
o inIcrmcdaIc avcracs as componcnI parIsimpIics IhaI in IaIIicc-
as auIomaIa Ihc imacs dspIaycd on Ihc compuIcr scrccn arc Ihc
rcsuIIoIhcavcracbchavioromany individuaIauIomaIanoIsinIc
auIomaIa sIaIcs as in Iic. P sccond dicrcncc has Io do viIh Ihc
arrovoIimc. U Iic IhcuIurc o a ivcn ccII iscnIircIydcIcrmncd
byIhc ruIcsbuI iIspasI is noI: a ivcn Iic paIIcrnmaybc arrivcd aI
by a varicIy o scqucnccs o prccursor paIIcrns. In IaIIiccas auIo-
maIa, on Ihc oIhcr hand, Ihc IvodirccIions oIimcmus| bc cquaIIy
dcIcrminisIicbccausc Ihc moIccuIar coIIisionsIhcysimuIaIc cxhibiI
Ihis indcrcncc Io a dirccIion o Iimc. IassicaI Ihcrmodynamcs noI
onIy IaKcs IcmpcraIurc or prcssurc or ranIcd iI aIso assumcs Ihc
cxisIcnccoan arrovoIimcpoinIinrom pasIIouIurc, IhaIIhis,iI
assumcs IhaI Ihc Icmpora! ordcr in vhich a scrics o cvcnIs occurs
in!Iucnccs iIs ouIcomc. BuI i vc vanI Io cxpIain Ihc cmcrcncc o
Iis dcpcndcncc on IcmporaI ordcr vc musI sIarI viIh componcnI
parIs in vhich Ihc arrov o Iimc docs noI ycI cxisI. U Ihc casc o
IaIIiccas auIomaIa Ihis impIicsIhaIIhc ruIcs dchnn Ihc inIcrac-
IionsbcIvccnccIIsmusIbcuIIyinvcrIibIc.''
IinaIIy, unKcIicinvhichaIidcrscapaciIyIomovciscmcrcnI
and in vhich Ihc ouIcomcs o coIIisions bcIvccn Iidcrs arc aIso
cmcrcnI, in IaIIiccas auIomaIaboIh Ihc moIion o moIccuIcs and
Ihc rcsuIIs o Ihcir coIIisions arc drccIIy spccibcd by ruIcs and arc
Ihcrcorc noI cmcrcnI.' On Ihc oIhcr hand, Ihc Iarcr scaIc phc-
nomcna IhaIrcsuIIromIhcsc coIIisions do cmcrc sponIancousIy.
II is rcIaIivcIysimpIc, or cxampIc, Io crcaIc a IaIIiccas auIomaIon
IhaI simuIaIcs Ihc dssipaIion o a radcnIandIhc rcachin o cqui-
Iibrium BuI vhaI happcns in siIuaIions in vhich Ihc radicnI is
LLLLULK U1M1 N LKNb LW
noIa!IovcdIocanccIiIscI, IhaIis, vhcniIccdsonanoIhcrradcnI
and cxpor|s iIs ovn dsordcr cIscvhcrc? U Buid dynamics Ihcsc
siIuaIions arc caIIcd "insIabiIiIics bccausc Ihc cxisIcncc o a Iivc
radicnI is unsIabIc |ivcn iIs IcndcncyIo dssipaIc) and bccausc Ihc
rcc cncry iIconIains canampIiysmaIIcaIurcIcss!IucIuaIionsinIo
compIcxpaIIcns. Thc convccIion ccIIsvc dscusscd in Ihc prcvious
chapIcr as componcnI parIs o IhundcrsIorms arc onc cxampIc o a
!IovpaIIcrn induccd by aninsIabiIiIy. AnoIhcrinsIabiIiIyamiIiar in
IaboraIorics |Ihc KcIvin-HcImoIIzinsIabiIiIy) is drivcn by a radicnI
o vcIociIy crcaIcd vhcn Ivo !Iovs o dcrcnI dcnsiIics rub aainsI
onc anoIhcr, such as air movinovcr a vaIcr suracc. Thc cmcrcnI
!IovpaIIcrns in Ihis casc arc vorIiccs. cIanoIhcrvcII-IUovnIabo-
raIory cxampIc |Ihc KayIcih-TayIor insIabiIiIy) is crcaIcd vhcn a
dcnsc, hcavy BuidisacccIcraIcdby a IihI onc. This dcnsiIy radcnI
produccs characIcrisIic bncrin or branchin paIIcrns as Ihc hcavy
!Iuid invadcs and dispIaccs Ihc IihI onc. AII o Ihcsc !Iov paIIcrns
havcbccnobscrvcdIocmcrcsponIancousIyinIaIIicc-asauIomaIa. '
AIIhouhvisuaIIysIriKinIhccmcrcnccoIhcscpaIIcnsisIcsso
a mysIcry Ihan IhaI vhich IaKcs pIacc insimuIaIions usin maIhc-
maIicaI cquaIions. Thc rcason isIhaIvhiIc cquaIions donoIincIudc
any inormaIion abouI Ihc microscopic dynamics bchind !Iov paI-
Icrns in IaIIiccas auIomaIaIhcisomorphism viIhmoIccuIar popu-
IaIions is parIiaIIybu!I in. NosIccIIuIar auIomaIa, or cxampIc, rcIy
on sIricIIyIocaIinIcracIions |onIyccIIs IhaI arc spaIiaIncihborscan
accIcachoIhcr) IhaIarcdcbncdbyIhccxacIsamcruIcs | aIIccIIsarc
IrcaIcd unormIy) . BuI Ihc propcrIics o IocaIiIy andunormiIy aIso
dchncmoIccuIarpopuIaIionsB vhichIhcrcisnoacIionaIa dsIancc
and in vhich cvcry mcmbcr is idcnIicaI Io cvcry oIhcr onc. OIhcr
propcrIics, suchasIhcinvcrIibiIiIyoIhcruIcs, arccxpIiciIIydcsincd
Io imiIaIc Ihc rcvcrsibiIiIy o moIccuIar inIcracIions. This impIics
IhaI aIaIccas auIomaIonsIarIcdviIharandomconhuraIion viII
rcmain cnIircIyrandommimicKinIhc acIhaIB IhcmaIcriaIvorId
noIhin inIcrcsIin happcns i vc sIarI a proccss viIh a maxima!Iy
dsordcrcdsIaIc, such as IhaIo a uIIydcradcd radicnI.'' To obIain
Ihc !IovpaIIcrnsjusIdcscribcd, or insIancc, vc musI sIarIIhcsimu-
IaIion in aconhuraIionIhaIconIainsaIivc radicnI.
Two oIhcr Kcy propcrIics o moIccuIar popuIaIions, conscrvaIion
andsymmcIry,aIsohavccorrcspondcnccsinciIhcrIhcruIcsorinIhc
JJ
J+
NLbN N bMUL1 N
comcIry ol Ihc cc!Is. II s vc!I ovn, lor cxamp!c. IhaI ccrIain
quanIIcs |such as Ihc quanIIy ol cncry) arc conscrvcd n physca!
proccsscs: cncry cannoI sponIancousIybccrcaIcdor dcsIroycd. Thc
ruIcs ol !aIIcc-as auIomaIa map cncry inIo Ihc numbcr ol sIaIcs
rcprcscnIn ndvdua! mo!ccuIcs and Ihs numbcr s a!vays con-
scrvcd. SymmcIry rclcrs Io Ihc ndllcrcncc IhaI Ihc rcuIarIcs n
Ihc bchavor ol moIccuIar popu!aIons dsp!ay Io bcin Iranslormcd
in ccrIainvays. KoIaInapopu!aIon, lorcxampIc, !cavcsIsbchav
ora!rcu!arIcsunchancd,oIhcrvsccxpcrmcnIsconducIcdnour
spnn1n pIancI vou!d vc rcsuIIs cnIrc!y dllcrcnI n IaboraIorcs
orcnIcd n dllcrcn drccIons. IaIIcc-as auIomaIausnccIIs viIh
asquarcshapcdonoIhavc cnouh roIaIona!symmcIry |Ihcyrcman
invaranI on!y lor roIaIons ol 0 dcrccs) inIroducin ar|lacIs nIo
a smu!aIon. HcxaonaI cc!!s, on Ihc oIhcr hand. havc cnouh roIa-
IonaInvaranccIo avodsucharIlacIs ccauscsomcsomorphsm
s a!rcady bu!I nIo !aIIcc-as auIomaIa Ihcr abiIiIy Io rcproducc
paIIcrns ol Bov s Icss probIcmaIc Ihan IhaI ol Ihc so!uIons Io Ihc
cquaIons ol Bud dynamcs. NcvcrIhc!css, Ihc cxpIanaon sI!! pro-
vdcs somc ph!osophca!inshIbccausc I dcnILcs somc Kcy prop-
crIcs |invcrIbiIiIy, conscrvaIon, andsymmcIry) IhaI arc sharcdby
Ihc smuIaIons and Ihc cquaIons and IhaI hc!p cxp!an vhy Ihc
so!uIonsIo Ihc!aIIcrcan cxhibIIhc rcu!arIcsIhaI Ihcy do.
U addIon, !aIIcc-as auIomaIacomp!cmcnImaIhcmaIcaImodc!s
in a novc! vay. U Ihc modc!s uscd in sIaIsIcaI mcchancs, lor
cxamp!c, ccrIan assumpIons musI bc madc abouI Ihc !iKcIihood ol
dllcrcnI spaIaI arrancmcnIs. U amodc!ol radcnI dsspaIon. lor
cxamp!c, vcmusIassumcIhaIIhcdsordcrcdsIaIcaIIhccndolIhc
proccss smuch morc probabIc Ihan Ihc ordcrcd onc aI Ihc sIarI, an
assumpIon IhaI musI D jusILcd on sparaIc rounds. uI vIh
IaIIcc-asauIomaIavc can acIua!!ylo!IovIhcmo!ccu!arpopuIaIon
unIiI I rcachcs a vcn spaIaI arrancmcnI. Ths s an mporIanI
capacIy ol smuIaIons noI sharcd by maIhcmaIca! cquaIons: Ihc
abiIiIyIo sIac a proccss andIracK I as I unlo!ds. Norcovcr, scvcra!
runs ol Ihc sam smuIaIon can bc carrcdouIchaninIhc va!ucs
assncd Io ccrIain paramcIcrs and IracKn Ihc unlo!dn proccss Io
chccKvhcIhcrIarrvcsaIIhcsamcLna!sIaIc. Sincc IhcvaIucsolIhc
paramcIcrs rcprcscnI assumpIons madc abouI Ihc cnvironmcnI on
vhch Ihc smuIaIcd proccss IaKcs p!acc, varyn Ihcm n dllcrcnI
LLLLULK U1M1 N LKNb LW
runs s cquvaIcnI Io chccKn vhcIhcr Ihc ouIcomc s robusI Io
chancs inIhc assumpIons. U oIhcr vords, cachrunolasimu!aIon
s !c an cxpcrmcnI conducIcd in a !aboraIory cxccpI IhaI I uscs
numbcrs and lormaI opcraIors as Is rav maIcra!s. Ior Ihcsc and
oIhcr rcasonscompuIcrsmu!aIons maybcIhouhI as occupynan
nIcrmcdaIc posIon bcIvccn IhaI ol lorma! Ihcory and !aboraIory
cxpcrmcnI.'"
IaIIcc-as auIomaIa provdc mporIanI!inKbcIvccnIhcsub|ccI
maIIcr ol Ihs chapIcr and IhaI ol Ihc prcvous onc. II s Imc, hov-
cvcr, Io movc bcyond popu!aIons ol dcnIca! cIcmcnIs vhosc dcn-
IIy snoI a!IcrcdbyIhcr nIcracIons. U oIhcrvords, vcmusI !cavc
Ihc rc!aIvc smp!cIy ol physcs bchind andmovc Io Ihc morc com-
p!cx vor!d ol chcmsIry, spccLcaIIy, Ihc chcmsIry ol Ihc prcboIc
soup. Thc mo!ccu!arpopu!aIons composn Ihc prmorda!mcdum
n vhch oransms and Ihcr cncIc codc cmcrcd dcparI n Ivo
vayslromIhc oncs cxamincdsolar. mosI olIhcmcmbcrsolapopu-
!aIon chanc dcnIIy n Ihc coursc ol Ihcr inIcracIons rcsu!Iin
in Ihc synIhcss ol cnIrc!y ncv compounds, vhIc Ihosc IhaI rcIan
Ihcr dcnIIy |caIaIysIs) arc capab!c ol allccIin IhaI synIhcss by
acccIcraIn I or dcccIcraIn I. ConIro!!n Ihc rcIaIvc rapdIy or
s!ovncssviIhvhch ncvcompounsarcproduccdsa vcrysinu!ar
capacIy, oncIhaIas vcv!!scc s as mporIanI lorIhccmcrcncc ol
Ivin crcaIurcs as s Ihc capacIylorsc!l-rcp!caIonIsc!l.
Jb
J
LHPJLJHLL
||IICId| 0O0|5||I O5 d0 |0O ||O0I O|IC bOJO
So lar vc havc cxamncd qucsIons ol cmcrcncc n Ihc rcIaIvcIy
simp!c casc ol a body ol vaIcr n vhch radcnIs ol propcrIcs !iKc
IcmpcraIurc, dcnsIy, orspccd causcIhcsponIancousoranzaIon ol
IhcmoIccuIcsnIoco!!ccIvcpaIIcrnsol!Iov.WccanconInucIousc
a body olvaIcr as our basc cnvronmcnI buI n ordcr Io add a ncv
Iaycrolcomp!cxIy vcmusImaKcIscomposuon !css homocnous.
Ths mcans IhaI ncv radicnIs musI bc nIroduccd: jradinrsc]ccn-
cnrraricn ol subsIanccs oIhcr Ihan vaIcr. Thcsc ncv radicnIs arc
aIso characIcrzcd bya Icndcncy Io dsspaIc buI unIiKc Ihc oId oncs
counIcrn IhaI Icndcncy nvo!vcs Ihc njccIon ol a !Iov ol maIIcr
noI jusI a Lov ol cncry. Thc cllccI ol IhaI njccIon vi!I. n Iun,
dcpcndonIhcspcchc chcmcaI subsIanccsbcninjccIcd. Somcsub-
sIanccs, lor nsIancc, havc a Icndcncy Io donaIc or cxporI somc ol
Ihcr proIons, basca!!yhydrocn aIoms dcprvcd ol Ihcr cIccIrons.
Thcsc subsIanccs arc caIIcd acds. OIhcr subsIanccs havc Ihc Icn-
dcncy Io acI as acccpIors or imporIcrs ol proIons and arc caIIcd
"bascs. Whcn conccnIraIons olsubsIanccs vIhIhcsc opposIc Icn-
dcnccss comc inIo conIacI, lormn an acd-basc or Ih radicnI, a
sponIancous !IovolproIons lrom oncchcmcaIspcccsIo anoIhcrs
cncraIcd asIhcmcnsuscdbyIhcradicnIIocancc!IscIl.AnoIhcr
cxampIc ol a ccmcaI radicnI s an oxdaIon-rcducIon or rcdox
radicnI crcaIcd vhcn a subsIancc IhaI has a Icndcncy Io oxdzc,
IhaI s, Io donaIc cIccIrons, comcs nIo conIacI vIh onc IhaI has a
Icndcncy Io rcducc or acccpI cIccIrons. A rcdox radicnI can drvc
anc!ccIronLovacross chcmcaIspccicsvhcnI occurs, lorcxamp!c,
K1LL LNLMb1KLb N 1NL KL 1L bU
n an aqucous cnvronmcnI n Ihc prcscncc ol conccnIraIons ol
mcIa!!c subsIanccs.
ChcmcaI radcnIs !iKcIhcscarcIhouhIIohavcp!aycdanmpor
IanI roIc n Ihc dynamcs ol Ihc prcboIc soup. ' uI n adduon Io
ncv radicnIs cxpIorn Ihc primordaI mcdum n vhch !vn crca-
Iurcs hrsIcmcrcdnvo!vcs consdcrn nIcracIons n vhch dillcr-
cnImo!ccuIcscanbondandlormancvcompoundor,onIhcconIrary,
invhch compounds canbcbroKcndovnnIoIhcr componcnIparIs.
Ths !cads noI omy Io muchIarcr combnaIoraI spaccs vIh aIoms
andmoIccu!cs oldillcrcnIspcOcscnIcrninIonnumcrab!cpossbIc
combnaIons, buI morc imporIanI!y, lorccs us Io nvcnI ncv mcans
Iocxp!orcpossib!IyspaccsIhaIarcinrrinsicallycrn-ndd. lchcmca!
nIcracIons !cadIoIhc lormaIonol a compoundnoI orna!!y prcs-
cnI n a mxIurc, and l Ihs compound s produccd n !arc cnouh
quanIucs Io yc!d a conccnIrauon radicnI, Ihcn Ihc vcrynaIurc ol
IhcmIurc asvcIIas IhaIolIs possbIIyspacc s chancd.U oIhcr
vords, Ihc inhcrcnI opcn-cndcdncss ol chcmcaI nIcracIons a!!ovs
possbi!IcsnoIornaIIyprcscnInaspaccIobcsubscqucnIIy addcd
IoI.Thc phIosophcaI nIcrcsIolIhcprc boIcsoupsIhaIbydchn-
Ion I docs noI conIan anyIvn crcaIurcs n I and ycI I has Ihc
capacIy Io!cadIoa novc! sIuaIon n vhch such crcaIurcs docxsI.
Thus,boIhmcchansmsandmcchansm-ndcpcndcnIsIrucIurcchanc
radca!!y oncc vc rcachIhc Icvc!ol chcmsIry and Is hcIcrocncous
moIccu!arpopuIaIons.
WhaIIypcs ol moIccuIcsinaddiIonIo vaIcr can vcassumcvcrc
parI ol Ihcsc prcbouc popuIaIons? Thc mosI promsn canddaIcs
arcmoIccuIcs ca!IcdrclymrsIhaIarcarrancd nIon Iincar chains.
U conIcmporary oransms boIh cncs and Ihc proIcns Ihcsc cncs
codclorarcpoIymcrsdillcrnomyinIhcrcomposIon:proIcnsusc
amino acds as componcnI parIs vh!c cncs usc nucoudcs. Today
mosIproIcnsarcsynIhcszcd lrominsIrucuonsconIancdincncsso
vclacca chcKcn-and-cdiIcmmavhcnIrvnIodccdcvhaIcamc
hrsI: cncs arc nccdcd Io crcaIc proIcnsbuIproIcins arcnccdcd Io
consIrucI andmanIanIhcbodics IhaIcarryIhosccncs.IroIcns,on
IhcoIhcr hand, canaIsobcsynIhcszcd vIhouIIhchc!polcncsand
Ihcr componcnI parIs havc a hhcr probabIIy Io lorm spnIanc
ous!yIhanIhoscolcncs.Sovcvi!IassumcIhaIpoIymcrsmadcouI
ol amno acds vcrc Ihc Ivpc ol moIccuIc orna!Iy popuIaIn Ihc
J
Jd
NLbN N bMUL1N
prcboIc soup.Thcscpo!ymcrs musIhavcbccnmuch!css comp!cx
Ihan Ihosc composn Ivin crcaIurcs Ioday buI Ihcy cou!d havc
ncrcascdn comp!cxIysmp!yby addn an cxIra amno acdaI onc
o! Ihcr cnds. Thc prob!cm vIh Ihs rouIc Io rcaIcr comp!cxiIy s
IhaI Ihc chcmica! rcacIon IhaI adds IhaI cxIra componcnI |ca!!cd
"po!ymcrzaIon") s noI !avorcd by an aqucous cnvironDcnI, Ihc
prcscncco!vaIcrpromoInnsIcadIhcbrcaKindovno!Ionchains
nIo shorIcr oncs. uI ! vc assumc IhaI Ihcsc primordia! po!ymcrs
manacd Io Iap nIo physca! or chcmca! radicnIs Ihcy cou!d havc
uscdIhaIcncryIoaccc!craIcpo!ymcrzaIonproducnIoncrchains
!asIcrIhanIhcraIc aI vhchIhcy vcrc broKcndovnbyvaIcr.
Achcmca! rcacIon may nvo!vc scvcra! sIcps, somc o! vhchIaKc
p!acc n IhcopposIcdirccIon o!radicnIdisspaIon and arc Ihcrc
!orccncrcIca!!yIcss!avorab!c.Addncncry!romancxIcrnaIra-
dicnIcanIrans!ormasIcp!rom"uphII"Io"do, "IhaIs, !roma
sIcp IhaI docs noIIaKcp!accsponIancousIyIo oncIhaIdocs, rcmov-
n I as an obsIac!c and accc!craIn Ihc ovcra!! rcacIon. Thc on!y
dravbacKIo Ihs so!uIonIoIhcpoIymcrzaIon prob!cm sIs !acK o!
spcchcIy: cxIcaI radicnIscan accc!craIcmanydi!!crcnIchcmcaI
rcacIions, somc o! vhch may nIcr!crc vIh po!ymcr rovIh. So a
bcIIcr so!uIon vou!d combnc Ihc cncry sIorcd n radicnIs vIh
somc !orm o! mclcular rccjniricn, Ihc smp!csI Kind o! vhch s
acomcIrcaIcomp!cmcnIarIybcIvccnIvodi!!crcnImo!ccuIcs, onc
acIin as a "!ocK" Ihc oIhcr as a "Kcy." IuIIn IocIhcr Ihcsc vo
capacIcs yc!ds a caralysr. NcIa!!c crysIa!s, !or cxamp!c, can ac| as
povcr!uIcaIaIysIs: Ihcy canaIIachIhcmscIvcsIo amo!ccu!c IvsIin
I or oIhcrvsc dc!ormin I pushn I avay !rom cquIbrium and
crcaIn a radicnI. Ths physca! radicnIssm!arIoIhoscIhaIcxsI
nmosI !oad-bcarin sIrucIurcs, suchasbuiIdns orbrdcs, invhch
Ihc sIruc|ura! c!cmcnIs carryin Ihc !oad arc Iypca!Iy dc!ormcd-
sIrcIchcd !Ihc Ioad s carrcd nIcnson, shrunK !in comprcsson-
and arc !or IhaI rcason Ioadcd vIh sIran cncry. A!Icr a mc|aIIic
caIa!ysIaIIachcsIo nddc!ormsIsIarcIasIcpinachcmcaIrcac|on
IhaI vascncrca!!yuphIIbc!orcIhcdc!ormaIons novdovnh!!
andvIIhavcabcIIcrchancc o!occurrn sponIancous!y.
WccanimancIhaIvIhIhcrhImIa!!iccaIaIysIsaddcd as nrc-
dicnIsIoIhcprcboIcsoup poIymcrzaIon cou!dhavcbccn cnhanccd
a!!ovin Ihc cxsIn po!ymcrs Io rov n comp!cxIy. uI an cvcn
K1LL LNLMb1KLb N 1NL KL1L bU
bcIIcr so!uIon vou!d bc H Ihc po!ymcrs Ihcmsc!vcs had caIa!yIc
capacIcs bccausc as Ihcy !aci!iIaIcd Ihcir ovn rovIh n !cnIh and
compIcxIy Ihcy cou!d a!so ncrcasc Ihcr caIa!yIc povcrs !urIhcr
cnhancin Ihcr comp!cxihaIon. U oIhcr vords, ! po!ymcrs couId
acI as Ihcr ovn caIa!ysIs Ihcy vou!d cncraIc a sc!!-sIimu!aIn
proccss IhaI vou!d ncrcasc Ihcr comp!cxiIy cxponcnIa!!y. Somc
conIcmporaryproIcns, ca!!cdnzms, can acI as morcpovcr!u!and
spcchc caIa!ysIs Ihan mcIa!!ic crysIa!s. Thcy can manipu!aIc Ihc
Bovs o! proIons and c!ccIrons drvcnbyIhand rcdox radicnIs buI
vIhouI rc!yin on nIcnsc conccnIraIons o! subsIanccs IhaI vouId
bc harm!u! Io an oransm. Somc o! Ihc amno acds IhaI composc
dcsIvc cnzymcs, !or nsIancc, can inIcracI vIh onc anoIhcr Io
acI as a proIon shuII!c, Irans!crrn posIvcIy harcd parIcIcs Io
and !rom a IarcI. Thcsc aminoacds may !orm Ihc acIvc sIc o! Ihc
cnzymc |IhcparIIhaI!ocKs nIoIsIarcI) soIhcycombncIhccapac-
Iy !or moIccuIar rcconiIon vIh IhaI !or radcnI manpuIaIon.
AndsmiIarIy!or rcdox radicnIs. ! a pccc o! amcIa! aIom, an ron
or coppcr on, !or cxampIc, s nscrIcd nIo an cnzymc I can n com-
bnaIon vIh Ihc am1no acds n Ihc acIvc siIc acI as an cIccIron
shuII!c. oIhproIon andc!ccIron !!ovs canbcuscd Io Iurnanuphi!I
sIcp n achcmica!rcacIioninIoa doonc acccIcraIn Ihcovcr-
a!!rcacIon viIhhhspccihciIy.'
Thcsc radicnI-manpu!aIon mcchansms brn us c!oscr Io a
soIuIion Io Ihc poIymcrzaIon probIcm buI Ihcrc s sI!!an obsIac!c.
boIho!IhcscmcchansmshavcbccndcnIhcdncnzymcsbc!onn
Io conIcmporary oransms buI Ihcsc arc Ihc producI o! m!!ons o!
ycarso! cvo!uIon, an cvo!uIon IhaIprcsupposcsIhccxisIcncco!Ihc
cncIic codc. Thc sponIancous!y asscmb!cd po!ymcrs n Ihc primor
diaI soup cou!dnoIhavcposscsscdIhcschnc!y Iuncd capaciIicssncc
Ihcy vcrc much shorIcr and Icss comp!cx Ihan Ihcr counIcrparIs
Ioday. Wcsccm Io !acc anoIhcr chcKcn-and-c dIcmma: po!ymcrs
nccd radicnIs Io ncrcasc n !cnIh and pasI a Ihrcsho!d o! !cnIh
Ihcy IhcmscIvcs an acqurc Ihc capacIy Iomanpu!aIc Ihosc radi-
cnIs, buI I s uncIcar hov Ihcy rcach Ihc crIica! !cnIh in Ihc hrsI
pIacc. Onc possb!c vay ouI o!IhcdiIcmma s cccrraricn. a po!ymcr
couId caIa!yzc Ihc !ormaIon o! anoIhcr po!ymcr vhch, in Iurn,
caIa!yscs Ihc!ormaIono!IhchrsI. OnccIhs c!oscd crcuiIs !ormcd
oIhcr poIymcrs can bc inscrIcd inIo I ! Ihcy caIa!yzc a rcacIon
JV
+
NLbN N b MUL1N
producnoncoIhccxsInmcmbcrsandIhcrovnproducIons
caIa!yzcd by anoIhcrmcmbcr. U Ihs vay an aurccaralyriclccrmay
sponIancous!yormandIhcnrov.Thcc!osurcosuchacoopcrauvc
ncIvorKocaIa!ysIs vou!d cndov IviIh sIab!Iy aansI d\srupIon
bypcrIurbauons anda!!ovIIoconccnIraIcIhcrcsourccs nccdcd or
rovIh |amno aOds, shorI po!ymcrs) nIo Isc!. To Ihc cxIcnI IhaI
Ihcsc rcsourccs arc uscd or Ihc bcnchI o a!! Ihc coopcraInmcm-
bcrs Ihc !oop as a vho!c bchavcs as a proIo-mcIabo!sm, Irappn
and d\csun "ood." 1 s Ihcsc cmcrcnI auIocaIa!yIc !oops IhaI
arcIhcrca!so!uuonIoIhcpo!ymcrzauonprob!cmandhcnccconsI-
IuIcIhcmancharacIcrs n IhcprcboIcdrama.
IcIssummarzcvhaIhasbccnsadsoar.Thcprob!cmoIhccmcr-
cncc o !vn crcaIurcs in an noranc vor!d has a vc!!-dchncd
causa!sIrucIurc. Thcmcchansmsinvo!vcdarca!!chcmca!nnaIurc:
Ihc crcaIon o bonds bcIvccn amno acds Io crcaIc proIcins: Ihc
accc!crauon o bond crcaIon Io compcnsaIc or Ihc sponIancous
dcsIrucIon promoIcd by an aucous cnvronmcnI, an accc!craIon
cxp!ancdbyIhcmanpu!auono radcnIs: Ihcuscoa!ocK-and-Kcy
mcchansm Io pcrorm mo!ccu!ar rcconuon and ocus Ihc cccIs
o radcnI manpu!aIon on spcOhc IarcIs: andhna!!y, Ihc orma-
Ion o coopcraIvc ncIvorKs o caIa!yuc po!ymcrs ac|n as proIo-
mcIabo!sms. A!Ihouh Ihcsc chcmca! mcchansms arc rc!aIvc!y
vc!!KnovnIhcrcarcsu!!apsnourundcrsIandn. Thcrcconuon
mcchansm,orcxamp!c,hason!ybccnparI!ycxp!aincd.Iorcnzymcs
Io havc Ihs capacIy Ihcr onc-d\mcnsona! sIrucIurc musI bc o!dcd
inIoaIhrcc-d\mcnsona!shapc vIhIhcrhIcomcIrccaIurcs. Thc
o!d\n prob!cm has noI ycIbccn so!vcd, Is sIudy consumin morc
supcrcompuIcr Imc Ihan any oIhcr prob!cm. U vhaI o!!ovs Ihc
comp!cxIcso Ihco!dn proccss v!!bcnorcd Io conccnIraIc on
Ihc ncrcasc n !cnIh o Ihc chans. U oIhcr vords, vc v!! assumc
IhaI!oncr chains havcmorccomp!cxcmcrcnIcaIa!yuc and rcco-
nuoncapacIcsIhanshorIcroncs. U add\Ion, asIhasbccn arucd
n prcvous chapIcrs, a causa! cxp!anaIon n Icrms o mcchansms
musIbc supp!ccnIcd viIh a mcchansm-indcpcndcnI componcnI:
an c!ucdaIon o Ihc sIrucIurc o Ihc spacc o possb!ucsassoOaIcd
viIh Ihc prcbouc soup Io dcIcrmnc vhcIhcr Is snu!arucs maKc
Ihc cxsIcncc o auIocaIa!yIc !oops noI jusI possb!c buI a!so hh!y
probab!c.
K1LL LNLMb1KLb N 1NL KL 1L bU
U acI, vc nccd Io cxaminc Ivo possbIy spaccs: Ihc spacc o
possb!cpo!ymcrsodcrcnI!cnIhsandcomposIonsandIhcspacc
o possb!cchcmca! rcacIons IhaI can producc Ihosc po!ymcrs. Thc
probabIy IhaI an auIocaIa!yuc !oop can sponIancous!y orm can
Ihcn bc asscsscd by a IhcorcIca! cxamnauon o Ihcsc Ivo spaccs
and Ihc acIua! cmcrcncc o such a !oop IracKcd usin compuIcr
smu!aIons. ThchrsIIhnIhaInccdsIobccva!uaIcdsIhcszcoIhc
Ivo possb!Iy spaccs. Thc numbcr o possb!c po!ymcrs o a vcn
!cnIhsIhcnumbcroava!ab!ccomponcnIsrascdIoIhcmamum
!cnIh. U Ihc proIcns uscd by conIcmporary oransms Ihc rcpcr-
Iorc o componcnIs s !mIcd Io on!y 20 amno acds. NcvcrIhc!css
cvcn a vcry shorI po!ymcr hvc amno aOds !on can cxsI in ovcr
Ihrcc m!!\on dcrcnI combnaIons |Ihc numbcr 20 rascd Io Ihc
hIh povcr). Thc numbcropossb!cpo!ymcrs300amnoaOds !on,
Ihc!cnIhoasma!!conIcmporary cnzymc, s!Icra!!ynhnIc. Thus,
vc arc consdcrn possb!Iy spaccs IhaI rov cxp!osvc!y as Ihc
!cnIh o Ihc po!ymcrs ncrcascs. NcxI vc nccd Io csImaIc Ihc
numbcr o possb!c chcmca! rcacIons IhaI can producc Ihs inhnIy
o combinaIons, IhaI s, Ihc numbcr o rcacIons IhaI can !uc |or
condcnsc) IvoshorIcr chains inIo a !oncr onc, and vcc vcrsa, cuI
|orc!cavc)a!onchannIoIvoshorIcroncs.Ioranyvcnpo!ymcr
Ihcrc arc many dcrcnI rcac|ons IhaI cou!d producc I: a sin!c
amno acd can bc aIIachcd aIonc cnd o an cxsun chain: or Ivo
shorIcrchainscanbccondcnscdIocIhcr: orIhrcccvcnshorIcroncs
canbcuscd, cachpcrhapsIhcproducIoIhcc!cavaco!oncroncs.
ThccxsIcnccomu!up!c vaysIoproducccachpo!ymcrmp!csIhaI
Ihcnumbcropossb!cchcmca!rcacIonsrovscvcnasIcrIhan Ihc
numbcr oproIcinspcccs asIhc!cnIho Ihc chans ncrcascs.'
Iina!!y vc nccd csumaIc Ihc dsIrbuuon o caIa!yIc capacucs,
a IasK IhaI s mpossb!c Io pcrorm viIhouI so!vn Ihc o!dn
prob!cm. A possb!c vay ouI o Ihs dhcu!Iy vou!d bc Io assumc
IhaI vcn Ihc inhnIy o boIh possb!c proIcn spcccs and chcmca!
rcacIons Ihc spcchc d\sIrbuIon o caIa!yIc capacucs docs noI
maIIcr much. Wc can assumc randomd\sIrbuIons, acKnov!cdn
ournoranccoIhcdcIa!s,andsI!!rcachsomcva!uab!cconUusons.
A!! IhaI vc nccd Io !Uovs Ihc !Kc!hood IhaI a scrcs o caIa!yzcd
rcacIonsnIcrconnccIcdIo o a oscd!oopcancmcrc sponIanc-
ous!y.ThscanbccsImaIcdbycrcaInaraphin vhchcachproIcn
+I
+Z
NLbN N b MUL1N
spccics bccomcs a poinI and cach chcmicaI rcacIion bcIvccn Ivo
proIcins bccomcs a cdc | oinin Ivo poinIs. Thc acI IhaI as IcnIh
incrcascs Ihc numbcr o possibIc rcacIions rovs asIcr Ihan IhaI o
proIcinspccicsmcansIhaIIhcnumbcrovaysoinIcrconnccIinIhc
poinIs rovs asIcr Ihan Ihc numbcr o poinIs. AI a ccrIain criIicaI
IcnIhIhcnumbcropossibIc connccIions rcIaIivc Io IhaIopoinIsis
soIarc IhaIIhc probabiIiIyIhaIsomc caIaIyIicproIcinsarcconnccIcd
inIoa cIoscdIoop bccomcsvcry hih.' ThisarumcnIissiDiIarIoIhc
onc uscd Io cxpIain Ihc Icndcncy o radicnIs Io dissipaIc. uI as in
IhaI casc, our conhdcncc inIhcconcIusionvouIdbcrcaIIysIrcnIh-
cncdi vc couIdacIuaIIyoIIovIhc proccss in a simuIaIion, Irackin
Ihc popuIaIion ol poIymcrs as Ihcy cI Ioncr andchcckini cIoscd
Ioops havc indccd lormcd. Thcsc simuIaIions arc coIIccIivcIy kovn
as "arIihciaI chcmisIrics. Somc arIihciaI chcmisIrics arc discrcIc
simuIaIions IhaI cxpIiciIIy rcprcscnI ca individuaI poIymcr and in
vhichIhcinIcracIionsbcIvccnchainsdcpcnd on Ihcir composiIion.
OIhcrs usc conIinuous dicrcnIiaI cquaIions inorin Ihc dcIai!s
o individuaI inIcracIions and ocusin insIcad on raIcs o chanc:
IhcraIcaIvhich cIcavac andcondcnsaIion rcacIions IakcpIacc, Ihc
raIcsaIvhichconccnIraIionsopoIymcrs o aivcnIcnIhorm, and
soon.
Thc conIinuous approach, caIIcd mradynamio. modcIs Ihc pri-
mordiaI body o vaIcr in vhich poIymcr cvoIuIion Iook pIacc as a
chcmicaIrcacIionvcsscI |or rcacIor) . ThcrcacIorisdrivcnbya ra-
dicnI o "ood," IhaI is, by an cxIcrnaI Bov omonomcrs and shorI
poIymcrs. AI any onc poinI u Iimc Ihc rcacIor conIains a Iarc buI
niIc numbcr ochcmicaI rcacIions, a numbcrIhaIcanbcrcduccdi
vc imposc aIhrcshoId o conccnIraIion bcIov vhichIhcprcscncc o
aparIicuIar spccicscanbcconsidcrcdinsinihcanI.Thus,aIanyivcn
insIanI vc can modcI Ihcdynamics oIhc rcacIorbyahniIc numbcr
o cquaIions, onc or cach chcmicaI rcacIion, Ihc cnIirc scI o cqua-
Iions posscssin a sinIc sIaIc spacc. Assumin IhaI Ihc chcmicaI
rcacIions arcrcvcrsibIc aIIovs usIourIhcrsimpIiyIhinsbccauscin
IhacascIhc possibiIiIy spacc or Ihc rcacIor is sIrucIurcd bya sinIc
sinuIariIy: a sIcady-sIaIc aIIracIor dchnin a sIabIc disIribuIion o
conccnIraIions o poIymcrs o dicrcnI spccics. On Ihc oIhcr hand,
unIikcIhcsIaIc spaccs o physics invhichIhc numbcr o dimcnsions
isivcninadvanccanddocsnoIchanc,inmcIadynamicsIhcnumbcr
K1LL LNLMb1KLb N 1NL KL 1L bU
o rcacIions docs chanc and so docs Ihc numbcr o dimcnsions o
Ihcspacc.KccpinIrackoIhcincrcascin Ihc numbcr o dimcnsions
can bc donc by coupIin a sIaIc spacc Io a raph Ic Ihc onc |usI
discusscd, a raph capIurin Ihc cxisIinpoIymcr spccics and Ihcir
chcmicaI inIcracIions. Thus, mcIadynamic simuIaIions cxpIorc nc
ccurldrcssibilinsracs.IhcraphivcsusIhcsIrucIurco Ihcspacco
possibIc chcmicaIrcacIionsvhiIc Ihc sinuIariIics o Ihc sIaIc spacc
providcIhcsIrucIurcoIhc spacc opossibIcouIcomcsoraivcnscI
o rcacIions.
AIypicaIruno amcIadynamicsimuIaIion procccdsIikcIhis: in|ccI
somc ood inIo Ihc rcacIorIocrcaIc a radicnI and cI Ihc dynamics
oin oIIov Ihc proccss unIiI iI rcachcs Ihc sIcady-sIaIc aIIracIor
nd chcck vhaI poIymcr spccics arc prcscnI abovc and bcIov Ihc
IhrcshoId o conccnIraIion: add Io Ihc raph any ncv spccicsIhaIis
abovcIhc IhrcshoId, crascanyoIdspccicsIhaIisbcIoviI, andupdaIc
Ihc connccIincdcs IoincIudcIhc ncvpossibIcchcmicaIrcacIions:
chanc Ihc scI o cquaIions |and iIs associaIcd sIaIc spacc) Io rcBccI
IhcchancincomposiIionoIhcrcacIor:oIIovIhcncvproccssunIi!
iI rcachcsancv sIcady sIaIc, ad|usIin IhcraphandIhccquaIionsIo
incIudc ncv poIymcr spccicsandrcacIions.This scricsosIcpsis Ihcn
rcpcaIcdmany Iimcs unIi!IhccomposiIionoIhcrcacIor sIopschan-
in. AI IhaI poinI anoIhcr sIcady-sIaIc aIIracIor is rcachcd |caIIcd a
"mcIadynamicaI hxcd poinI") buI Ihis Iimc iI is a sinuIariIy rcprc-
scnIinIhc Ion-IcrmIcndcncicso IhccnIircscricsorcacIions,and
iI is Ihcrcorc much morc imporIanIIhaI aII Ihc inIcrvcnin oncs.'
Oncc aI Ihc mcIadynamicaI xcdpoinIvcIookaI Ihc raphIoscci
iI conIains nodcs connccIcdby cdcsinIo a cIoscd ncIvor I such a
cIoscd ncIvorkcxisIsiI impIics IhaI, had Ihc simuIaIcdrcacIor bccn
popuIaIcdby individuaIpoIymcrs, IhcyvouIdhavc ormcd an auIo-
caIaIyIicIoop.IinaIIy, IhcsimuIaIionispcrormcdscvcraIIimcsvary-
in Ihc vaIucs o ccrIain conIroI paramcIcrs-Ihc dcrcc Io vhich
vaIcr accIsIhc rcacIions: IhcraIcaIvhich ood is cdinIoIhc soup:
Ihc disIribuIion o caIaIyIic capaciIics in Ihc poIymcr popuIaIion-Io
chcck IhaI Ihc cmcrcncc o Ihc Ioop occurs ovcr a vidc ranc o
vaIucs and iI is noI |usI an arIiacI. Thc rcsuIIs o Ihc mcIadynamic
simuIaIions IhaIhavc acIuaIIy bccnpcrormcd shov IhaIIhc sponIa-
ncous cmcrcncc o a proIo-mcIaboIism is indccd a IikcIy ouIcomc,
onc IhaI couIdhavcoccurrcdin prc bioIic condiIions.
+J
++
NLbN N b MUL1N
Jo lurIhcrrcduccIhc!Kc!hoodIhaIIhcrcsuIIsanarIlacIvccan
IryIo rcproducc IusnoIhcrsmu!aIonsbascdon cnIrc!ydllcrcnI
prncp!cs. Jobc Iru!ydllcrcnI Ihs oIhcr smu!aIon shou!d noI rc!y
on macro-propcrIcs !Kc ovcra!! raIcs ol conccnIraIon buI raIhcr
cnacImcronIcracIonsbcIvccn cxp!cI rcprcscnIaIons ol ndvd-
uaI poIymcrs and IcI Ihosc nIcracIons cncraIc Ihc conccnIraIons.
IKc mcIadynamcs Ihc ncv smu!aIon can cxp!oI Ihc smIarIy
bcIvccn!ncarchansolmoIccuIcs and!ncarsIrnsol symbo!s, buI
I musI noI cmpIoy Ihc sIrns as mcrc !abc!s lor an cnIrc spcccs
ol poIymcrs. InsIcad, I musI usc Ihcm Io capIurc Ihc bchavor ol
ndvdua! po!ymcrs. InparIcuIar, a symboI sIrn musI capIurc Ihc
doub!cIlc ol ancnzymc: on onchand, ancnzymcs smpIya chan
ol amno acds IhaI canbc acIcd upon Io bc cIcavcd nIo parIs or bc
condcnscd nIo !arcr vho!cs: onIhc oIhcr hand, alIcrIhs chan has
loIdcd nIoaIhrcc-dmcnsona! lorm, an cnzymc san cnIIyvIhIhc
capaIy Io acI on oIhcr chans brcaKn Ihcm up or !un Ihcm
IocIhcr.Jhc pcrlccI canddaIclorasymbo!sIrnvIhsuchadoub!c
!lc vou!dbc a smaI! compuIcr proram. Yhcn a compuIcr proram
s copcd lrom onc hard dsK Io anoIhcr, lor cxampIc, I s IrcaIcd as
amcrcchanoloncsandzcrocs, buIvhcnIscxccuIcdIbccomcsa
proccdurcIhaIcanpcrlorm avarcIyol opcraIons onoIhcrchansol
oncs and zcrocs. Jhus, compuIcrprorams !cad Ihc Knd ol doub!c
!lc IhaI vc nccd lorIhc ncvarIhca!chcmsIry.
NoIa!!prorams, hovcvcr, !cndIhcmsc!vcsIoIhsIasKbccauscvc
nccdIoconsdcrnoIon!yIhccapacIyolaproramIoallccIIhcdaIa
buI a!so Ihc capacIy ol Ihc daIa Io bc alccIcd byIhc proram: pro-
rams IhaI pcrlorm arIhmcIca! opcraIons, lor cxamp!c, dcmand
IhaI Ihc daIa conssIs ol numbcrs noI vords n En!sh, vh!c pro-
rams IhaI pcrlorm !nusIc opcraIons nccd vords noI numbcrs.
Sm!ar!y, n an arIhca! chcmsIry vhcn a proram s IrcaIcd as a
mcrcsIrnolsymbo!sImusIhavcIhcrhIcapacIcsIobcallccIcd:
Ihcsymbo!sIrnmusIbcab!cIoundcrorandomc!cavacsandcon-
dcnsaIonsandsIi!!rcIancnouhcohcrcnccIobccxccuIcdasa pro-
ram.IroramsvrIIcnnconvcnIona!!anuacsconIanmcchancaI
rccpcs spcchcd sIcp by sIcp-usn spccaI consIrucIons Io rcpcaI
scrcsolsIcpsandIoIranslcrconIroIlromoncrccpcIoanoIhcr-and
Ihcscrccpcsarccomp!cIc!ydcsIroycdlvcaddIoIhcm,ordc!cIclrom
K1LL LNLMb1KLb N 1NL KL 1L bU
Ihcm, random pcccs olcodc. uIa dllcrcnI lam!y olprorammin
!anuacs, ca!!cd "rccursvc luncIon Ianuacs, docs noI spccly
compuIcr prorams as sIcp-by-sIcp rccpcsbuIsmpIy as Iranslorma-
Ions ol npuIs nIo ouIpuIs, IhaI s, as luncIons, and morcovcr, as
luncIons IhaI can IaKc oIhcr luncIons as npuIs and producc ycI
oIhcr luncIonsas ouIpuIs. Jhs mcans IhaI a comp!cx proram can
bc bu!I usn a parI-Io-vhoIc rc!aIon: comp!cx luncIons can bc
crcaIcdouI ol simpIc luncIons vhcharccomposcdlcvcnsmp!cr
oncs, aIIIhcvaydovn Io opcraIonsIhaIarcdrccI!ycxccuIab!c. Jhc
rcsuIIn compIcx luncIons can bc cxprcsscd as symboI sIrns, vIh
parcnIhcscs scparaIn Ihc dllcrcnI ncsIcd componcnIs, sIrns IhaI
havc a much hhcr chancc ol rcmann cohcrcnI alIcr bcn acIcd
uponbyoIhcr luncIons."
Usnarccursvc luncIon!anuacaloIdcdpoIymcrvIhcaIaIyIc
capacIcscanbcrcprcscnIcdbyasIrnol symbo!s vIhIhcab!IyIo
allccI oIhcr sIrns, vh!c Ihc unloIdcd poIymcr IhaI s Ihc IarcI ol
Ihc caIa!ysI canbcrcprcscnIcdbyasymboIsIrnvIhIhc capacIyIo
bc allccIcd by oIhcr sIrns. Yhcn Ihcsc Ivo sIrns nIcracI a Ihrd
sIrnrcsu!IsrcprcscnIn Ihc producI ol Ihc chcmca! rcacIon caIa-
!yzcdbyIhc cnzymc. Josimp!ly Ihc smu!aIon spaIaIrc!aIonsarc
noImodcIcd cxp!cI!y, IhaIs, Ihcsmu!aIcdpoIymcrsarcnoIrc!aIcd
Ioonc anoIhcrbyrc!aIonsolproxmIy. Jhsscquva!cnI Ioassum-
nIhaIIhcchcmca! rcacIorssubjccIcdIoa consIanI sIrrnmoIon
IhaIdcsIroys a!! nchborhood rc!aIonsbcIvccn po!ymcrs. IIavcI!-
sIrrcd rcacIor a!! co!!sons bcIvccn moIccu!cs arc basca!!y random
sonIcracIonsbcIvccnpo!ymcrscanbc modc!cd byrandom!ypcK-
n Ivo symboI sIrns lrom Ihc cxsIn popu!aIon, IrcaIn onc
as a proram |cnzymc) and Ihc oIhcr as daIa |IarcI subsIraIc) , Ihcn
rc!casn Ihcr producI bacK nIo Ihc popu!aIon. Jhcsc chancc
cncounIcrs arc n!!ucnccd byIhc chann conccnIraIons ol symbo!
sIrns ol Ihc samc "spcccs, jusI as n a vcI!-sIrrcd rcacIor Ihc lrc-
qucncyolco!!sonbcIvccnIvopo!ymcrsoldllcrcnI spcccsvou!d
dcpcnd onhov manymcmbcrsolcachspcccs cxsI nIhcchcmca!
soup. ccausc rccursvc luncIon !anuacs havc Ihc compuIaIona!
capacIy ol Ihc mosI sophsIcaIcd auIomaIa, and bccausc ol Ihc ran-
dom characIcr ol Ihc co!!sons, Ihs arIhcaI chcmsIry s rclcrrcd Io
as a 1urinjj

. '

+b
+
NLbN N bMUL1N
AIypca!Turnassmu!aIonsIarIsvIhapopuIaIonolrandom!y
consIrucIcd luncIons. Thc popu!aIon, conssIn ol abouI I . 000
mcmbcrs, sIhcn a!IovcdIonIcracIproducnncvluncIons. Somc
ol Ihcsc producIs arc dcnIcaI Io oIhcrs aIrcady prcscnI, oIhcrs arc
novcI n Ihc scnsc ol noIhavn counIcrpar|s n Ihc cxsIn popuIa
Ion. vhIc ycI oIhcrs arc abso!uIc!y novc! n IhaI Ihcy havc ncvcr
cxsIcdnIhc hsIory rcprcscnIcdbyaparIcu!arrunolIhcsmu!a-
Ion.Thcpopu!aIonsdc!bcraIc!yKcpIconsIanI,byrandom!ydrain-
n cxccss symbo!ssIrns lrom Ihc as, and Ihs acIs as a sc!ccIon
prcssurc: Ihc onIy vay lor a smu!aIcd po!ymcr Io rcman in Ihc
popuIaIonlorcxIcndcd pcrods olImcsbybccomnIhcproducIol
a sIab!c scrcs ol inIcracIons bcIvccn luncIons. And, in Iun, Ihc
onIyvay lora scrcsol nIcracIonsIobc sIab!c sIodspIaycaIa!yIc
c!osurc. Thc smp!csI auIocaIa!yIc !oop s composcd ol a luncIon
IhaI acIs on a symbo! sIrn Io producc anoIhcr symbo! sIrin IhaI,
vhcnIrcaIcdasaluncIon, produccsIhcsymbo!sIrncorrcspondn
IoIhc hrsIluncIon. Such c!oscd scIsol nIcracInluncIonshavcin
lacI bccn produccd in Turin as smu!aIons and I can bc shovn
IhaI Ihcy corrcspond Io snu!arIcs |invaranI hxcd ponIs) ol Ihc
dynamcsol Ihc sysIcm.
Thc rcsu!Is lrom Turin ascs and mcIadynamcs shov IhaI auIo-
caIa!yIc !oops acIn as proIo-mcIabo!sms arc a rccurrcnI lcaIurc
n arIhca! chcmsIrcs usn vcry dllcrcnI lorma! rcsourccs. Ths
somorphsm, nIun,maybccvdcnccolycIoIhcrsomorphsm,onc
bcIvccn smu!aIcdpoIymcr popu!aIons andIhc rca!po!ymcrpopu
!aIons nhabnprcscnIday chcmca! rcacIors or anccnIprc boIc
soups. ThaI s, Ihc spaccs ol possb!Ics ol Ihc smu!aIons and Ihc
rca! cnvronmcnIs may cxhbI sulhccnI ovcr!ap so IhaI vhaI vc
!ca lrom Ihc lormcr can bc uscd as Ihc mcchansmndcpcndcnI
parIolIhccxp!anaIonolIhc!aIIcr.OnIhcoIhcrhand,IhcsponIanc
ous cmcrcncc ol auIocaIa!yIc !oops cannoI lu!Iy accounI lor Ihc
orinol!lcbccaus Ihcsc!oopsdonoIhavcIhcmcansIosIorcnlor-
maIon andIo pass vhaIcvcr advanIacsIhcymay havc acqurcd n
caIa!yIc clhcccy and spcchcIy Io luIurc cncraIons. A possb!c
so!uIon Io Ihs prob!cm rc!cs on Ihc lac| IhaI Ihc rcsu!Is ol Ihcsc
smuIaIonsa!soapp!yIopo!ymcrsmadcouIolnuc!coIdcs,po!ymcrs
IhaIdohavcIhcab!IyIo ac|asIcmpIaIcsnordcr Ioproducccopcs
ol Ihcmsc!vcs. Norcovcr vh!c somc chains ol nuc!coIdcs, such as
K1LL LNLMb1KLb N 1NL KL 1L bU
IhosccharacIcrznconIcmporaryNA, donoIhavccaIaIyIccapac-
Ics,IhcchanscomposnsomclormsolKAdohavcIhcm,andIhs
mcansIhaI anccnIKAmayhavc combncd Ihc ab!Iy IoIransmI
inlormaIon acrosscncraIonsvIh Ihcpropcr|yol caIaIyIccIosurc.
ThcloIIovinchapIcrv!IcxpIorcIhssccnaroandIhcncvcapacIy
IhaI cmcrcs vhcn a popuIaIon olscIl-rcp!caIinmoIccu!cs scou
p!cd Io any proccssIhaI bascsIhaI rcp!caIon onc vay or anoIhcr:
IhccapacIyIopcrlorm a scarchin a spacc ol possb!Ics.
+
+d
LHPJL
OOIC |O|I005 d0 0O ||O0| OIC bOJO
Achocc ola smp!c buI snhcanI sIarInponIlor an cxamnaIo
ol cmcrcncc has provcd mporIanI n Ihc casc ol boIhphyscs and
chcmsIry. A sm!ar sIraIcy v!I bc usclu! vhcn vc movc Io Ihc
rca!m olbo!oy. Wc musI rcmann an aqucouscnvronmcnIpopu-
!aIcdbymo!ccu!csbuIaddancvcapacIy.Ihcab!IyIosc!lrcpIcaIc.
Thc smp!csI sc!l-rcp!caIncnIIcsarc KNAmo!ccu!cscxsInn a
lrccsIaIc, IhaIs, noIcncapsu!aIcdbyamcmbrancorsubordnaIcdIo
Ihcnccdsolanoransm.Tocxp!anIhcrmcchansmolsc!l-rcpIcaIon
vc do noI havc Io consdcr Ihc comp!cx proccss uscd by Ivn
crcaIurcs, a proccss n vhch Ihc cncIc codc and a!I Is Irans!aIon
machncry arc cruca!!y nvo!vcd. A!!IhaI vc nccd Io accounI lor s
Ihc rc!aIvc!y smp!c proccss Ihrouh vhch an KNA mo!ccu!c can
scrvcasaIcmp!aIc lorIhccrcaIonolacopyolIsc!l. Apopu!aIonol
IhcscrcpIcaIors, B Iun, nIroduccs a ncvkndolradcnI, a rad-
cnI olhIncss. IIncssrclcrsIoIhc dllcrcnIa!rcproducIvcsucccssol
cmbodcd oransms buI I can a!so bc appIcd Iomo!ccu!ar rcp!ca-
Iors and IhcrdllcrcnIcapacIcs Io producc copcs ol Ihcmsc!vcs. 1
can bc rcardcd as a radcnI bccausc hIncssdllcrcnccs acI jusI Ikc
IcmpcraIurc dllcrcnccs. as !on as Ihc dllcrcnccs cxsI Ihcy luc! a
proccss ol sc!ccIon avorn Ihc rcp!caIon ol onc knd ol po!ymcr
ovcr anoIhcr: Ic momcnI Ihc dllcrcnccs dsappcar Ihc sc!ccIon
proccss sIops.'
IocusnonnakcdKNAmo!ccu!cshasIhcrclorc IhcadvanIacol
addnon!yamnma! !aycrolcomp!cxIyIoIhcvor!dvc arccxp!or-
n buI a !aycr IhaI a!rcady hasmany ol IhcpropcrIcsIhaI scparaIc
LNL1lL LK1NMb N 1NL KL1L bU
bo!oy lrom chcmsy. An KNAmo!ccu!c ssmp!y a scqucncc ol
lour possb!c nuc!coIdcs, cach mcmbcr ol Ihc scI dsp!ayn c!car
IcndcnccsIobondvIhjusIanoIhcrmcmbcr.U vc rclcrIo Ihc lour
nuc!coIdcs usn Ihc convcnIona! !cIIcrs |A, C, G, U) vc Knov
IhaIs IcndIobondvIhUs, andGsvIhCs.ThcscIcndcnccs, ol
coursc, nccd Io bc cxp!ancd by ycI oIhcr mcchansms buI lor Ihc
purposc olaccounInlorsc!l-rcpIcaIonIhcycansmp!ybcassumcd
Io cxsI. Gvcn Ihcsc Icndcnccs a scqucncc ol nuc!coIdcscan crcaIc
a copy ol Isc!l usn a mcchansm sm!ar Io IhaI ol convcnIona!
phoIoraphy. hrsI a ncaIvc s crcaIcd as comp!cmcnIary mono-
mcrsaIIachIhcmsc!vcsoncby oncIoIhcIcmp!aIc, andIhcna pos-
Ivc prnI s produccd lrom IhaI ncaIvc. Wc a!so Knov IhaI Ihc
amounI ol cncry nvo!vcd B lormn Ihc comp!cmcnIary chcmca!
bonds snoI IhaI !arc, rouh!y cqua!IoIhaI ol Ihcrma! !ucIuaIons
n an aqucous cnvronmcnI, andIhsmpIcsIhaI crrors nIhc copy-
n proccss v!I Icnd Io occur sponIancous!y. U oIhcr vords, Ihc
copyn crrors or muraricns IhaI provdc Ihc ncccssary varab!Iy Io
prcvcnI hIncss radcnIs lrom dsappcarn do noI nccd a scparaIc
cxp!anaIon.
scusson ol Ihc mcchansm-ndcpcndcnI sIrucIurc s a!so smpI-
hcd by sIarIn vIh nakcd KNA bccausc Ihc possbIIy spaccs
assocaIcd vIh !vn oransms can bccomc vcry numcrous as Ihc
comp!cxIyol Ihcr parI-Io-vho!c oranzaIon ncrcascs. U Ihc casc
ol uncc!Iu!ar oransms !kc bacIcra, lor cxamp!c, vc musI consdcr
aI !casIIhrccdllcrcnIspaccs. Ihc spaccolpossb!c cncs:Ihcspaccol
possb!c sIrucIura! proIcns and cnzymcs IhaI Ihcsc cncs codc lor:
and Ihc spacc ol possb!c spaIa! sIrucIurcs and mcIabo!c paIhvays
IhaI Ihc sIrucIura! proIcns and cnzymcs can lorm. U oIhcr vords,
vc nccd onc possbIIy spacc lor Ihc hcrcdIary nlormaIon |Ihc
cnoIypc) and Ivo possb!Iy spaccs lor Ihc bodIyIraIsproduccd
lromIhaI nlormaIon |Ihc phcnoIypc) . nmu!Icc!!u!aroransms
scvcra! oIhcr possbIIy spaccs musI bc addcd sncc Ihc phcnoIypc
a!so nc!udcs Ihc spacc ol possb!c cc!! Iypcs |such as musc!c, bonc,
ncrvc, b!ood) : Ihc spacc ol possb!cIssucsandoransIhcsccc!Iscan
lorm: and Ihc spacc oI possb!c oransms Ihcsc Issucs and orans
can composc. Norcovcr, n !arc anma!s Ihc mappn ol cnoIypc
nIophcnoIypc nvo!vcsacomp!cxcmbryo!oca!proccssIhaIson!y
mpcrlccI!y undcrsIood. By conIrasI, nakcd KNA has a vcry smp!c
+V
b
NLbN N bMUL1N
rcIaIon bcIvccn cnoIypc and phcnoIypc. Ihc unloIdcd po!ymcr
consIIuIcsIhc lormcrvhi!cIhc lo!dcdvcrsonandIs caIa!yIccapac-
Ics consIIuIcs Ihc !aIIcr.Thsmcans IhaIvc nccd Io consdcronIy
Ivo possb!Iy spaccs. Ihc spacc ol possb!c scqucnccs ol Ihc lour
nuc!coIdcs, Ihcszcolvhchs Ihc numbcrlourrascdIoIhc !cnIh
ol Ihc po!ymcr, and Ihc spacc ol lo!dcd lorms, a sma!!cr spacc vcn
IhaIscvcra!dllcrcnIscqucnccsol nuc!coIdcs cancndupvIhcquv-
a!cnIlo!dcdlorms.'
Kc oIhcr dscrcIc combnaIora! spaccs Ihc spacc olpossb!c KA
po!ymcrs s nIrnsca!!y amorphous. uI vc can imposc an ordcr on
Ias Ion asIhs s jusIhcd by rclcrcncc IoIhc mcchansm ol rcpIca-
Ion and as !on as I scvcs Io rcvca! Ihc sIrucIurc ol Ihc possbi!Iy
spacc. Thus, !ovnIhaIIhc varaIonna popu!aIon olmo!ccu!ar
rcpIcaIors nvo!vcs copyn crrors aI spcchc ponIs n a scqucncc
vc can arran Ihcm so IhaI cach mo!ccuIc has as nchbors oIhcr
mo!ccu!csdllcrnlromIbyonIyoncmuIaIon.Thsspaccsmu!I-
dmcnsona!bccausc ImusInc!udc a!!Ihc varanIs IhaI canbc cr-
aIcdbyvarynoncmonomcra!onIhcluII!cnIholavcnpo!ymcr,
and cach ol Ihcsc onc-muIanI varanIs musI bc assncd a dllcrcnI
dmcnson ol Ihc spacc. uI vh!c Ihc shccr numbcr ol dmcnsons
maKcs Ihc spacc vcry comp!cx I a!so rcaI!y simp!hcs Ihc dsIrbu
Ionolpossb!c mo!ccu!cs: cvcrypossb!cpo!ymcrs ndrccIconIacI
viIh aI! Is onc-muIanI nchbors and a scrcs ol nchbors lorms
a connccIcd paIh lor cvo!uIon Iolo!!ov. U oIhcrvords, vcn Ihs
spaIa! arrancmcnI mo!ccu!ar cvo!uIon can bc vsua!zcd as a ccn
rinucuswalklromonc nchborIoIhc ncxI drvcnby cvcnIsproduc
n oncmuIaIon aI a Iimc.Tocomp!cIc Ihc characIcrzaIon ol Ihc
possb!IyspaccvcmusIassnahIncssva!ucIo cacholIhc possb!c
po!ymcrs.IIncssrclcrsnIhscascIoIhc conscqucnccs lorrcproduc-
Ivc succcss ol Ihc caIa!yIc capacIcs IhaIcachpo!ymcrvou!d havc
l I vcrc lo!dcd nIo a Ihrcc-dmcnsona!shapc. Wcshovcd n Ihc
prcvous chapIr hov caIa!yIc capacIcs cxcrcscd n coopcraIvc
ncIvorKs can !cad Io Ihc locusn ol rcsourccson a parIcuIar scI ol
mo!ccu!csso vcan novIaKc Ihs lor ranIcdIo conccnIraIconIhc
cllccIsolcaIa!yssonraIcs ol rcpIcaIon.
A possb!Iy spacc ol sc!l-rcpIcaIn cnIIcs Io vhch hIncss
va!ucshavcbccn assncds caIIcda]rnsslandscar.Thc sIrucIurc ol
a vcn !andscapc dcpcnds boIh on Ihc dsIrbuIon ol sinu!arIcs,
LNL1L LK1NMb N 1NL KL1L bU
scqucnccs ol nuc!coIdcs vIhopIimum hIncss, and on hovhIncss
varcsnIhcnchborhoodolIhoscsnu!arIcs. Iorcxamp!c,lonc-
muIanInchborshavcsim!arhIncssIhcnIhcsnu!arIcswlorm
adsIrbuIonol!ocaIopIimarcscmbIinIhc "pcaKs olcnI!ys!opn
h!Is. IlhIncssvarcsrcaI!y bcIvccn nchbors, onIhc oIhcrhand,
Ihcn Ihc !andscapc vi!I bc morc rucd vIh Ioca! opIima pcaKs
surroundcdbysIccp cIlls. IIncss!andscapcsarcmporIanIbccausc
Ihcy comp!cmcnI Ihc IradIona! modc! ol cvo!uIon. U IhaI modc!
scarcc cnvronmcnIa! rcsourccs causc somc rcp!aIors Io bc sc!ccIcd
aIIhccxpcnscoloIhcrs, asc!cc|onproccsscxp!ancdbyIhccapacIy
ol Ihc cnvironmcnI Io allccI Ihc rcp!caIors. uI vc a!so nccd Io
consdcr Ihc capacIyol Ihc rcpIcaIors Iobc allccIcdand Ihs s vhaI
a hIncss !andscapc capIurcs. Norc spcchca!!y, n a !andscapc vIh
many !oca! opIima scparaIcd by !on dsIanccs, dsIancc bcn mca-
surcdnIhscascbyIhcnumbcrolmuIaIonsnccdcdIoconvcrIonc
scqucncc nIo anoIhcr, sc!cc|on prcssurcs a!onc cannoI accounI lor
parIcu!ar ouIcomcs. A cvo!vn popu!aIon may, lor cxamp!c, bc
Irappcd n a !oca! opIimum l Ihc paIh Io a snu!arIy vIh rcaIcr
hIncsspasscsIhrouhponIs olmuch !csscr hIncss. InIhs cascsc!cc-
Ion prcssurcs a!onc cannoI ds!odc Ihc popu!aIon lrom Ihc Irap.
Thus, Io Ihc cxIcnI IhaI mo!ccu!ar cvo!uIon can bc modc!cd as a
va!Ka!onaconInuous paIhol onc-muIanInchborsIhc "Iopora-
phy olIhchIncss!andscapcmusIbc parIol Ihcmodc!snccI laci
IaIcs or obsIrucIsIhoscva!Ks.
A prccsc characIcrzaIon ol Ihs Ioporaphy vou!d imp!y Knov!
cdcolIhccxacIhIncssva!uc olcachlo!dcdpo!ymcr. uI cvcnvIh
ouIIhs quanIIaIvcnlormaIonvc cancIa scnsc olIhcqua!IaIvc
charac|crsIcs ol a!andscapc. vhcIhcr Ihas asin!c oba!opImum
or many !oca! opIima, lor cxamp!c, or vhcIhcr Ihc nchborhood
ol Ihosc snu!arIcs s smooIh or rucd. A imporIanIlacIordcIcr-
mninIhs quaIIaIvc sIrucIurc s Ihcdcrcc Io vhch Ihc dllcrcnI
componcnIs ol a poIymcr nIcracI Io dcIcrmnc hIncss. Whcn rcpro-
ducIvc succcss s dcIcrmncd by caIa!yIc capacIy inIcracIons
bcIvccncomponcnIsmaIIcrbccauscas anKNApo!ymcrlo!dsIvoor
morc nuc!coIdcs lrom dllcrcnI parIs ol Ihc scqucncc may cnd up
IocIhcr lormn Ihc par| ol Ihc caIa!ysI IhaI rcconzcs Is IarcI,
causn Ihc hIncss olIhc po!ymcrIo dcpcnd on scvcra!componcnIs
aI oncc. y sysIcmaIca!!y varyn Ihc dcrcc ol nIcracIon amon
bI
bZ
N LbN N bMUL1N
componcnIs vc candcIcrmnc IhccllccIs IhaI Ihs lacIorhas on Ihc
Ioporaphy ol Ihc !andscapc. AI onc cxIrcmc Ihcrc arc no nIcrac-
Ions andIhc!andscapc s sIrucIurcdbyasn!c!oba!opImum: onc
bcsI caIa!ysI surroundcd by muIanIs dcc!nin smooIh!y n hIncss.
AI IhcoIhcrcxIrcmc, vhcnIhchIncss ol Ihcpo!ymcrnvo!vcsnIcr-
acIons bcIvccn a!! Is componcnIs, Ihc !andscapc s sIrucIurcd by
many !oca! opImaaroundvhchhIncss dccrcascssIccp!y.Thc hrsI
cxIrcmc maKcs cvo!uIon cnIrc!y prcdcIab!c vh!c Ihc sccond onc
maKcsIcnIrc!yunlcasb!c.ThcinIcrcsIncascsarc, Ihcrclorc, Ihosc
corrcspondn Io nIcrmcdaIc dcrccs ol nIcracIon. A parIcu!ar!y
mporIanI casc s obIancd vhcn Ihc numbcr ol componcnIs IhaI
nIcracI s sma!! rc!aIvc Io Ihc !cnIh ol Ihc po!ymcr: n IhaI casc
Ihcrc arcmany!oca! opIma vIh rc!aIvc!y hhhIncss, Ihcr sdcs
s!opcsmooIh!y,andIhchhcsIpcaKsIcnd IobroupcdIocIhcrn
parIcu!ar arcas ol Ihc !andscapc.
To undcrsIand hov Ihc sIrucIurc ol Ihc possb!Iy spacc allccIs
cvo!uIonaryva!Ksvc a!sonccdIodcIcrmncvhaIcxacI!ypcrlorms
Ihc va!K. Is I, lor cxamp!c, ndvdua!rcp!caIorscompcIn aansI
cachoIhcrors I cnIrc roupscompcInvIhoIhcr roups? Yhcn
consdcrnIhccvo!uIonol!arcanma!sandp!anIsIhchrsIa!Icna-
Ivc s morc p!ausb!c vh!c Ihc sccond onc s morc adcquIc Io dca!
vIh mo!ccu!ar popu!aIons Ihc mcmbcrs ol vhch canbc convcrIcd
inIo cach oIhcr by a sn!c muIaIon. IcIs manc a parIcu!ar KNA
po!ymcr vIh vcry hh hIncss !ocaIcd aI a !oca! opImum vIh
smooIh s!opcs. SponIancous muIaIons v!! Icnd Io producc many
oncmuIanI nchbors vIhsmiIarhIncss. Thcsc nchbors can a!so
muIaIcIoproduccs!hIIy!csshIIvomuIanInchborsIhaI,nIun,
can cncraIc IhrccmuIanI nchbors and so on. As vc movc avay
lroIhc hIIcsIpo!ymcrIhcnumbcr olvab!cmuIanIsdccrcascsunI!
Ihcr numbcr ladcs Iozcro, IhccnIrc scI lormn acohcrcnIclcudc]
muranrs. AIIhouh Ihc c!oud conIans a hIIcsI mcmbcrIhsdomnanI
Iypc maynoI bc Ihc onc IhaIrcproduccsIhcmosI, as I vou!d lIhc
compcIIon vas amon sn!c rcp!icaIors. Thc rcason s IhaI a non-
domnanI muIanI IhaI s surroundcdbyvcryhI oncs can ouI rcpro-
ducc Ihc domnanI onc, noI bccausc ol Is ovn capacIy Io maKc
copcsol IscIlbuIbccausc Is nchborscanbyasn!cmuIaIoncrc-
aIc copcs ol I. Ths amp!ihcaIon ol rcproducIvc succcss by ncarby
LNL1L LK1NMb N 1NL KL1L bU
muIanIsmcansIhaIIhcc!oudas avho!c, aso-caI!cd"quas-spcccs,
bccomcsIhcIarcIolsc!ccIon
IcIs manc a muIanI c!oudmovn ovcr a hIncss !andscapc n
vhch !oca! opIma ol dllcrcnI hchIs arc dsIrbuIcd n rc!aIvc
proxmIy, Ihc c!oud nIaI!yc!usIcrcd arounda !oca! opImum vIh
IhcdomnanIscqucnccaI Ihc pcaK.Ascvo!uIon procccdsparIolIhc
c!oudmaysIumb!c upon anoIhcr ncarby opImum ol hhcr hIncss.
AI IhaI momcnI Ihc inIa! c!oud viI! mc!I andrc-condcnsc around
Ihc ncvsinu!arIy.ThsmcansIhaI vhcn avho!cc!oud olmuIanIs
movcs in a hIncss !andscapc Ihc ovcra!! movcmcnI s noI a random
va!K, as onc vou!dcxpccI vcnIhcrandomncssolmuIaIons, buI a
sarch. noI a scarch vh a oa!, ol coursc, buI a ropn in Ihc darK
IhaI s ncvcrIhc!css bcIIcr aI hndn !oca! opIma ol hhcr hIncss
Ihana random va!K. Acruca!lacIor bchndIhsscarchn capacIy s
Ihc raIc aI vhchcopyncrrorsarcmadc, IhaI s, IhcmuIaIon raIc.
Il Ihc muIaIon raIcszcro Ihc c!oud docs noI lorm vh!c l I s Ioo
hh Ihc nIcrna!cohcrcncc olIhcc!ouds dcsIroycd. Asbclorc, I s
Ihc nIcrmcdaIc va!ucs IhaI yc!d Ihc nIcrcsIn cascs. U addIon,
Ihc cnIh olIhc po!ymcrs musIbcIaKcn nIo accounIsnccIhc !on-
cr Ihc mo!ccu!ar chanIhcmorc possb!c muIanIs I can havc. Thcsc
Ivo lacIors IocIhcr dcIcrmnc vhaI s

caI!cd Ihc rrcr rhrshcld, a

snu!arIysm!arIoaphascIransIonbccauscbcyondIhc Ihrcsho!d
Ihc "!qud c!oud Icnds Io "vaporzc. Thc Ihcory ol quas-spcccs
prcdcIsIhaIc!oudsvIhIhccapacIyIoscarchahIncss!andscapcvI!
Icnd Io !c car Ihc crrorIhrcsho!d, IhaI s, Io havc as many muIa-
Ionsaspossb!cvIhouIsullcrndsnIcraIon. EvdcnccIhaIIhss
nlacIIhc casc comcslromIaboraIorysIudcsol Ihc on!yconIcmpo-
rary oransms IhaI usc KNA Io sIorc cncIc inlormaIon, vruscs.
SIudcs ol vra! popuIaIons, asvcI! as ol popu!aIons ol naKcd KNA
cxIracIcd lrom Ihosc vruscs, conhrm IhaI Ihcy Icnd Io lorm quas-
spcccscsInnIhc vcnIy olIhccrror Ihrcsho!d.
Asn prcvous chapIcrs vc candcrvc lurIhcrph!osophca!nshI
l naddIonIoconsdcrnIhc IhcorcIcaI IrcaImcnIol cvo!uIonary
va!Ks on hIncss Iandscapcs vc cou!dacIua!!y loI!ovIhoscva!Ks n a
smu!aIon. Thc rcp!caIors n Ihcsc smu!aIons cou!d bc cquaI!y
"naKcd, IhaI s, cxsI n a dscmbodcd sIaIc and havc Ihc smp!csI
rc!aIonshpbcIvccn cnoIypc andphcnoIypc. Smu!aIonsusnIhc
bJ
b+
NLbN N b MUL1N
smp!csI sIns ol symbo!s as cnoIypcs | sIrns ol oncsandzcrocs)
nIo vhch poIcnIa!so!uIons Io phcnoIypc prob!cms arccodcd by
hand vou!d bc adcquaIc lor Ihc IasK. Thcsc naKcd cvo!uIonary
smu!aIons, cacdjnric aljcrirhms. arc so smp!c IhaI nonc ol Ihc
componcnIs ol an cvo!uIonary proccss-hIncss, sc!ccIon, sc!l-
rcp!caIon-s n lacIcmcrcnI.IIncsscan bcconsdcrcd Iocmcrc
sponIancous!y n a smuaIon l Ihc rcproducIvc succcss ol a vcn
cnoIypc dcpcnds on Ihc vay Ihc phcnoIypc I codcs lor acIua!!y
cxcrcscs Is capaIy Ioso!vcsurvva! prob!cms, aI Ihc vcry !casIIhc
prob!cm ol hov Io cxIracI cncry lrom cnvronmcnIa! radcnIs. U
oIhcrvords, IIhcphcnoIypchas a smu!aIcdmcIabo!sm andl Ihc
capacIyoIIhc cnoIypc Io crcaIc copcs ol Isc!l dcpcnds on posscs-
son ol amnmum olmcIabo!zcd cncry, Ihcn Ihc cva!uaIon ol Is
hIncss s pcrlormcd vIhn Ihc smuaIon Isc!l. U cncIc a!o-
riIhms, byconIrasI, hIncssscva!uaIcd lrom IhcouIsdcby a procc-
durcIodscrmnaIcoodIrombadso!uIonsIoaprob!cm,aproccdurc
ca!cda hIncssluncIon. Oncccachsymbo!sIrnhasbccnassncd
a prob!cm-so!vn va!uc anoIhcrproccdurc, ca!!cda scccIonIunc-
Ion, uscs Ihosc va!ucs Io choosc vhaI sIrns v!! bcncorporaIcd
nIo Ihc ncxI cncraIon and hov many copcs ol cach W bc pro-
duccd.U oIhcrvords,IhcIcndcncyoldllcrcnIrcp!caIorsIoncrcasc
nlrcqucncyvIhcachcncraIndocsnoIcmcrclromcompcIIon
buI s dcIaIcd by ru!cs IhaI Irans!aIc prob!cm-so!vn ab!Iy nIo
rcproducIvc succcss. Ina!!y, Ihc symbo! sIrns arc mosI!y ncrI
Ihc copynproccssspcrlormcdIorIhcm by anoIhcr proccdurc IhaI
app!cs Io Ihcm a muIaIon opcraIor Io producc varaIon pror Io
rc!casn Ihcm bacK nIo Ihc popu!aIon. Ths mcans IhaI muIaIons
do noI occur sponIancous!y as Ihc sIrns crcaIc lau!Iy copcs ol
Ihcmsc!vcs.
Thcon!yIhnIhaIcmcrcsnsmuaIonsusincncIca!orIhms
s spcchc so!uIons Io spcchc probcms. Thc lacI IhaI Ihc !aIIcr arc
poscd by Ihc cxpcrmcnIcr maKcs Ihc prob!cms Ihcmsc!vcs somc-
vhaI arIhca! buI vIhn Ihs !mIcd doman Ihc so!uIons arc
ndccd lound by cvo!uIonary scarch. A Iypca! smu!aIon ol Ihs
IypcIaKcsIhcIormolacycc.app!yIhchIncss luncIonIoIhccurrcnI
popu!aIion ol symbo! sIrns Io cva!uaIc Ihcr probcm-so!vn ab-
Ics: usc Ihc sc!ccIon luncIon Io dccdc vhaI hI sIrns v!! maKc
copcs oI Ihcmsc!vcs: app!y Ihc muIaIon opcraIor Io Ihc sc!ccIcd
LNL1L LK1NMb N 1NL KL 1L bU
sIrnsIocncraIcvaraIonandlormIhcncxIcncraIon:rcpcaIa!!
loursIcpsunI!a!oba!!yor!oca!yopIma!so!uIonhasbccnlound.
l
Thc prob!cms Io vhch cncIc a!oriIhms arc app!cd havc olIcn
noIhn Io doviIhbo!oy. A ood !!usIraIon n an ndusIra! con-
IcxI s Ihc conIro ol a ppc!nc lor naIura as. A ppc!nc musI !nK
IhcponIolsupp!yolnaIura!asIoIsponIol consumpIon ordsIr-
buIon usn a scrcs ol comprcssorsnKcd by ppcs. Thc prob!cm s
IodcIcrmincIhc rc!aIonbcIvccnIhc sucIonprcssurcol cach com-
prcssor IoIsdscharcprcssurc |IhcprcssrcradcnIbcIvccnnpuI
andouIpuI) nsuchavayasIomnmzcIhcovcra!!c!ccIrca!povcr
consumcd. U oIhcr vords, Ihc prob!cm s Io hnd Ihc combnaIon
olprcssurcvaucslorcachcomprcssorIhaIopImzcsIhcclhccncyol
cncry usc..l Ior Ihc saKc ol smp!cIy vc manc IhaI cach com-
prcssor can havc prcssurc va!ucs bcIvccn zcro and hlIccn Ihcn a
sIrnollouroncsandzcrocsv!!bccnouhIorcprcscnII,IhcsIrn
0000"rcprcscnInIhcminmumva!ucand "I I I I rcprcscnInIhc
maxmum onc. Sncc vc havc many comprcssors scvcra! ol Ihcsc
"cncs musI bc concaIcnaIcd Io lorm Ihc cnIrc "chromosomc.
Ina!y, a hIncss luncIon musI bc crcaIcd Io cva!uaIc Ihc povcr
consumpIonol cach combnaIon ol va!ucs lordllcrcnIcomprcssors
as vc! as Io cnlorcc !oba! consIranIs. such as Ihc minmum or
maxmumolprcssurca!ovcdnIhc ppc!nc. Whcn a popuaIon ol
Ihcsc varab!c rcp!caIors s uncashcd n a compuIcr I can hnd Ihc
opIma! combnaIon ol prcssurc va!ucs lor Ihc cnIrc ppc!nc n a
hhyclhccnIvay.
UnKcIhsndusIra!app!caIonusncncIc a!orIhmsIomodc!
naKcd KA n Ihc prmorda! soup runs nIo somc unso!vcd prob-
!cms. Wh1cIhcsmu!aIcdcnoIypcssI!rc!aIvc!ysmpcIobu!d
Ihc lour nuc!coIdcs bcn cas!y cxprcsscd n bnary lorm as 00. 0I .
I 0. ! !-crcaInaphcnoIypcmp!csIranslormnaonc-dmcnsona!
chanol smu!aIcdnuc!coIdcs nIo aIhrcc-dmcnsona!lo!dcd lorm.
BuI asvcsavinIhcprcvouschapIcr arc!abcmcchanca!rccpcIo
pcrlorm Ihc lo!dn opcraIon docs noI ycI cxisI. cIhcr docs a
mcIhod Io cvauaIc Ihc caIa!yIc capacIy IhaI such a lo!dcd lorm
vou!d havc, an cvauaIon IhaI vou!d bc nccdcd Io crcaIc a hIncss
luncIon. So smu!aIons ol prcboIc condIons musI scII!c lor !css.
vh1cIhclu!Ihrcc-dmcnsona!shapccannoI bc cncraIcdannIcr-
mcdaIcsIcpnIhclo!dnproccssnvo!vnaIvo-dmcnsona!shapc
bb
b
NLbN N blMUL1lN
|a "sccondarv sIrucIurc") can bc rcIiabIv produccd. And vhiIc vc
cannoI caIcuIc Ihc caIaIvIic capaciIv lrom Ihis inIcrmcdiaIc lorm
vc can caIcuIaIc Ihc cncrcIic sIabiIiIv IhaI Ihc loIdcd lorm vouId
havc This mcasurc ol hIncss is jusIihcd bccausc an unsIabIc shapc
vouId noI IasI Ion cnouh Io cxcrcisc iIs caIaIvIic capac|Iics. Il vc
Ihink ol a loIdcd KA poIvmcr as Ihc lorm produccd bv a radicnI
as iI dissipaIcs, Ihcn Ihc sIabIc loIdcd lorms vouId bc Ihosc IhaI
manacd Io achicvc a combinaIion ol bonds IhaI luIIv dissipaIc Ihc
radicnI.ThissIabiIiIvcan bc caIcuIaIcdlromIhcsccondarvsIrucIurc
and uscdIocva!uaIc hIncss.
Givcn Ihis simpIihcd rcIaIion bcIvccn cnoIvpc andphcnoIvpc a
cncI|c aIoriIhm can bc uscd Io loI!ov a popuIaIion ol simuIaIcd
KAmoIccuIcs across manv cncraIions and chcck vhcIhcr sIabIc
lormscmcrcIhaI can bccomc Ihc ccnIcrs olquasi-spccics. Onc sim-
uIaIion, lor cxampIc, sIarIcd viIh a popuIaIion ol 3, 000sIrins, cach
70 componcnIs Ion, buI in vhich aII componcnIs vcrc idcnI|caI.
Sincc such a homocnous sIrin vouId bc incapabIc ol loIdin Ihc
popuIaIion sIarIcd iIs cvoIuIionarv vaIk aI a zonc in Ihc Iandscapc
IhaIvasa "!IaIpIain" ol vcrvIovhIncss, andiIhadIohndandcIimb
a "hiII" rcprcscnIin a IocaI opIimum. Thc mosI imporIanI paramcIcr
IhaIcouIdbcvaricdin IhcsimuIaIionvas IhcmuIaIionraIc, Ihc lrc-
qucncvviIhvhichcopvincrrorsvcrcmadc,sinccasvc savbclorc
IhisparIIvdcIcrmincsIhccohcrcncc olIhcmuIanIcIoud. ThcmuIa-
Iion raIc is normaIIv kcpI hxcd in mosI appIicaIions ol cncIic aIo-
riIhmsbuIinIhisonciIhadIobcvaricdlromoncrunolIhcsimuIaIion
Io anoIhcr Io IcsI Ihc cxisIcncc ol Ihc crror IrcshoId. Thc rcsuIIs ol
Ihc simuIaIion conlormcd Io Ihc IhcorcIicaI prcdicIion: Ihc popuIa-
Iion olsIrinscvcnIuaIIvlounda IocaIopIimumandbccamc capabIc
ol loIdin |as inlcrrcd lrom Ihc sccondarv sIrucIurc) buI onIv il Ihc
crrorraIcvasbcIovIhc criIicaI IhrcshoId. ' Thisis arcIaIivcIvIimiIcd
rcsuII parIicuIarIvil vc considcr vhaIIhccvcnIuaI oaI ol Ihiskind
ol simuIaIionsvouIdhavcIobc: IoshovhovIhcscarchincapaciIv
olmoIccuIarrcpIica orscouIdhavcIcdIoIhccmcrcnccolIhccncIic
codc. And noI nIv Ihc codc iIscIl, IhaI is, Ihc riid corrcspondcncc
bcIvccn Ihrcc nucIcoIidcs in onc Ivpc ol poIvmcr |cncs) and a
sinIcaminoacidinanoIhcrIvp |proIcins) , buIIhcmcansIorcIiabIv
androuIincIvIransIaIc onc inIo Ihc oIhcr.
LNL1lL L1NMb N 1NL L 1L bU
WhiIcIhcnakcdrcpIica IorsolcncIicaIoriIhmsarcidca!IomodcI
Ihc primordia!soup priorIoIhc cmcrcncc olIhccncIiccodc a dil-
lcrcnIapproachisnccdcdIoIrackcvoIuIionalIcrIhaIsinuIarcvcnI.
InparIicuIar, vcnccdIhcmcansIoincrcascIhcIcvcIolcmbodimcnI
olIhc rcpIicaIors ivcn IhaI Ihc coupIin ol cncs andproIcins couId
noI havc occurrcd viIhouI Ihc cncapsuIaIion ol Ihcsc inIcracIin
poIvmcrs viIhin primiIivc mcmbrancs. And as vc considcr morc
cmbodicd rcpIicaIors Ihc rcIaIion bcIvccn cnoIvpc and phcnoIvpc
musIbc modcIcdin morc dcIaiI. vcn Ihc carIicsI ccIIuIar oranisms
had Io soIvc mcIaboIic probIcms bv usin Ihcir IimiIcd rcpcrIoirc ol
proIcinsandcnzvmcsasrccomb|nabIccomponcnIsIolormprimiIivc
chcmicaI circuiIs. So simuIaIions ol ccI!uIar cvoIuIion musI havc Ihc
ncccssarv rcsourccs Io ivc risc Io simpIc parI-Io-vhoIc rcIaIions
bcIvccn dillcrcnI IcvcIs ol Ihc phcnoIvpc. And il Ihcsc rcIaIions arc
Io bc inhcriIabIc Ihc cnoIvpc |IscIl musI bc modcIcd bv svmboI
sIrins capabIc olincrcmcnIaIcomposiIionaIiIv. n Ihcprcvious chap-
Icr vc sav IhaI simpIc lorms ol chcmicaI circuiIrv | auIocaIaIvIic
Ioops) couId cmcrcina soup olsvmboI sIrinsbcIoninIo arccur-
sivc luncIion Ianuac, IhaIis, in aTurinas. Wc aIsosavIhaIIhosc
Ianuacs aI!ov lor compIcx compuIcr prorams Io bc cncraIcd
usin a parI-Io-vhoIc rcIaIion: IhcsimpIcsIluncIions can bc uscdIo
crcaIcmorccompIcxluncIionsIhaI,inIun, can cnIcrinIocvcnmorc
compIcx oncs. ThccompuIcrproramssocncraIcdcanbc cxprcsscd
as svmboI sIrins in vhich Ihc ncsIcd luncIions aI dillcrcnI IcvcIs
arc scparaIcd bv parcnIhcscs. Thc branch ol cvoIuIionarv compuIa-
Iion usin Ihcsc morc sophisIicaIcd svmboI sIrins is caIIcd jnric
rrcjrumminj.
U lacI, incncIicproramminIhc simuIaIcdchromosomcs arcnoI
impIcmcnIcd as sIrins buI as raphs havin Ihc lorm ol a Ircc. This
cIiminaIcs IhcpIcasin simiIariIv bcIvccn a chain ol moIccuIcs and
a sIrin ol svmboIs buI compcnsaIcs lor iI bv cIcarIvdispIavin Ihc
parI-Io-vhoIcrcIaIionbcIvccnprobIcmsandsubprobIcms: Ihc ovcr-
aIImcIaboIic Iask lorms IhcrooI ol Ihc IrccvhiIcprorcssivcIv morc
dcIaiIcdsub-Iasks lormIhcbranchcsolIhc Ircc. U indusIria!appIica-
Iions Ihc povcr ol cncIic prorammin has bccn cIcarIv dcmon-
sIraIcd bv oin bcvond Ihc soIuIion Io opIimizaIion probIcms Iikc
Ihc as pipcIinc discusscd abovc. In parIicuIar, cncIic prorammin
b
bd
NLbN N bMUL1N
can IackIc morc chaIIcnin dsijn rrcblms. Onc cxampIc is Ihc
dcsin ol anaIo cIccIricaI circuiIs, anarca in vhich human cxpcrIisc
hasIradiIionaIIy bccn rcquircd andinvhichIhc dcrcc oloriinaIiIy
ol a ivcn dcsin can bc rouh!y asscsscd by Ihc lacI IhaI a paIcnI
olhcc has acccpIcd iI. Usin cncIic prorammin popuIaIions ol
rcpIicaIorsscarchin Ihc spacc ol possibIc circuiIshavc rcdiscovcrcd
scvcraI circuiI dcsins IhaI had bccn prcviousIy paIcnIcd: havc dis-
covcrcd novcI csins IhaI maIch Ihc luncIionaIiIy ol paIcnIcd
dcsins: and in aI IcasI onc casc Ihcy havc lound an cnIircIy ncv
dcsin, IhaIis, onc noIIoicaIIydcducibIclrom aprcviousIypaIcnIcd
invcnIion. '' This abiIiIy Io cncraIc non-obvious dcsins in vhich
componcnIs arc inIcrconnccIcd in compIcx vays canbc vcry uscluI
IocxpIorcIhc cvoIuIionaryoriinolIhcmcIaboIiccircuiIryoluniccI-
!uIar oranisms.
To shov hov Ihis couId bc donc IcIs bcin by dcscribin hov
cncIicproramminvorks in appIicaIionsnoIinvoIvinIhc modcI-
in ol bioIoicaI proccsscs. IirsI ol aII, ivcn IhaI compIcx prorams
arc cvoIvcd by rccursion, IhaI is, by luncIions IhaI rcsuII lrom Ihc
composiIion ol simpIcr oncs, vc nccd a scI ol cIcmcnIary luncIions
IocIIhcproccsssIarIcd.ThcscprimiIivcluncIionslormIhc nodcs or
branchin poinIs olIhcIrcc.Wc aIsonccdIospccilyIhcvariabIcsand
consIanIs IhaI Ihosc cIcmcnIary luncIions usc as Ihcir inpuIs |Ihc
"Icavcs" or IcrminaIs ol Ihc Ircc) . Thcsc basic componcnIs musI bc
choscnbya dcsincr IomaIchIhcIypc ol probIcm Iobc soIvcd. ' `U
Ihc casc ol anaIo cIccIricaI circuiIs, lor cxampIc, Ihc cIcmcnIary
opcraIionsincIudcluncIionsIhaIinscrIancvcomponcnI |a rcsisIor,
a capaciIor, aninducIor):luncIionsIhaI aIIcrIhcconnccIiviIyolIhosc
componcnIs |Ihc IopoIoy ol Ihc circuiI) : and luncIions IhaI scI Ihc
inIcnsiIy |or sizin) ol acomponcnI, IhaI is, Ihc dcrcc ol rcsisIancc
ol arcsisIor, Ihc capaciIancc ol a capaciIor, and so on. U Ihc casc ol
diiIaI circuiI dcsin Ihc cIcmcnIary luncIions shou!d bc opcraIors
Iikc "And, "Or, and "NoI. U Ihc casc ol roboIic moIion, IhaI is,
vhcn Ihc probIcm Iobc soIvcd is Ihc synIhcsis ol moIion paIhs, Ihc
cIcmcnIary rccrIoircmusIconIain luncIions Iikc "TurnIclI, "Turn
KihI, and "Novc Iorvard. Thcsc cIcmcnIary luncIions arc Ihc
componcnI ol Ihc cvoIuIionary proccss IhaI is noI cmcrcnI sincc
Ihcir bchavior |Ihc vayIhc luncIions allccIIhcir inpuIs) is dicIaIcdby
ruIcs. AnycompIcx luncIinIhaIcanbccrcaIcdlrom Ihc cIcmcnIary
LNL1L LK1NMb N 1NL KL l1L bU
oncsby rccursion, onIhcoIhcrhand,canbc considcrcd cmcrcnIil
iIcvoIvcs sponIancousIyas aspccihsoIuIionIoa dcsinprobIcm.'
ThchnaIsIcp is Io crcaIc a hIncss luncIion dchnin Ihc probIcm
Io bc soIvcd. Sincc Ihc cvoIvin cnIiIics in Ihis casc arc compuIcr
prorams cvaIuaIin Ihcir hIncss dcmands acIuaIIy runnin Ihcm
bccausc vhaI isasscsscdis Ihc bchavior ol Ihc proram, or Ihc con-
sIrucIion capaciIics viIhvhich IhaI bchavior cndovs Ihc proram.
ThiscvaIuaIionispcrlormcdbycomparinIhcrunninproramviIh
a scI ol IcsI cascs IhaI aIrcady posscss Ihc rcquircd capaciIics, Ihc
dcrcc ol hIncsscxprcsscdas a disIancc lrom IhaI IarcI. In Ihc casc
ol anaIo cIccIricaIcircuiIs Ihc cvaIuaIion ol hIncss is, in lacI, morc
compIcx bccausc vc nccdIochccknoI onIy IhaI a runnin proram
canconsIrucIIhc circuiIbuIaIso IhaIIhc circuiIiIscIl bchavcs as iIis
inIcndcd Io: as a hIIcr, as an ampIihcr, as a scnsor. To do Ihis an
"cmbryocircuiI|ancIccIricaIsubsIrucIurcviIhmodihabIcvircsand
componcnIs) is pIaccdinIo aIarcrcircuiIinvhichnocomponcnIis
modihabIc. OnIyIhc cmbryo cvoIvcsbuI iIs pIaccmcnIinIoa Iarcr
luncIionaIscIIinaIIovsiIIobcchcckcdlorviabiIiIy. ' 'AsiIhappcns,
Ihc insihIs and Icchniqucs dcvcIopcd in Ihc appIicaIion ol cncIic
prorammin Io cIccIricaI circuiIs can bc uscd Io modcI ccIIuIar
mcIaboIismsbccauscinmany cascs Ihcrc isanisomorphismbcIvccn
IhcIvo.
IcIs cxaminc a concrcIc cxampIc. Thc IarcI ol a simuIaIion ol
IhisIypc is anacIuaIpiccc ol biochcmicaI machincry IhaIhas bccn
sIudicdin IhcIaboraIory. ThisIypicaIIyincIudcsa ncIvorkolchcmi-
caIrcacIins: a scIolcnzymcsacIinascaIaIsIs: andaniniIiaIscIol
conccnIraIionradicnIsolIhcsubsIanccsscrvinassubsIraIcs.In Ira-
diIionaI biochcmisIry dillcrcnIiaI cquaIions arc uscd Io modcI Ihc
raIcsaIvhichIhc subsIraIcsarcIranslormcdinIociIhcrinIcrmcdiaIc
or hnaIproducIs, asvcIIasIhc vayinvhichIhccaIaIysIsallccIIhosc
raIcs by acccIcraIin Ihcm or dcccIcraIin Ihcm. Thcsc cquaIions
arc Ihcn uscd Io prcdicI Ihc conccnIraIion ol Ihc hnaI producIs, a
prcdicIionIhaIcanbc IcsIcdby carryin ouIacIuaI cxpcrimcnIs. Thc
qucsIion lor cncIic proramminis Ihis: ivcn daIa abouIIhc iniIiaI
and hnaI subsIancc conccnIraIions can vc rcvcrsc cninccr Ihc
chcmicaI ncIvorkIhaI!nksIhc Ivo? ThisvouIdinvoIvc discovcrin
Ihc IopoIoyansizinolIhcncIvork, IhaIis, Ihc connccIiviIyolIhc
dillcrcnI rcacIions and IhcvaIucs olIhc raIcs aIvhich Ihcy procccd.
bV

NLbN N b MUL1N
U onc smu!aIon, lor cxamp!c, an acIua! mcIabo!c paIhvay usin
lourdllcrcnIchcmca!rcacIonsvasuscdasaIarcI. ThcIopo!oyol
Ihc paIhvay vas comp!cx: I conIaincd an nIcrna! lccdbacK!oop a
blurcaIon ponI aI vhch onc subsIraIc vas uscd lor Ivo dllcrcnI
rcacIons and an accumu!aIon ponI aI vhch Ihc conccnIraIon ol
asubsIanccvasallccIcdbyIvodllcrcnI sourccs.
l8
ThcprmIvcluncIonsuscdnIhssmu!aIonnc!udcdmaIhmaI-
ca!cquaIonsdcscrbnchcmca!rcacIonsvh!cIhcIcrmna!svcrc
Ihc npuIs Io IhosccquaIons, such as subsIraIcs and caIa!ysIs. oIh
ol Ihcsc camc lrom modc!s uscd by bochcmsIs and vcrc Ihcrclorc
noI cmcrcnI. On Ihc oIhcr hand, Ihc nIa! popu!aIon ol random
proramshad no inlormaIon abouI hov many rcacIons Ihc mcIa
bo!c paIhvayvas supposcd Iohavc, hov Ihcsc rcacIons shou!d bc
coup!cdIooncanoIhcr, orvhaIsubsIraIcscachrcacIonoIasnpuIs
and vhaI producIs I produccd as ouIpuIs. Thc on!y nlormaIon Ihc
cvo!uIonary proccss had abouI Is IarcI vas bu!I nIo Ihc hIncss
luncIon: Ihc conccnIraIon ol Ihc hna! producI ol Ihc mcIabo!c
paIhvay. Thc rcsu!Is ol Ihc smu!aIon vcrc cncouran. A nIa!
popu!aIonol I00,000rcp!caIorsvasab!cIodscovcrIhcIopo!oyol
Ihcchcmca!ncIvorK |nc!udnIhclccdbacK !oop, andIhcblurca
Ion andconccnIraIon ponIs) n abouI I20 cncraIons, vh!cIhc
sznol IhcncIvorK |Ihc va!ucs lor raIcs ol conccnIraIon andraIcs
ol caIa!yss) vas c!osc!y approxmaIcd in an addIona! I 00 cncra
Ions.
lV
Thccva!uaIonolIhchIncssolcachcncraIonvaslac!IaIcd
by cxp!oIn Ihc somorphsm ol chcmca! and c!ccIrca! crcuIs:
vo!Iac radcnIs p!aycd Ihc ro!c ol conccnIraIon radcnIs, vh!c
crcuIs IhaIpcrlormcd addIon andsubIracIon rcprcscnIcdIhcpro
ducIon andconsumpIonol avcnsubsIancc.Onccansomorphc
c!ccIrcacrcuI vas crcaIcd I cou!d bc IcsIcd lor hIncss usn Ihc
samc approach dscusscd abovc: p!acn an cvo!vn cmbryo crcuI
nIo a !arcrnon-cvo!vn onc and usn sIandard solIvarc Io chccK
lorvab!Iy.
Thc conc!uson vc may drav lrom IhsandIhc prcvousbo!oca!
smu!aIon s IhaI Ihc capacIy ol a popu!aIon olvarab!crcp!caIors
Ioscarchapossb!Iyspaccsndccdrca!. OnIhcoIhcrhand, dcmon
sIraInIhccxsIcnccol acapaIysnoIIhcsamcIhinascxp!ann
I. Thc smp!csI cxp!anaIon, Ihconc app!yin IonaKcdK vou!d
rc!y on Ivo assumpIons: IhaI Ihc spacc ol possb!c po!ymcrs docs n
LNL1L LK1NMb N 1NL KL 1L bU
lacIposscssIhcordcrvcmposcdonI,IhaIs,IhaIIhcscqucnccsarc
arrancd so IhaI onc-muIanI varanIs occupy nchborn posIons,
and IhaI Ihc scarch s carrcd ouI n para!!c! by an cnIrc c!oud ol
muIanIs. uI oncc Ihc rc!aIon bcIvccn cnoIypc and phcnoIypc
sIaKcn nIoaccounI Ihs cxp!anaIon ccascs Io bcsulhccnI. Evcn n
Ihc casc ol naKcd KA Ihc Iopo!oy ol Ihc possb!c bccomcs morc
comp!caIcd. InparIcu!ar, IoundcrsIandhovonclo!dcdcaIa!ysIIhaI
IarcIsaccrIanchcmca!rcacIoncou!d cvo!vc nIoanoIhcroncvIh
a dllcrcnI IarcI, Ihc paIh lrom onc phcnoIypc Io Ihc oIhcr musI
bc a!so rcprcscnIcd n Ihc vay Ihc spacc s ordcrcd. Onc vay ol
achcvn Ihs s Io mposc an ordcr IhaI cnsurcs IhaI bcIvccn Ivo
mo!ccu!ar scqucnccs, cach onc occupyn a !oca! hIncss opImum,
Ihcrc arc "rdcs onn IhcIvo "mounIan pcaKs, rdcs n vhch
Ihc scqucnccs a!! havc Ihc samc hIncss. Ts vay a muIaIon cou!d
Iranslorm onc scqucncc nIo a nchborn onc vIh Ihcsamc hIncss
vh!c Kccpn Ihc IranslormaIon nvsb!c Io sc!ccIon prcssurcs.
A scrcs ol such ncuIra! muIaIons cou!d Ihcrclorc !cad lrom onc
lo!dcdcaIa!ysIIoavcrydllcrcnIoncbyva!KnacrossIhcrdccvcn
uIhcrcarczoncsol !ovcrhIncssbcIvccn IhcmounIan pcaKs.''
AdllcrcnIKndolprob!cmviIhIhcordcrvcmposconIhcoIhcr
vsc amorphous spacc ol possb!c po!ymcr scqucnccss IhaI n add
Ion Io muIaIon somcuncc!!u!aroransmsuscsxalrccmbinaricn.
WhcnIvodllcrcnIchromosomcs arcscxua!!ycombncdcachparcnI
conIrbuIcsha!l ol Ihc cncscomposnIhc ollsprns chromosomc.
Il vc manc IhaI IhcanccsIors ol cach parcnI hadc!imbcda dllcr
cnI !oca! hIncss opImum Ihc rcsu!In ollsprn vou!d la!!bcIvccn
IhcIvopcaKs,IhaIs, nanarcaol!ovcrhIncss, andvou!dIhcrclorc
bcc!mnaIcd by naIura! sc!ccIon. ThsncaIvccllccI maybc amc-
!oraIcd lIhc !andscapc has Ihc rhI Ioporaphy, IhaI s, la!!!oca!
opIma arc roupcd IocIhcr lormn a IhI c!usIcr ol pcaKs. uI
a morc cncra! so!uIon may lorcc us Io rcIhnK hov Ihc spacc ol
possb!cpo!ymcrsshou!d bc ordcrcd so IhaI Is connccIvIy s com
paIb!cvIhboIhmuIaIonandscxua!rccombnaIon.Thss parIcu
!ar!y mporInI vhcn vn an accounI ol Ihc cmcrcnI scarchn
capacIy ol smu!aIons bascd on cncIc a!orIhms or cncIc pro
rammn bccausc B mosI ndusIra! app!caIons Ihcsc lavor Ihc
uscola crcsscvrcrrarcrovcramuIaIonopcraIor asamcansIocn
craIcvaraIon. Thc crossovcropcraIor vorKs Ihs vay: I IaKcs Ivo
I
Z
NLbN N bMUL1N
scparaIc simu!aIcdchromosomcsandpicks aparIicu!arpoinIincach
onco!Ihcm:iIIhcnbrcaksboIhaIIhaIpoinIcrossinonco!IhcIwo
ha!vcs !rom onc parcnI ovcr Io Ihc oIhcr: hna!!y iI rcaIIachcs Ihc
IwoparcnIa!ha!vcsbc!orcrc!casinIhc o!!sprininIoIhcpopu!aIion.
In oIhcr words, Ihc crossovcr opcraIor mimics Ihc c!!ccI o! scxua!
rccomb

naIion.
Onc p!ausib!c cxp!anaIion !or Ihc supcrioriIy o! crossovcr ovcr
muIaIionsIarIswiIhIhcassumpIionIhaIIhcsurviva!prob!cmposcd
Ioancvo!vinpopu!aIioncanbc dccomposcdinIosubprob!cmscach
o!whichcanbc so!vcd scparaIc!y. InoIhcrwords, Ihc assumpIionis
IhaIcomp!cx adapIivc prob!cms IhaI havc a modu!arsIrucIurc can
bc so!vcd cn buildinj blcck ar a rim. wiIh scxua! rccombinaIion
p!ayin Ihc ro!c o! brinin scparaIc bui!din b!ocks IocIhcr inIo
a !u!! so!uIion. To cxp!orc Ihis sccnario a spccia! mcIhod has bccn
dcvc!opcdIokccpIracko!IhcpropaaIiono!bui!dinb!ocksinsimu-
!aIionsusincncIica!oriIhms.I!wcimaincIhcsimu!aIcdchromo-
somcs Io bc Icn biIs !on, Ihc popu!aIion conIainin sIrins !ikc
I I 0I I 00000" or 00I I 0I I I I 0, " wc can idcnIi!y Ihc simi!ariIics
amonsIrinswiIhanoIaIionusinwi!dcards. "Thc IwosIrins|usI
mcnIioncd, !or cxamp!c, havc simi!ar cncs" aI posiIions !ourand
Icn, a simi!ariIyIhaIcanbc rcprcscnIcdby a sIrin usin awi!dcard
symbo! !ikc #. " Thc sIrin ###I #####0"is ca!!cdaschcma Oncc
cvcry cncraIion cachmcmbcr o! Ihc popu!aIion o! sIrins is cva!u-
aIcdbyIhchIncss!uncIion!oriIspcr!ormanccasaso!uIionIoaprob-
!cm. I! Ihcpopu!aIionconIaincd,say, 50 sIrins, IhaIwou!d covcr a
rc!aIivc!y sma!! arca o! Ihc possibi!iIy spacc. BuI i! in addiIion wc
cva!uaIcdIhcavcracpcr!ormancco!cachschcmawcwou!dcxpand
Ihc arca scarchcd. For cxamp!c, i!Ihc IwosIrinsabovc bc!oncdIo
Ihc popu!aIion buI noI Ihc sIrins I 00I I 000I 0" or 0I I I 0 I I I 00. "
Ihcn Ihc !aIIcr Iwo wou!d noI bc cxp!iciI!y cva!uaIcd. BuI bccausc
Ihcsc Iwo sIrins arc a!so insIanccs o! ###I #####0" wc wou!d cI
in!ormaIion abouI Ihcm whcncva!uaIinIhc avcrac hIncss o! Ihc
schcma.
U wc Ihink ocachsIrin inIhc popu!aIionas a poinIin a mu!Ii-
dimcnsiona! possibi!iIy spacc Ihcn a schcma wiIh many insIanccs is
!ikcas!icc [orhypcrp!anc) o!IhaIspacc.

Thus,apopu!aIioninwhich
sIrins sharc bui!din b!ocks wou!d noI bc !ikc a c!oud o!poinIsmov-
inIhrouhIhcpossibi!iIyspaccbuIraIhcr!ikcascIo!s!iccscovcrin
LNL1L LK1NMb N 1NL KL L bU
a muchwidcr arca and Ihcrc!orc incrcasin Ihc para!!c!ism o! Ihc
scarch. Thc crossovcr opcraIor, onIhc oIhcr hand, cancasi!ydcsIroy
bui!din b!ocks i! Ihc cuI poinIhappcns Io scparaIc iIs componcIs.
This imp!ics IhaI an insIancc o! a schcma in which Ihc hxcd posi-
Iions arc c!osc Io onc anoIhcrhas a bcIIcr chancc o! prcscrvin iIs
bui!din b!ocks a!Icr many rccombinaIions. insIanccs o! Ihc schcma
#I 0#######," !or cxamp!c, havc a bcIIcr chancc o!prcscrvinIhc
bui!dinb!ock I 0" IhanIhoscbc!oninIo ###I #####0. "II canbc
shown IhaI a schcma wiIh many wi!d cards [rcprcscnIin bicr
s!iccso!Ihcpossibi!iIyspacc) andinwhichIhc!cwhxcdposiIionsarc
c!usIcrcd IocIhcr can propaaIc Ihrouh a popu!aIion as !on as
Ihc hIncss o! Ihc schcma is rcaIcr Ihan Ihc avcrac hIncss oIhc
popu!aIion. This rcsu!I is known as Ihc schcma Ihcorcm. " Thc
schcma Ihcorcm, howcvcr, says noIhin abouI how bui!din b!ocks
arc brouhI IocIhcr ovcr many cncraIions Io bc combincd inIo a
!u!! so!uIion Io Ihc prob!cm. In oIhcr words, Ihc schcma Ihcorcm
cxp!ainsIhc propaaIion o! oodbui!dinb!ocks buI noI Ihcir accrc-
Iion.

` BuI Ihc!aIIcris cqua!!yimporIanI bccausc as bui!din b!ocks

comc IocIhcr Ihc cvo!uIionary scarch is conducIcd on spaccs o!
cvcr dccrcasin dimcnsiona!iIy and cvcr incrcasin hIncss, IhaI is,
cach schcma rcprcscnIs an incrcasin!y !arcr s!icc o!Ihc spacc and
conIains scvcra!parIia!so!uIionsIoIhc ovcra!!prob!cm.
Is Ihcrc any cvidcncc !or Ihc cxisIcncc o!bui!dinb!ocks in rca!
popu!aIions o! rcp!icaIors? To scparaIc buI rc!aIcd discovcrics
in biochcmisIry and cncIics bcar on Ihis qucsIion. On onc hand,
biochcmisIssIudyinproIcinsandIhcir!o!dcdsIrucIurcshavcidcnIi-
hcdIhoscparIso!Ihcirsur!accsIhaIp!ayaro!c incnzymaIicacIiviIy.
Ihcir bindin siIcs and Ihcir caIa!yIic siIcs. Bccausc Ihcsc siIcs arc
rc!aIivc!y compacI and cxhibiI spccihc !o!dcd moIi!s [shccIs, hc!iccs,
ribbons) Ihcy arc namcd domains. " Simi!ar domains in di!!crcnI
proIcins Icnd Io havc simi!ar!uncIions, a !acI IhaI sucsIs IhaI Ihcy
arc indccd rcusab!c bui!din b!ocks.

On Ihc oIhcr hand, cncIicisIs

havc discovcrcd IhaI Ihc cncs o!unicc!!u!aroranismsinwhichIhc
cncIic maIcria!s arc cncapsu!aIcd inIo a nuc!cus arc noI sIrucIurcd
as onc conIinuous uniI as Ihcy arc in bacIcria IhaI !ack a nuc!cus.
InsIcad, Ihcir cncs havc a mosaic sIrucIurc in which scqucnccs
o!nuc!coIidcsIhaIcodc!orproIcins [ ca!!cd cxons") arc inIcrspcrscd
wiIh scqucnccs IhaI do noI codc !or anyIhin [ ca!!cd inIrons") .
J
9
NLbN N bMUL1N
U manycascsoncormorccxonscodc!oraspccihcdomaininapro-
Icin,IhaIis,!orabui!dinb!ock, socxonscanbc rcshu!BcdIhrouh
scxua!rccombinaIionIoproducc novc!proIcins aIrc!aIivc!y!ow cosI,
ahypoIhcsismadcmorcp!ausib!cbyIhc!acIIhaIcachproIcindomain
has iIs own !o!din capabi!iIics so IhaI ncw combinaIions can bc
cxpccIcdIo!o!dIocIhcrncaI!y.

'AddiIiona!!y,IhcprcscnccoinIrons
shou!d !owcr Ihc risk o! disrupIion o! cxons. i! Ihc cuI poinI uscd
inscxua!rccombinaIion!icsonaninIronIhcnIhc cocvo!vcdcompo-
ncnIs o!abui!dinb!ockwi!!noIbc disrupIcd.
cspiIc iIs morc c!aboraIc mappin bcIwccn cnoIypc andphcno-
Iypc Ihc rcp!icaIors uscdincncIicproramminrcmaindiscmbod-
icd andarcnoI siIuaIcdinspacc. AndIhc !acIIhaIIhccva!uaIiono!
hIncssispcr!ormcd cxocnous!y, IhaIis, IhaIIhcIarcI o!Ihc scarch
isscIbyIcsIcascsprovidcdbyIhc dcsincr,imp!icsIhaIIhcproccssis
morc !ikc Ihc conIro!!cd brccdin o! !arm anima!s and p!anIs by
humanbcinsIhan an cvo!uIionaryproccss in which somc spccics
acIsassc!ccIionprcssurcsonoIhcrspccics.ThismcansIhaIqucsIions
invo!vinIhccocvo!uIiono!spccicsorIhccmcrcncco!!oodwcbsin
ccosysIcmscannoIbccxp!orcdusinIhcsc simu!aIions. H Ihc!o!!ow-
inchapIcrwcwi!!rcmovcIhcsc!imIaIionsbycmbodyinIhcrcp!i-
caIors,siIuaIinIhcminspacc,andmakinIhcirrcproducIivcsucccss
dcpcnd on Ihcir capaciIy Io mccI IhcirmcIabo!ic nccds by Iappin
inIo rcsourcc raicnIs disIribuIcdinIhaI spacc.
LHPJL VL
OOIC |O|I005 d0 CI O |d|505
WiIhIhc cmcrcncc o!Ihc hrsI!ivincrcaIurcsIhc aqucouscnviron-
mcnI wc havc bccn cxp!orin acquircd a ncw !caIurc. a c!aIinous
sIraIum o! co!onia! bacIcria aI Ihc inIcr!acc bcIwccn boIIom scdi-
mcnIs and waIcr !ormin Ihc car!icsI biosphcrc. This ncw sIraIum
rcmaincd dcccpIivc!y simp!c !or ovcr Iwo bi!!ion ycars bccausc Ihc
moIion!cssmicrooranisms IhaI composcdiI hard!y chancdin Ihcir
anaIomy.' BuI conIrary Io whaI a supcrhcia! !ook aI Ihcir cxIcrna!
appcaranccmaysucsIIhoscancicnIoranismshadmanacdIodis-
covcrovcrIhaIpcriodo!Iimc a!!Ihcbiochcmica!proccsscs o!cncry
cxIracIionIhaI cxisIIoday.A!IcrIhaI, cvo!uIionproduccdmanyncw
mcchanismsIo usc cncryIopcr!ormwork, !orcomp!cx!ocomoIion
or!orncura!conIro!, !orinsIancc,buI nc nwma]crwaysc]x!racrinj
nrjy ]rcm jradinrs.' Kouh!y, Ihc car!icsI bacIcria appcarcd on
Ihis p!ancI Ihrcc and a ha!! bi!!ion ycars ao scavcninIhc producIs
o!non-bio!oica!chcmica!proccsscs:abi!!ionycars!aIcrIhcycvo!vcd
IhccapaciIyIoIapinIoIhcso!arradicnI,producinoxycnasaIoxic
byproducI: and onc bi!!ionycarsa!IcrIhaIIhcy cvo!vcdIhc capaciIy
Io usc oxycnIorcaI!yincrcascIhc c!hcicncyo!cncryandmaIcria!
consumpon. ByconIrasI,IhcrcaIdivcrsiIyo!mu!Iicc!!u!aroransms
IhaI popu!aIc Ihc p!ancI Ioday was cncraIcd in abouI six hundrcd
mi!!ion ycars. ThusIhc hisIory o!Ihc car!icsIbiosphcrc is anarraIivc
o!howIhc capaciIy o!Ihc cnvironmcnI Io susIain !i!c was cxIcndcd
byIhcdiscovcryIhrouhcvo!uIionary scarch o!Ihcbiochcmica!cir-
cuiIrynccdcd!or]rmnraricn,rhcrcsynrhs, andrsriraricn.
b

N LbN N blMUL1N
Toivcanicao!Ihcincrcascdc!hcicncyrcprcscnIcdbyIhcscmcI-
abo!ic !andmarks wccanusc somc numbcrs obIaincd !rom sIudyin
conIcmporary microoranisms. Usin Ihc !casI c!hcicnI proccss, !cr-
mcnIaIion, I 80 rams o! suar can bc brokcn down Io yic!d 20.000
ca!orics o!cncry. Tc suar uscdasrawmaIcria!!orIhis chcmica!
rcacIion was oriina!!y Iakcn rcady-madc !rom Ihc cnvironmcnI by
Ihc car!icsI bacIcria buI wiIh Ihc discovcry o! phoIosynIhcsis Ihosc
oranisms cou!d now producc iI usin 264 rams o! carbon dioxidc,
I 08rams o!waIcr, and 700.000 ca!orics Iakcn !rom sun!ihI, Ihcy
cou!dproduccIhc samc I 80ramso!suar,p!us I 2ramso!oxycn
as wasIc producI. WiIhIhc advcnIo!rcspiraIion, inIurn, IhaIwasIc
producI cou!d bc uscd Io burn Ihc I 80 rams o! suar Io producc
700.000ca!oricso!cncry.Thus, add\nphoIosynIhcsisIo!crmcnIa-
Iion madc Ihc rowIh o! Ihc car!icsI popu!aIions o! !ivin crcaIurcs
sc!!-susIainin, whi!c addin rcspiraIion produccd a ncI surp!us o!
bacIcria!t!csh [or biomass" ) .GivcnIhaIsurp!usiIbccamcpossib!cIo
o!rommcrcpopu!aIionrowIhIoincrcascdspccicsdivcrsiIyIhrouh
Ihc comp!cxihcaIion o!!ood chains.U oIhcrwords, bacIcria!biomass
iIsc!! bccamc a radicnIIhaI cou!dbc Iappcd inIobyncw!y cvo!vcd
prcdaIory spccics, Ihc anccsIors o!conIcmporary unicc!!u!ar oran-
isms!ikcparamcciaoramocbac.
WhcIhcr prcdaIory or noI, morc comp!cx !ivin crcaIurcs didnoI
havcIocon!ronIIhcsamc mcIabo!ic prob!cmsIhaIhadIakcnbacIcria
bi!!ions o! ycars Io so!vc. KaIhcr, ancicnI anima!s incorporaIcd as
a who!c microoranisms IhaI had a!rcadymasIcrcd rcspiraIion [Ihc
anccsIors o!miIochondria) whi!c ancicnIp!anIs absorbcdIhosc IhaI
had acquircd Ihc capaciIy !orphoIosynIhcsis [ ch!orop!asIs) . U oIhcr
words, Ihc p!anIs and anima!s IhaI wou!d cvcnIua!!y bccomc mu!Ii-
cc!!u!ar camc inIo bcin by usin bacIcria and Ihcir mcIabo!ic cir-
cuiIryasbui!d\nb!ocks.ThcinIcracIionsbcIwccnIhcncwcrcaIurcs
and Ihc o!dcr bacIcria may havc sIarIcd as parasiIic and on!y !aIcr
dcvc!opcdinIoamorcmuIua!isIicrc!aIionasIhcinIimaIcknow!cdc
IhaIparasiIcshadabuIIhcirhosIswasuscd!orIhcircommonbcnchI.
Thc crucia!c!cmcnI in Ihc rc!aIion was simulrancus rrlicaricn. Ihc
!ormcrparasiIcshadIo bcrcp!icaIcdinsynchronywiIhIhcirhosIsso
IhaIIhcirrcproducIivc inIcrcsIsdidnoIdivcrc. `UmanyconIcmpo-
raryoranismsIhcrcarc cndurinsymbioscs inwhichbacIcriainIhc
uIs o! !arcr anima!s a!!ow Ihc !aIIcr Io dicsI !ood Ihcy cou!d noI
LNL1L LK1NMb N NLLN1 KNlbMb
oIhcrwiscproccss,buIIhcmicrooranismsmusIbcrc-incsIcdcvcry
cncraIion.ThcdccrcascdinIimacyIhisimp!icsmcansIhaImuIaIions
may arisc IhaI swiIch Ihc symbiosis back inIo parasiIism. UIhc casc
o!ancicnIan\ma!sandIhcirmiIochondria, orancicnIp!anIsandIhcir
ch!orop!asIs,onIhcoIhcrhand,anyrcIaincdmuIaIioninIhcabsorbcd
bacIcria had Io a!so bc bcnchcia! Io Ihc hosI. This dccpcr !orm o!
muIua!isIicrc!aIionis ca!!cdndcsymbics.
ndosymbiosisisancxoIici!!usIraIiono!IhcparI-Io-who!crc!aIion
incco!oy.Thc morc ordinarycxamp!cs!ormscvcra!!cvc!so!orani-
zaIioncachonc sIudicdbyiIsownsub-discip!inc.bchaviora!, popu!a-
Iion. communiIy, and ccosysIcm cco!oy. AI Ihc sma!!csI sca!c Ihc
ob|ccI o!sIudyis Ihc bchavior o! individua! oranismsin Ihcircnvi-
ronmcnI. moIi!c bacIcria scnsin nuIricnI radicnIs in Ihcir viciniIy
andmovininIhc dirccIion o!hihcr conccnIraIion, !or cxamp!c, or
anccsIra!amocbac usin Bowin proIop!asmic cxIcnsions Io cncirc!c
bacIcria!prcyprior UincsIin iI.ThcncxI sca!c !ocuscsonIhcpopu-
!aIion o!whichIhcsc oranisms arccomponcnIparIsandsIudicsIhc
wayiIrows!imiIcdon!ybyIhc capaciIy o!!oca!radicnIs Io susIain
iI.cIai!sabouIIhcbchavioro!individua!oranismsbccomc causa!!y
rcdundanI B an cxp!anaIion o! Ihis rowIh. AI Ihc ncxI sca!c cnIirc
communiIics composcdo!scvcra!inIcracIinpopu!aIionso!di!!crcnI
spccisbccom

Ihc IarcI o! ana!ysis. Andhna!!y, aIIhc !arcsI sca!c,

Ihc inIcracIions bcIwccn scvcra! o! Ihcsc communiIics arc sIudicd,
disrcardin Ihc disIincIionsbcIwccn spccics and dca!inon!y wiIh
Ihc !uncIion Ihcy pcr!orm, asproduccrsor consumcrs, in Ihc ovcra!!
t!ow o! maIIcr and cncry in an ccosysIcm. A!Ihouh cach o! Ihcsc
spaIia!sca!cs is disIincI cnouhIo bc sIudicdby a di!!crcnI hc!diI is
imporIanI Io cmphasizc IhaI wc arc dca!in hcrc wiIh di!!crcnccs
in rlarivscal, IhaI is, wiIh sca!c as is cncraIcd by Ihc rc!aIion o!
parI-Io-who!c, noI wiIh abso!uIc sca!c. a sin!c conIcmporary!arc
p!anI,!orcxamp!c,mayhousc cnIircccosysIcmo!microoranisms
disp!ayina!!!our !cvc!s o! oranizaIion.
A ood sIarIin poinI !or a phi!osophica! discussion o! cco!oica!
issucs is Ihc sca!c sIudicd by popu!aIion cco!oy, a !cvc! o! oraniza-
Iion characIcrizcd by cmcrcnI propcrIics !c Ihc paIIcrn o! rowIh
o!apopu!aIion.This paIIcrnisproduccdbyIwodi!!crcnI!acIors. On
onc hand, Ihcrc is Ihc rowIh raIc o! Ihc oranisms, basica!!y Ihcir
birIh raIc minus Ihcir dcaIh raIc, a!Ihouh raIcs o! cmiraIion and

d
NlLbN N blMUL1N
immiraIiona!soa!!ccIIhcrapidiIyors!owncsso!popu!aIionrowIh.
OnIhc oIhcrhandIhcrc isIhcavai!ab!c spacc, maIIcr, and cncryin
Ihc !ocaIion inhabiIcd by Ihc popu!aIion. Thcsc Iwo !acIors IocIhcr
dIcrminc Ihc carrinj caracin. Ihc maximum popu!aIion IhaI can
bc susIaincdindchniIc!ywiIhouIdcp!cIinavai!ab!c rcsourccs. Sincc
oncandIhcsamccnvironmcnImayhousc manydi!!crcnIspccicsIhc
carryin capaciIy varics dcpcndin on Ihc cncrcIic, maIcria!, and
spaIia! rcquircmcnIs o!cachspccics. apopu!aIionIhaIcansurvivc on
sma!!amounIso!cncry, !orcxamp!c,wi!IrowIo amuch!arcrsizc
Ihan onc whosc cncry nccds arc hihcr. Bccausc rowIh paIIcrns
maybc a!!ccIcdby ovcrcrowdiniIisprc!crab!c Iomcasurc Ihc ouI-
comc o! rowIhnoI B abso!uIc numbcrs buI in dcnsiIics, IhaI is, by
Ihc avcrac popu!aIion sizc in a ivcn amounI o! spacc. Carryin
capaciIy canIhcnbc IrcaIcd as asinu!ariIyin Ihc spacc o!possibi!i-
Iics !orIhc inIcracIionsbcIwccnpopu!aIion dcnsiIics and rcsourccs,
Ihcspccia!va!uc o!dcnsiIyIhaIcnsurcsapopu!aIionwi!Ibc aI!on-
Icrmcqui!ibriumwiIhiIscnvironmcnI.
Kca! popu!aIions, o! coursc, nccd noI cxisI aI Ihc sinu!ariIy buI
t!ucIuaIcaroundiIorpcriodica!!y ovcrshooIiI.Howarca!popu!aIion
wi!! bchavc rc!aIivc Io Ihc sinu!ariIy dcpcnds on which o! Ihc Iwo
!acIors IhaI dcIcrmincs carryin capaciIy dominaIcs iIs rcpoducIivc
sIraIcy. A sIraIcy bascd on rowIh raIc imp!ics IhaI a spccics wi!!
IcndIoproducc many o!!sprinquick!y, o!!sprinIhaIwi!!IcndIobc
sma!! and shorI!ivcd. AsIraIcyIarcIcdaIcxisIinrcsourccs, onIhc
oIhcr hand, wi!I producc !cwcr, !arcr, !oncr !ivcd o!!sprin morc
ab!c Io!cam Io conIro!IhcircnvironmcnI, morc cncry c!hcicnI, and
morc capab!co!hnc-IuniniIsnumbcrsIocnvironmcnIa!condiIions.
InsccIs and wccds arc o!Icn uscd as cxamp!cs o! Ihc hrsI sIraIcy
whi!c !arc mamma!s and Irccs cxcmp!i!yIhc sccond onc. GivcnIhc
sma!! sizcandrapidrcproucIion o!microoranisms onc wou!dIhink
IhaI Ihcy shou!d a!ways usc Ihc hrsI sIraIcy buI sIudics o!microbia!
cco!oics havc shown IhaI boIh sIraIcics cxisI. somc popu!aIions
inorc carryin capaciIy, ovcrshooI iI, Ihcn crash-pcrhaps oin
dormanI as spocs or cysIs Io waiI !or ncw rcsourccs-whi!c oIhcrs
row !asIaIhrsI buI Ihcnshow dcnsiIy-dcpcndcnI saIuraIion c!!ccIs
and cnI!y convcrc on Ihc sinu!ariIy. In addiIion, chancs B Ihc
avai!abi!iIy o! rcsourccs can conIribuIc Io shapin rowIh paIIcms,
as whcn scasona! variaIions a!!ccI rcsourcc abundanccsmakinIhc
LNL1L LK1NMb N NLLN1 KNbMb
dcnsiIy o! apopu!aIionvaryina pcriodicway.Thcsc cyc!icpaIIcms
can ccomc cvcn morc comp!cx whcn wc movc Io Ihc !cvc! o!
communiIycco!oybccausc aIIhaIpoinIIhcdcnsiIyo!aprcyspccics
bccomcs Ihc main !acIor dcIcrminin Ihc carryin capaciIy !or a
prcdaIory spccics.
UsindcnsiIyinsIcado!abso!uIcnumbcrsIocharacIcrizcapopu!a-
IionmcansIhaIwc can Ihink o!Ihc!aIIcrinIcrms o! radicnIs. !ikc
IcmpcraIurc orprcssurc dcnsiIy is an inIcnsivc propcrIy. InIhis casc,
Ihc radicnI is a conccnIraIion o!biomass!romwhichcncrycanbc
cxIracIcdbyoIhrspccics.co!oica!rc!aIionscan,in!acI,bcdchncd
by Ihc c!!ccIIhaIIhc dcnsiIy o! onc spccics has on Ihc dcnsiIy o!
anoIhcr. I! an incrcasc inIhcdcnsiIy o! oncspccicsdccrcascsIhaI o!
anoIhcrIhcnIhc rc!aIionisciIhcrprcdaIoryorparasiIic. U Ihc dcnsi-
Iicso!boIhspccicsdccrcascIhcrciscompcIiIion.I!IhcyboIhincrcasc
Ihc rc!aIion is onc o! muIua!ism or symbiosis. Ad hna!!y, u Ihc
dcnsiIy o! onc popu!aIion rows wiIhouI a!!ccIin IhaI o! Ihc oIhcr
Ihcn Ihcir rc!aIion b onc o! commcnsa!ism. To sIudy Ihcsc rc!aIions
communiIy cco!oy musI add Io Ihc mcchanisms dcIcrminin Ihc
rowIh raIc o! a popu!aIion a raIc coup!in mcchanism, IhaI is, a
cndurininIcracIionmakinIhc raIcs o!rowIh o! Ihc Iwo popu!a-
Iions dcpcndoncachoIhcr.'IcIsuscIhcinIcracIionbcIwccnprcda-
Iorsandprcy as ancxamp!c. InsIandardmodc!sIhc prcypopu!aIion
b assumcdIo havc acccss o un!imiIcdrcsourccs soIhcprimarypro-
duccrs arc noIcxp!iciI!y modc!cd. InIhc casc o!ancicnI !ood chains
Ihc prcy popu!aIion can bc imaincd Io bc bacIcria usin !crmcnIa-
Iion Io consumc Ihc oranic producIs o! phoIosynIhcIic produccrs.
In Ihc abscncc o! prcdaIors such abacIcria!popu!aIionwou!d row
cxponcnIia!!ybuIinIhcirprcscncciIsdcnsiIywi!!dcpcndonIhcraIc
o!prcdaIion. Thc dcnsiIyo!Ihc prcdaIorpopu!aIion, inIun, dcpcnds
oniIsown rowIhraIc as wc!!asonIhc raIc o!prcy capIurc. Fina!!y,
Ihc dcnsiIics o! cach popu!aIion a!!ccI Ihc probabi!iIy IhaI Ihcy wi!!
cncounIcrcach oIhcr and inIcracI, Ihc hihcrIhc dcnsiIics Ihc morc
!ikc!yIhc inIcracIion.
In suchascnarioiIisrc!aIivc!ycasyIo obIainpopu!aIiondcnsiIics
IhaI do noI scII!c inIo a sIcady sIaIc buI insIcad cyc!c rcpcaIcdJy. as
IhcprcdaIorpopu!aIionrowsiIrcduccsIhcpopu!aIiono!prcyupIo
Ihc poinI whcrc Ihcrc is noI cnouh prcy Io hunI: Ihis makcs Ihc
prcdaIor popu!aIion crash a!!owin Ihc prcy popu!aIion Io rccovcr
V

N LbN N b MUL1N
andsIarIIhc cyc!c aain. OnIhc oIhcrhand, a rhyIhmicpaIIcrn o!
boom and busIin Ihc rcspccIivc dcnsiIics docs noI imp!y IhaI Ihc
spacc o! possibi!iIics !orIhc coup!cd sysIcm prcdaIor-prcy conIains a
cyc!icsinu!ariIy [a pcriodicaIIracIor) .Thiswi!!bcIhccascon!yi!Ihc
osci!!aIion in Ihc dcnsiIics is asympIoIica!!y sIab!c, IhaI is, on!y i!
Ihc pcriod o! Ihc osci!!aIion has a Icndcncy Io rcIurn Io iIs oriina!
va!uc a!Icr an cxIcma! shock. NaIhcmaIica! modc!s wiIh pcriodic
aIIracIors can bc cncraIcd u Io Ihc prcvious modc! wc add Ihc
assumpIion IhaI prcdaIors can bccomc saIiaIcd, IhaI is, IhaI as Ihc
prcy popu!aIion bccomcs morc abundanI raIcs o! prcdaIion s!ow
down bccausc prcdaIors arc a!rcady a!! Ihc prcy Ihcy nccd.
aIa!rom acIua!popu!aIionsinIhcirnaIura!habiIaIs, aswc!!as!rom
popu!aIions sIudicdundcr conIro!!cd!aboraIory condiIions, Icnds Io
conhrm IhaI sIab!c cyc!cs in dcnsiIy arc indccd common. U somc
cascs cvcn morc comp!cxaIIracIors, cyc!cs oncyc!cs or cvcn dcIcr-
minisIicchaos,havc bccndocumcnIcd. "Thus,IhcinIcrp!ay o!!orma!
ana!ysisand!aboraIory cxpcrimcnIaIionis ivin communiIy cco!o-
isIs Ihc mcans Io cxp!orc Ihc mcchanisms and Ihc mcchanism-
indcpcndcnIsIrucIurco!Ihcirsub|ccImaIIcr.ToIhisiImusIbc addcd
Ihcro!c IhaI simu!aIions canp!ay as inIcrmcdiaricsbcIwccnIhcory
and cxpcrimcnI a!!owin us Io !o!!ow popu!aIion dcnsiIics as Ihcy
movc Ioward asinu!ariIy.
Top!ayIhisro!c a simu!aIionmusI cmbody iIs simu!aIcd oranisms
by ivinIhcm a mcIabo!ism andsiIuaIc Ihcm in a spacc wiIha ccr-
Iain disIribuIion o! rcsourccs. Un!ikc Ihc uscs o! cncIic a!oriIhms
discusscd in Ihc prcvious chapIcr, in which discmbodicd rcp!icaIors
wcrc capab!c o! hndin a sin!c opIima! so!uIion, cmbodicd and
siIuaIcd crcaIurcs Icnd Io hnd a ranc o! acccpIab!c so!uIions ivcn
IhaI rcsourccs can vary rcaI!y in Ihcir spaIia! disIribuIion and IhaI
Ihcso!uIions!oundbycvo!uIionaryscarchnccdnoIbc opIima!!orIhc
cnIirc popu!aIion. U addiIion, hIncss cva!uaIions musI noI bc pcr-
!ormcd !rom Ihc ouIsidc buI nccd Io bc ndcjncus. rcproducIivc
succcss musIdcpcndonIhcabi!iIyo!Ihcsimu!aIcdoranismsIomccI
Ihcir mcIabo!ic rcquircmcnIs. Whcrcas wiIh cxocnous hIncss an
cvo!uIionaryscarchhndsso!uIions Io prob!cms poscdbyIhc cxpcri-
mcnIcr in Ihcsc simu!aIions Ihc prob!cms arc poscdby Ihc cnviron-
mcnIin Ihc casc o!popu!aIion cco!oy, orby oIhcroranisms in Ihc
casc o!communiIycco!oy. IcIsbcinwiIhIhchrsIcasc.simu!aIions
LNL1L LK1NMb N NLLN1 KNbMb
o! popu!aIion cco!oy in whch Ihc IarcI o!Ihc sIudy is cmcrcnI
rowIh paIIcms and hIncss is dchncd by carryin capaciIy. On
onc hand, Ihc rowIhmay Iakc p!acc bccausc Ihc cnvironmcnIhas
bccn cnrichcdwiIhncwradicnIs, an in|ccIion o!mincra!rcsourccs
causcd by an cxIcma! cvcnI !ikc a vo!canic crupIion, !or cxamp!c.
OnIhc oIhcr, Ihc rowIhmaybc causcd byIhc cvo!uIionardiscov-
cry o!novc! mcIabo!ic sIraIcics, !ikc phoIosynIhcsis or rcspiraIion.
ThaI is, wc wanI Io modc! Ihc capaciIy o! Ihc cnvironmcnIIo a!!ccI
Ihc oranisms as wc!! as Ihc capaciIy o!Ihc oranismsIo a!!ccI Ihcir
cnvironmcnI.
U onc approach, ca!!cd IaIcnI ncry nvironmcnIs" [I) , a
spaIia!dimcnsionb addcdIocncIica!oriIhmsinIwodi!!crcnIways.
Ihc simu!aIcd oranisms inhabiI an cxIcma! spacc wiIh a ccrIain
disIribuIion o! rcsourccs and Ihcy havc an inIcma! spacc [oruI")
inwhich Ihcy cancarry rcsourccs a!Icr incsIin Ihcm. U addiIion,
un!ikc sIandard cncIic a!oriIhms in which ancw cncraIion com-
p!cIc!y rcp!accs Ihc o!dcr onc and in which popu!aIion numbcrs
rcmainhxcd, inIdi!!crcnIcncraIionscan cocxisIandpopu!aIion
sizc can vary, a!!owin Ihc cxp!oraIion o! dcnsiIydcpcndcnI c!!ccIs
onrowIhpaIIcrns. BuIIhc mosIimporIanI!caIurc o!IisIhcway
inwhichiIcnab!csusIodisIinuishIhc c!!ccIso!rcsourccavai!abi!iIy
!rom Ihosc o! cvo!uIionary adapIaIions inIhc !aIc o!a ivcnpopu!a-
Iion. To Ihis cnd I docs noI disIribuIc rcsourccs as rcady-madc
!oodbuI raIhcrasbiIs andpicccs IhaImusIbc combincdinspccihc
ways!orIhcir!aIcnIcncry"Iobccomc usc!u!cncry."Thcpossib!c
mcIabo!ic combinaIions o! Ihcsc biI and picccs arc prcdchncd in a
Iab!c,IhaIis,IhcyarcnoIcmcrcnI,buIIhcyarcrichcnouhIoyic!d
noI on!y di!!crcnI amounIs o! cncry buI a!so byproducIs IhaI can
cnIcr inIo !urIhcr combinaIions. Thus, an cxpcrimcnIcr can conIro!
Ihcprob!cmIhaIIhc cnvironmcnIposcsIoanoranisminIwoways.
bymakinIhcpossib!cmcIabo!iccombinaIionsmorcor!csscomp!cx
[ chanin Ihc cnIrics in Ihc Iab!c) orby dcIcrminin how spaIia!!y
conccnIraIcdordispcrscdIhc !ood componcnIs arc. SinccIhcIab!c o!
rcacIionsisnoIcvo!vab!c andIhc crcaIurcs uIisasimp!cconIaincr
Ihc so!uIionsIhcymusI discovcr arc noIncwmcIabo!icproccsscsbuI
ncw bchaviors. Norc spccihca!!y, Ihcy musI cvo!vc novc! ]crajinj
srrarjis. Scarchin !or !oodin an c!hcicnI way is imporIanIbccausc
movcmcnI invo!vcs an cncry cosI. This mcans IhaI Ihc simu!aIcd
I
Z
NLbN N bMUL1N
oranisms havc an inccnIivc Io discovcr bchaviora! sIraIcics IhaI
minimizc Ihc amounI o!work nccdcdIo hnd Ihc rihI combinaIion
o! !oodbiIsIhaIwi!! yic!d a!u!!mcIabo!icrcacIon.
A!IhouhI canbcuscdindi!!crcnIways!orIhcprcscnIdiscus-
sionwcshou!dIhink o!Ihcsc!orainsIraIcicsas discovcrcdbyan
cnIirc popu!aIion ovcr many cncraIions. Wc can Iakc Ihc mobi!c
oranisms in I Io rcprcscnI unicc!!u!ar prcdaIors-Ihc di!!crcn
c!asscs o! which arc disIinuishcd by Ihcir !ocomoIivc machincry
usin ci!ia, tIac!!a, orpscudopodia-whi!c Ihc sIaIic !ood biIs rcprc-
scnI di!!crcnI Iypcs o! immobi!c bacIcria, cach providin a prcdaIor
wiIhdi!!crcnInuIricnIs.IikcancicnIprcdaIorsIcrcaIurcshavcan
oricnIcdbodydchncdbyIhc!ocaIiono!simu!aIcdscnsorsandmoIors.
Thcscnsorsranc!romcxIrcmc!ysimp!c,scnsindirccIconIacIwiIh
a!oodparIic!cinIhcsamc!ocaIion, IomorcsophisIicaIcdoncs a!!ow-
in scnsin aI a disIancc. Thc crcaIurcs a!so havc uI scnsors" Io bc
ab!cIoIc!!whaIIhc currcnIinIcrna! conIcnIs arcandmakc !orain
dccisions onIhc basis o!whaI c!sc is nccdcdIo comp!cIc a mcIabo!ic
rcacIion. Thcsc scnsors corrcspond Io Ihc ro!c IhaI proIcins [di!!uscd
inIhccyIop!asmorcmbcddcdonIhcmcmbranc)p!ayinmicrooran-
isms. Thc componcnI o! Ihc simu!aIcd oranisms IhaI Irans!orms
in!ormaIion !rom Ihc scnsors inIo di!!crcnI Iypcs o! movcmcnI is a
simp!c modc! o! a ncuron ca!!cd a ncura! ncI." This aspccI o! Ihc
imp!cmcnIaIion has advanIacs and disadvanIacs !rom Ihc prcscnI
poinIo!vicw. IhcadvanIacisIhaIncura!ncIs cancasi!ybccoup!cd
Io cncIic a!oriIhms, as wc wi!! discuss in Ihc !o!!owin chapIcr,
a!!owininhcriIcd bchaviorIocvo!vc ovcrmanycncraIions:Ihcdis-
advanIac is IhaIancicnIprcdaIors did noI usc ncurons Io connccI
scnsors and moIors buI comp!cxbiochcmica! mcchanisms. BuI sincc
wc arc inIcrcsIcd in cxp!orin mcchanism-indcpcndcnI qucsIions
Ihis way o!mappin scnsory inpuIs inIo moIor ouIpuIs nccd noI bc
aprob!cm.
Whcn I is uscd Io simu!aIc siIuaIions B popu!aIion cco!oy
whaI wc wou!d !ikc Io scc cmcrinis Ihc corrccI rowIhpaIIcrns.
Iikc ancicnI prdaIors Is crcaIurcs arc dcsincd Io cvo!vc morc
c!hcicnI !orain sIraIcics sucsIin a rowIh paIIcm in which
dcnsiIy is hnc-IuncdIo carryin capaciIy. ThaI is, Ihc rowIhpaIIcrn
o! Ihc popu!aIion shou!d noI pcriodica!!y ovcrshooI Ihc carryin
capaciIy and Ihcn crash buI approach iI asympIoIica!!y. And IhaI is,
LNL1L LK1NMb N NLLN1 KNbMb
indccd,IhcpaIIcrnobscrvcdinacIua!simu!aIions. UaddiIion, whcn
a novc! cvo!vcd bchavior cxIcnds carryin capaciIy Ihcrc wi!! bc a
pcriod o!Iimc inwhichIhc o!dcqui!ibriumis rcp!accdbyancw onc
and maIhcmaIica! modc!s havc !iII!c Io Ic!! us abouI Ihc dynamics
durin Ihc IransiIion. I on Ihc oIhcr hand can Irack popu!aIion
chancsasIhcyad|usIIoncwcondiIions. Fina!!y,havincsIab!ishcd
iIs compaIibi!iIywiIhIradiIiona!modc!sI canscrvc as amcansIo
conducI cxpcrimcnIs o!di!!crcnIkinds. Ihc Iab!c o! mcIabo!ic rcac-
Iions, !or cxamp!c, canbcchancdIo rct!ccIIhc scasona! avai!abi!iIy
o! di!!crcnI rcsourccs by makiD Ihc producIs o! di!!crcnI combina-
Iions varypcriodica!!y: orIhc spaIia! disIribuIion o!rcsourccs can bc
a!Icrcd !rom rc!aIivc!y uni!orm Io onc in which di!!crcnI biIs and
picccs o! !ood arc !ocaIcdindi!!crcnI p!accs, makiDIhc so!uIion Io
Ihc surviva! prob!cm hardcr. This wayI can IcsI whcIhcrin cnv-
ronmcnIs o!incrcascd comp!cxiIy cvo!uIionaryscarchis ab!c Io nd
so!uIionsIhaIarcopIima!!orIhccnIircpopu!aIionorwhcIhcrdi!!cr-
cnI subpopu!aIions wi!! dcvc!op adapIaIions IhaI arc ood compro-
miscs IoIhcir!oca! condiIions. '

IcIs pausc !or a momcnI Io summarizc Ihc arumcnI so !ar. Thc

carIhs car!icsI biosphcrc consisIcd o! t!aI co!onics o! bacIcria wiIh
unchanin cxIcrna! anaIomics IhaI IcndcdIo obscurc Ihc momcn-
Ious chancs IhaI wcrc Iakinp!acc wiIhin Ihcir mcmbrancs. ncw
biochcmica!circuiIryIhaIbasica!!y cxhausIcda!!Ihcknownways o!
cxIracIin cncry !rom cnvironmcnIa! radicnIs. Thcsc mcIabo!ic
!andmarks cvcnIua!!y !cd Io a ncI surp!us in Ihc producIion o! bio-
mass and Io Ic subscqucnI cmcrcncc o! cnIirc!y ncw crcaIurcs
capab!c o! survivin by consumin IhaI biomass. From Ihcn on Ihc
inIcracIion bcIwccn popu!aIion dcnsiIy and avai!ab!c rcsourccs, or
IhaIbcIwccncoup!cdpopu!aIion dcnsiIics, bccamc Ihc mainsourcc
o!cco!oica!cmcrcnIpropcrIics. Thc mainqucsIioncon!ronIin us
nowisIhccvo!uIionaryconscqucnccso!Ihcsccco!oica!inIcracIions.
Norc spccihca!!y, whcn onc spccics prcys on anoIhcr iI bccomcs a
sc!ccIion prcssurc on iIs prcy, and vicc vcrsa, whcn a prcy spccics
cvo!vcs adapIaIions Io counIcracI prcdaIory abi!iIics iI bccomcs a
sc!ccIionprcssurc oniIsprcdaIors. ThismcansIhaIovcrmanycncra-
IionsIhcIwospccicscnacinanarmsracc"inwhichancwadapIa-
Iionby onc spccics sIimu!aIcs Ihc cvo!uIion o! a counIcr-adapIaIion
byIhcoIhcr.Andsimi!ar!y!orcco!oica!inIcracIions oIhcrIhanIhosc
J
7
NLbN N b MUL1N
bcIwccn prcdaIors and prcy. SymbioIic spccics, !or cxamp!c, may
comcIodcpcndoncachoIhcrsoc!osc!yIhaIIhcircvo!uIionbccomcs
IihI!y coup!cd, as i!!usIraIcd byIhcwayinwhich many conIcmpo-
raryp!anIshavcdcvc!opcdanob!iaIoryrc!aIionwiIhIhcinsccIsIhaI
po!!inaIc Ihcm. vo!uIionaryinIcracIions arc di!!crcnI!rom cco!oi-
ca!oncsnoIon!ybccauscIhc!aIIcrIakcp!acc inrc!aIivc!yshorIIimc
sca!cs-Ihc pcriods o! Ihc osci!IaIions in dcnsiIy, !or cxamp!c, arc
mcasurcdinycars-whi!c Ihc!ormcrIakcp!acc in much!oncrIimc
sca!cs, buI a!so bccausc Ihcy arc associaIcd wiIh di!!crcnI possibi!iIy
spaccs.
U Ihc prcvious chapIcr wc saw IhaI an cvo!uIionary possibi!iIy
spacc canbc conccpIua!izcdasahIncss!andscapc. a spacc conIainin
d possib!c cnc scqucnccs on Iop o! which a va!uaIion in Icrms o!
cxpccIcdrcproducIivc succcss has bccn supcrimposcd. Thc disIribu-
Iiono!sinu!ariIics [hIncss opIima) BIhisspacc dchncsIhc comp!cx-
iIyo!Ihc surviva!prob!cmIhaIhasIobcso!vcd. aspaccwiIhasin!c
!oba! opIimum surroundcd by arcas o! minimum hIncss is a Iouh
prob!cm [a nccd!c in a haysIack) whi!c onc wiIh many !oca! opIima
roupcd IocIhcr dchncs a rc!aIivc!y casy prob!cm. In Ihc casc o!
prcdaIors and Ihcir prcy [or o! any oIhcr cco!oica! rc!aIion) wc
nccd Io coup!c Iwo o! Ihcsc spaccs bccausc a!IhouhprcdaIor and
prcy spccics havc Ihcir own cnoIypcs Ihcy int!ucncc cach oIhcrs
hIncss.whcncvo!uIionaryscarchhappcnsIohndancwcombinaIion
o! cncs IhaI improvcs Ihc prcys capaciIy Io cvadc iIs prcdaIor, !or
cxamp!c, Ihis wi!! !owcr Ihc hIncss o! somc cxisIin prcdaIor cncs,
andconvcrsc!y, ancwcncscqucnccIhaIimprovcshunIinbchavior
wi!I!owcrIhc hIncsso!somccxisIinprcycncs. U oIhcrwords,Ihc
cocvo!uIion o!prcdaIorsandprcyimp!icsIhaIIhcirhIncss!andscapcs
arc consIanIIybcin dc!ormcdwiIh somc hi!!s" bcin Irans!ormcd
inIo va!!cys. "' xIcndinIhis !inc o! IhouhI Io cnIirc !ood chains
imp!ics IhaI many hIncss Iandscapcs may bccomc scria!!y coupIcd,
raisinIhc qucsIion o!how!on !ood chains can bc bc!orc Ihc cou-
p!cd possibi!iIy spaccs conIain sinu!ariIy disIribuIions IhaI makc
surviva!probIms impossib!c Io so!vc. Thcsc considcraIionshavc !cd
IhcorisIs Io cxpccIIhaI cvoIuIioniIscI!may !imiI IhcIcnIh o! !ood
chains Ioavoida dcadcnd, ahypoIhcsis madcpIausibIc by cmpirica!
hndins o!rcIaIivc!y shorI!ood chains in acIuaIccosysIcms.'
LNL1L LK1NMb N NLlLN1 KNlbMb
Whcn dcsinin simu!aIions Io sIudy cocvo!uIion havin conIro!
o! Ihc dcIai!s o! mcIabo!ic proccsscs or o! Ihc spaIia! disIribuIion o!
rcsourccs is noI as ncccssary as bcin ab!c Io dcp!oy cmbodicd crca-
Iurcs o! di!!crcnI spccics IhaI can rcconizc and inIcracI wiIh onc
anoIhcr. Uoncapproach, ca!!cdEchc.IhcinIcrna!mcIabo!icmachin-
cry o! Ihc simu!aIcd crcaIurcs is madc so simp!c IhaI Ihcy in c!!ccI
caI cncs. " In cho cvo!uIion is simu!aIcdIhrouh a cncIic a!o-
riIhminwhichIhc sIandardsymbo!sIrino! oncs and zcrocs acIin
as a chromosomc has bccn chancd Io onc usin a !arcr a!phabcI.
Thc crcaIurcs!ivc and movc in a simpc spacc conIainin rcsourccs
IhaI arc symbo!s !rom IhaI a!phabcI and rcproducIion Iakcs p!acc
on!ywhcnIhccrcaIurcshavcaccumu!aIcdcnouhsymbo!sIo crcaIc
a copy o! Ihcir chromosomcs. ConsIrucIin cncrcIic rcsourccs and
chromosomcs!romIhc samc rawmaIcria!s a!!ows cho Io disrcard
mcIabo!ic proccssin and inc!udc on!y sIorac in an inIcma! uI"
whi!c simu!Iancous!y makin hIncss cva!uaIions cndocnous. To
simu!aIc cco!oica!inIcracIions symbo!s IhaI arc B Ihc uIs o! oIhcr
crcaIurcscanbccapIurcdinIhc casc o!prcdaIionorcxchancdinIhc
casc o! symbiosis. chos crcaIurcs arc css cmbodicd Ihan Ihosc o!
LnoIon!ybccasco!IhcidcnIiIyo!cncryandcncsbuIa!soduc
Io Ihc !ack o! aIrans!ormaIion o!cnoIypc inIo phcnoIypc. whi!cin
L Ihc simu!aIcd chromosomc cncodcs!ora ncura! ncI, Ihc inpuIs
andouIpuIso!whichacIas scnsors andmoIors, in choIhc chromo-
somc acIsboIhasinhcriIab!cmaIcria!andasbodiIyIraiIs.
Norc spccihca!!y, Ihc chromosomcs symbo!s p!ay Ihc roIc o! rujs
IhaI arc visib!c" Io oIhcr crcaIurcs. In rca! microoranisms Ihcsc
Iaswou!dbc mcmbranc-boundproIcinsa!!owin!ormo!ccu!arrcc-
oniIionusina!ock-and-kcymcchanism, orcvcnchcmica!byprod-
ucIs o!an oranisms mcIabo!ism IhrouhwhichiI canbc idcnIihcd
byoIhcrs,buIIhcscdcIai!sarcinorcdin cho. TasworknoIon!yas
a mcans o!idcnIihcaIionbuIa!so as Ihc mcans Io spcci!y Ihc capaci-
Iics o!acrcaIurcIoa!!ccIandbc a!!ccIcdby oIhcrcrcaIurcs.Thishas
Ihc conscqucnccIhaIchos crcaIurcsdonoIinIcracIwiIhcachoIhcr
dirccI!y buI nccd an cxIcrna! proram Io comparc Ihcir Ias nd
dcIcrmincIhc ouIcomc o!an inIcracIion."ThcIasarco!IwoIypcs.
o!!cnsivcanddc!cnsivc.ThcbasicproccdurcisIrandom!ypickpairs
o!crcaIurcsinhabiIinIhc samcsiIc andIhcncomparcIhc o!!cnscIa
b

N LbN N bMUL1N
o!onIoIhcdc!cnscIao!IhcoIhcr, andviccvcrsa.Thccomparison
aims onIy aI csIabIishin Ihc numbcr o! symboIs o! cach Ia IhaI
maIchIhosc o!anoIhcr Ia aIIhc samc posiIion inIhc chromosomc.
I!IhcmaIchisasymmcIric-Ihc o!!cnscIao!onccrcaIurcmaIchin
Ihc dc!cnsc Ia o! anoIhcr buI noI Ihc oIhcr way around-Ihc hrsI
crcaIurc is Iakcn Io bc a prcdaIor andIhc sccond onciIs prcy. I!Ihc
maIch is symmcIric, IhaI is, i! boIh Ihc o!!cnsc and dc!cnsc Ias o!
cach crcaIurc maIch cach oIhcr, Ihcn Ihc Iwo arc considcrcd symbi-
onIs. Thc numbcr o! maIchcs dcIcrmincs Ihc amounI o! rcsourccs
IhaIarccapIurcdbyIhcprcdaIor [iI can caIIhccnIircprcyorsim-
pIyIakca biIc !romiI) orIhc amounI o!rcsourccsIhaIarcmuIuaIIy
cxchancd in symbiosis. This aIIows !or compIcx ccoIoicaI rcIaIions
Io cmcrc. onc spccics mayprcy n anoIhcr IhaI is symbioIicwiIh a
IhirdoncIhaI,inIurn, prcys onIhc hrsI.' `
UamorccompIcx vcrsiono!choIhc chromosomc is dividcdinIo
Iwo parIs, onc acIin as Ias Ihc oIhcr pcr!ormin Ihc roIc o! condi-
Iion-acIion ruIcs, IhaI is, ruIcs spcci!yin IhaI i! a ivcn condiIion
obIains aparIicuIar acIionis Iakcn. U IhismorccompIcx vcrsionIhc
inIcracIions arc sIiIInoI pcr!ormcd by Ihc crcaIurcs buI Ihc Ias arc
comparcdIhrouh Ihc condiIion-acIion ruIcs aIIowiD !orouIcomc
IhaI arc morc tIcxibIc. Ihc crcaIurcs can rc!usc Io inIcracI i! Ihc
condiIionsspccihcdinIhcirruIcsarcnoImcI, !orcxampIc. 'ThcrcIa-
IivcIack o!cmbodimcnIo!chos crcaIurcs rctIccIs Ihc cmphasis o!
IhisapproachonIhcIon-Icrm ouIcomcs o!inIcracIions[ sIrucIurc o!
!oodchains, rcIaIivc spccics abundanccs, arms raccs) raIhcrIhanIhc
inIcracIionsIhcmscIvcs. AsinIIhchrsIIaskIo bcpcr!ormcdisIo
caIibraIc Ihcsc cmcrcnI ouIcomcs Io Ihosc o! ciIhcrmaIhcmaIicaI
modcIs ordaIa!romcommuniIyccoIoy. ForinsIancc, inmanycascs
Ihc rcIaIivc abundancc o! di!!crcnI spccics has a disIincI sIaIisIicaI
shapc.manyspccicshavca!cwrcprcscnIaIivcswhIca!cwhavcmany
rcprcscnIaIivcs. xpcrimcnIs wiIh cho dcsincd Io IcsI whcIhcr iI
canproducc Ihcsc abundanccpaIIcrns havc bccn conducIcd andIhc
iniIiaI rcsuIIs arc ccourain. Thc cxpcrimcnIs aIIcmpIcdIo maIch
noIonIy a IarI sIaIisIicaI disIribuIion buI aIso Iricd Io answcr Ihc
qucsIiono!whcIhcrIhcouIcomcwasproduccdbycocvoIuIion.ToIcsI
Ihis hypoIhcsis Iwo vcrsions o! cho wcrc ran !or cach cxpcrimcnI,
onc in which inIcracIions wcrc mcdiaIcd by Ias and condiIions
[hcncc sub| ccI Io cocvoIuIion) and anoIhcr in which inIcracIions
LNL1L LK1NMb N NLLN1 KNbMb
IookpIaccrandomIy. ThcIa-mcdiaIcdsimuIaIionhIIhcdaIabcIIcr
IhanIhc purcIyrandomonc
FromIhc poinI o! vicw o! simuIaIin Ihc ancicnI biosphcrc cho
providcs a mcans Io cxpIorc Ihc vcry di!!crcnI ways in which Ihc
dcsccndanIs o!Ihc carIicsI microoranisms cvoIvcd. Thcrc wcrc Iwo
disIincIcvoIuIionarydynamicsdcpcndinonwhcIhcrornoIamicro-
oranism posscsscd an inncr mcmbranc cncapsuIaIin iIs cncIic
maIcriaIswiIhin anucIcus.Iackinsuch anucIcarmcmbrancaIIows
Ihc !rcc sharin o! cncs by cnIircIy di!!crcnI oranisms. II is wcII
known, !or cxampIc, IhaI Irans!crabIc scIs o! cncs caIIcd pIasmids
arcrcsponsibIc!orIhcsprcado!anIibioIicrcsisIanccinbacIcriainIhc
pasI 70 ycars, buI Ihis cncIicpromiscuiIy is noIhin ncw. AncicnI
bacIcria may havc cnacd in Ihis !orm o! horizonIaI cnc Irans!cr
!romIhc sIarI usinpIasmids andoIhcrmovabIc cncs ascombinaIo-
riaI buiIdin bIocks Io rapidIy acquirc capaciIics Ihcy prcviosIy
Iackcd.ThcancicnIprcdaIorso!IhoscbacIcria,onIhcoIhcrhand, did
posscss a scparaIc nucIcus and wcrc noI capabIc o! sharin cncs
IhaI way. Thcy compcnsaIcd !or IhaI by acquirin Ihc capaciIy !or
rcrrcducrivciscluricn. UIarcroranismsIhc!ormaIiono!ncwspccics
occurs IarcIyIhrouh Ihc isoIaIiono!Ihcir cnc pooI !rom tIows o!
cncIic maIcriaIs !rom oIhcr spccics. This mcchanism o! spcciaIion
[Ihc birIh o!a ncw spccics) is a ma|or cvoIuIionary !orcc promoIin
divcrcncc andanaIomicaIdi!!crcnIiaIion.Thc cvoIuIionary hisIorics
o! microoraniss wiIh and wiIhouI a nucIcus [ cukaryoIs and
prokaryoIcs) IcsIi!y Io Ihc imporIancc o! Ihis disIincIion. whiIc Ihc
!ormcrwcnIonIo!orm aIIIhcdi!!crcnIpIanIsandanimaIsIhaIcxisI
Ioday, IhcIaIIcr!oIIowcdnoIa divcrcnIbuIaconvcrcnIpaIh.ThaI
is, insIcad o! yicIdin a pIuraIiIy o! cnc pooIs morc or Icss isoIaIcd
!rom cach oIhcr Ihcy cncraIcd whaI is basicaIIy a ianI cnc pooI
spanninIhccnIircpIancI. '
U addiIionIoIhc prcscncc orabscncc o!spcciaIionIhc oIhcrma|or
dcIcrminano!carIycvoIuIionary ouIcomcs was cndosymbiosis, Ihc
aIrcady mcnIioncd absorpIion o! cnIirc microoranisms and Ihcir
mcIaboIic capaciIics. SimuIaIin cndosymbiosis andspcciaIioncanbc
donc in cho simpIy by addin ncw Ias or condiIion-acIion ruIcs.
ThchrsImodihcaIionisIoaddancwIamcdiaIinadhcsioninIcrac-
Iions and a mcans Io manac Ihc boundarics or mcmbrancs o! Ihc
inIcracIin crcaIurcs. U asimuIaIiono!ancicnImicrobiaIccosysIcms
77
d
NLbN N blMUL1lN
adhcsion Ias cou!d p!ay scvcra! ro!cs dcpcndin on Ihc ouIcomcs
o! Ihc inIcracIions and Ihc rcsu!Iin boundarics. A!!owin adhcsion
bcIwccnsimi!aroranisms can casi!y yic!d co!onics arrancdin !ay-
crs,much!ikcIhcmaI-!ikcsIrucIurcsIhaInon-moIi!cbacIcria!ormcd
in Ihc hrsIIwobi!!ionycars o! cvo!uIion. WiIhIhisspaIia!oraniza-
Iion Ihc cvo!uIion o!individua!crcaIurcs may bc int!ucnccd byIhc
cnvironmcnI providcdby simi!arcrcaIurcs on Ihc samc !aycr, orby
di!!crcnI crcaIurcs in a ncihborin !aycr, promoIin spccia!izaIion.
ndosymbiosis, on Ihc oIhcrhand, can bc simu!aIcd i!Ihc rcsu!I o!
adhcsioninIcracIionsisnoI Ihc crcaIiono!asin!cboundary around
many crcaIurcs buI Ihc absorpIion o! onc crcaIurc by anoIhcr in
whichIhc absorbcd crcaIurc kccps iIs ownboundary. '"
To simu!aIc spcciaIion Ihc obvious choicc wou!d bc Ioadda ncw
Ia Io simu!aIc cxprcssions o! spccics idcnIiIy This choicc wou!d
bc |usIihcd by Ihc !acI IhaI many oranisms achicvc rcproducIivc
iso!aIion Ihrouh Ihc posscssion o! spccia! markins [co!ors, odors,
pimcnIaIion paIIcms) . BuI in cho sc!ccIivc maIinis achicvcd by
addin a ncw coniIion-acIion ru!c IhaI uscs an cxisIin Ia [Ihc
o!!cnsc Ia) . Whcncvcr Iwo crcaIurcs occupyin Ihc samc siIc arc
rcady !or rcproducIion, IhaI is, whcn Ihcy havc incsIcd cnouh
symbo!s Io makc a copy o! Ihcir chromosomcs, Ihc condiIion parI
o! Ihcir maIin ru!csarc comparcd Io Ihcir o!!cnsc Ias and i! boIh
condiIionsarcsaIishcdcopicso!Ihcirchromosomcsarcmadc, crosscd
ovcr,andIwo ncwo!!sprin arc addcdIoIhcpopu!aIion.Thisprocc-
durc rcscmb!cs IhaIuscd by paramccia Ihc anccsIors o!whichwcrc
amon Ihc car!icsI cukaryoIcs. ByvaryinIhc spccihciIyo!Ihc maI-
incondiIion!owcrdcrccso!rcproducIivciso!aIioncanbcachicvcd
Io yic!dIhc cuiva!cnI o!prokaryoIcs.`Thc disIincIionbcIwccnIhc
Iypcs o! cvo!uIion IhaI rcsu!I wiIh and wiIhouI Ihc possibiiIy o!
spcciaIion musI bc kcpI in mind whcn IcsIin chos cvo!uIionary
ouIcomcs. OnccxpcrimcnI,!orcxamp!c, aIIcmpIcdIochcckwhcIhcr
chossimu!aIcdcvo!uIiondisp!aycdIhc samc opcn-cndcd characIcr
asrca!cvo!uIionor whcIhcriIrcmaincdboundcdwiIhin!imiIs.Thc
rcsu!Is !ai!cd Io maIch Ihc dcrcc o! divcrcncc IhaI is cvidcnI in
Ihc !ossi! rccord a!IcrIhc rcaI bursI o! di!!crcnIiaIionknown asIhc
Cambrian cxp!osion. "`' BuI Ihc cxpcrimcnI did noI conIro! !or Ihc
dcrcc o! rcproducIivc iso!aIion. Sincc Ihc !ossi!s o! cukaryoIcs prc-
dominaIc inIhaIrccord a simu!aIion o! cvo!uIion IhaImaIchcs iIs
LNL1L LK1NMb N NLLN1 KNbMb
opcn-cndcdcharacIcrshou!dbc conducIcdwiIhpopu!aIionsIhaIcan
spccaIc IhrouhsIron rcproducIivc iso!aIion.
Oncc oursimu!aIions rcach Ih posI-Cambrian pcriod wc nccd Io
cnrich Ihcm wiIh ncw rcsourccs. So !ar Ihc simu!aIcd unicc!!u!ar
oranisms havc bccn ab!c Io !carn how Io so!vc surviva! prob!cms
on!yasaspccics.ThaIis, Ihc so!uIionsarc rcachcdbyancnIircpopu-
!aIionovcrmanycncraIions. BuIa!IcrcukaryoIcsundcrwcnIIhcir
cxp!osivc divcrcncc chancs in IhcircomposiIion crcaIcd Ihc condi-
Iions !or Ihc cmcrcncc o! prob!cm so!vin abi!iIics IhaI cou!d bc
cxcrciscd durina sin!c oranisms!i!cIimc. Ls crcaIurcs, in!acI,
canbc uscdin Ihis modc bya!!owinIhc sIrcnIh o!Ihc connccIions
o!Ihcirncura!ncIsIobccsIab!ishcdnoIbyacncIica!oriIhmbuIby
IhcIraininIhc crcaIurcscIasIhcymovc aroundscarchin!or!ood.
ThcncxIsIcpinIhisinvcsIiaIionshou!dIhcrc!orcbc a discussiono!
Ihc ncrvous sysIcm and o! Ihc cmcrcnI !camin capaciIics wiIh
which iIcndows!ivin crcaIurcs.
V
d
LHPJL b
NOJ|d| NO5 d0 5OC O|||OCO
Thc cxp!osivc divcrcncc IhaImu!Iicc!!u!aroranismsundcrwcnIsix
hundrcdmi!!ionycars ao hadmany conscqucnccs. Thc divcrsiIyo!
Ihcsurviva!prob!cmsIhaIIhcscoranismshadIoso!vc,!orcxamp!c,
incrcascd cnormous!y as Ihcir cnvironmcnIs h!cdwiIh oIhcr !ivin
crcaIurcscapab!c o!a!!ccIinandbcin a!!ccIcdbyIhcm. Tocon!ronI
Ihis divcrsiIy oranisms bcan Io dcvc!op inIcma! modc!s Io uidc
Ihcirbchavior. vcnhumb!cmoIi!c bacIcria, asIhcyswimup aradi-
cnI o!nuIricnIsIowardIhc poinI o!maximumconccnIraIion, canbc
said Io havc an inIcma! modc! o! Ihcir cnvironmcnI.' This inIcma!
modc!isnoI, o! coursc, bascd onrcprcscnIaIionso!anykindandiI is
noIa modc! o!Ihc wor!daI!arc buI on!yo!Ihc concrcIc opporIuni-
Iicsandrisksa!!ordcdIobacIcriabyIhcirimmcdiaIc surroundins. U
oIhcrwords,!romIhcbcinniD o!!i!cIhcinIcma!modc!smcdiaIin
IhcinIcracIionbcIwccnaprimiIivcscnsorysysIcmandamoIorappa-
raIus cvo!vcd in rc!aIion Io whaI was dirccI!yrc!cvanI orsinihcanI
Io!ivin bcins. WiIhIhc advcnI o! mu!Iicc!!u!ar oranisms andIhc
prorcssivc di!!crcnIiaIion o! Ihcir cc!!s inIo mu!Iip!c kinds Ihc scopc
!or inIcrna! modc!s was rcaI!y cn!arcd. U parIicu!ar, a ncw kind
o!bio!oica!maIcria!,ncurona!maIcria!,bcanIorowandinIcrpcn-
cIraIc Ihc !ivin|c!!y accumu!aIin aI Ihc boIIom o! Ihc occan. And
wiIhIhc avai!abi!iIy o! ncurons Ihc capaciIyIo disIinuish Ihc rc!c-
vanI!romIhcic!cvanI, Ihc abi!iIyIo!orcround on!yIhc oporIu-
niIics and risks pushin cvcryIhin c!sc inIo an undi!!crcnIiaIcd
backround, wasvasI!yincrcascd.
Simp!cmu!Iicc!!u!aroranisms, !ikcIhchydraorIhc|c!!yhsh,havc
a !cwscparaIcncurons!inkcd IocIhcrinIoa ncIwork. U Ihc hydra
NLUKL NL1b N lNbLL1 N1LLLLNLL
Ihis ncIworkis conccnIraIcdaroundIhc mouIhwhi!c inIhc|c!!yhsh
iI is !inkcd Io simp!c rcccpIors !or !ihI and chcmica! subsIanccs.
Thcsc asscmb!ics o!ncurons, inIurn, susIain Ihc cmcrcnI capaciIy
Io modu!aIc inhcriIcd bchaviora! rcsponscs durin Ihc !i!cIimc o!
an oranism. Thc simp!csI o! Ihcsc capaciIics is ca!!cd hubiruuricn. iI
opcraIcs on scnsory-moIor acIiviIics IhaI arc innaIc buI iI a!ows
oranisms Io conIro! Ihcir inIcnsiIy. A oranism !ikc a hydra, !or
insIancc, can radua!!y dccrcasc iIs rcsponsc Io a sIimu!us-can
bccomc habiIuaIcdIoiI-as cxpcricncc hndsiIharm!cssoravoidab!c.
U oIhcr words, habiIuaIion Irans!orms a sinihcanI sIimu!us inIo
aninsinihcanI onc. Thc opposiIc capaciIy, scnsiIizaIion, a!ows Ihcsc
oranisms Io bchavc Ioward a prcvious!y irrc!cvanI sIimu!us as a
poIcnIia! sourcc o! an opporIuniIy or a risk, IhaI is, iI makcs Ihc
sIimu!usbchaviora!!y rc!cvanI. U a scnsc Ihcsc IwoancicnI!orms o!
!carninmark Ihc cmcrcncc o! sub|ccIivc radicnIs. bc!orc ahydra
bccomcs habiIuaIcd Io a novc! sIimu!us, !or cxamp!c, iI may bc
IhouhI o! as bcin surpriscd" by iI, a proIo-sub|ccIivc sIaIc IhaI
s!ow!yd1minishcsunIi!iI disappcarsasiIrcachcs cqui!ibrium. Bchind
Ihcscsub| ccIivcradicnIs,onIhcoIhcrhand,Ihcrcarcob|ccIivconcs.
conccnIraIion radicnIs o! c!ccIrica!y charcd subsIanccs consIanI!y
produccd and mainIaincd in Ihc Buids insidc and ouIsidc ncurons.
Thc capaciIy IobccomchabiIuaIcdIocxIcma!sIimu!i can scrvc as a
usc!u!sIarIinpoinI!oraphi!osophica!invcsIiaIiono!anima!!cam-
in bccausc cxp!ainin iI dcmands !ookin on!y aI whaI happcns
insidc a sin!c scnsory ncuron as iI inIcracIs wiIh a sin!c moIor
ncuron.
Thc sIandard modc! o! ncurons p!accs Ihc manipu!aIion o! radi-
cnIs o! mcIaic ions, poIassium orsodium ions, !or cxamp!c, aI Ihc
hcarI o! Ihcir !uncIion1D. Nanipu!aIin Ihcsc chcmica! radicnIs
kccpin poIassium aI hih conccnIraIions insidc Ihc cc!! and aI !ow
conccnIraIions ouIsidc o! iI-produccs ycI anoIhcr radicnI, onc o!
c!ccIrica! poIcnIia!. Thc kcy Io Ihc convcrsion o! onc radicnI inIo
anoIhcr is Ihc sc!ccIivc pcrmcabi!iIy o! a ncurons mcmbranc Io
mcIa!!icions. asmcmbrancpcrmcabi!iIychancsinst!owinandouI
o! Ihc cc! crcaIin variaIions inIhc radicnI o! c!ccIrica! poIcnIia!.
Ncuronsusc IhcscsmooIh!ow!cvcvariaIionsIorccordIhcc!ccIrica!
sina!s acIin asIhcir inpuI. Iroducin an ouIpuI invo!vcs chanin
Ihc c!ccIrica! radicnIs !rom conIinuous Io discrcIc, Irans!ormin
dI
dZ
N LbN N b MUL1N
IhcminIoachainorIraino!pracIicaIIyidcnIicaIcIccIricaIspikcs.This
spikc Irain is uscdIo producc a chcmicaI sinaI IhaI, whcn rcccivcd
by oIhcr ncurons, incrcascs Ihc pcrmcabiIiIy o! Ihcir mcmbrancs i!
Ihc chcmicaI subsIanccs arc cxciIaIory or dccrcascs iI i! Ihcy arc
inhibiIory.' Toprovidc a mcchanism !or habiIuaIion in simpIc crca-
Iurcs aII wc nccd Io assumc is IhaI Ihc suppIy o! an cxciIaIory
subsIancc in a scnsory ncuron bccomcs prorcssivcIy dcpIcIcd a!Icr
succcssivc sIimuIaIions, soIhaI Icss and Icss o!Ihis subsIancc crosscs
Ihc ap Io a!!ccI Ihc pcrmcabiIiIy o! a moIor ncurons mcmbranc
IcadinIoadccrcasciniIsrcsponsc.AsimiIarmcchanismcanaccounI
!or scnsiIizaIioni! wc add a sccond scnsory ncuron inIcracIinwiIh
Ihc hrsI causin iI Io incrcacproducIion o!Ihc cxciIaIory subsIancc
and Ihcrc!orc Ihc amounI o! Ihc IaIIcr IhaI rcachcs Ihc moIor
ncuron.
Thc capaciIy Io bccomc habiIuaIcd or scnsiIizcd impIics onIy Ihc
posscssion o! innaIc inIcrnaI modcIs. BuI wiIh Ihc advcnI o! morc
compIcx asscmbIics o! ncurons ncw !orms o!Icarnin cmcrcd IhaI
madc possibIc Ihc crcaIion o!inIcrnaI modcIs durin Ihc Ii!cIimc o!
an oranism. U parIicuIar, ancicnI insccIs wcrc capabIc o! Ihc kind
o! Icarnin rc!crrcd Io as clussiculccndiricninjin which an inhcriIcd
associaIion bcIwccn a sIimuIus and a rcsonsc providcs Ihc basis Io
!orm a novcI associaIion wiIh a prcviousIy insinihcanI sIimuIus.
SIudyin Ihis !orm o!Icarnin in a IaboraIory invoIvcs idcnIi!yin a
parIicuIar bchavior [ saIivaIion in a do, !o cxampIc) IhaI is aIways
Iricrcd by ivcn sIimuIus [!ood odor) . I! Ihc Iricr is prcscnIcd
immcdiaIcIyprcccdcdbyancuIraIsIimuIus [Ihc soundo!a bcII) Ihc
IaIIcr bccomcs capabIc a!Icrmany prcscnIaIions Io cIiciI Ihc dcsircd
rcsponscaIonc.AIIhouhmammaIswcrcoriinaIIyuscdIoprovcIhc
cxisIcncco!IhisassociaIivc capaciIyiI bccamccIcarIaIcrIhaIinsccIs
aIso posscss iI. Whcn Ihc anIcnnas o! honcy bccs, !or cxampIc, arc
Iouchcd by a drop o! suar insoIuIion Ihc bccs rctIcxivcIy cxIcnd a
parI o!Ihcir mouIh IowardIhc suar andIick iI. I!anodor ora coIor
whichwasprcviousIyirrcIcvanIIo Ihc bccs is madcIocoincidcwiIh
Ihc prcscncc o! suar iI W a!Icr somc Irainin Iricr Ihc Iickin
bchavior aIonc. TcchnicaIIy, Ihc sIimuIus Io which Ihc animaI is
cncIicaIIyprcdisposcd Io rcspond is caIIcdIhc "uncondiIioncd sIim-
uIus"whiIcIhc oncsIhaImaybc associaIcdwiIhiIarcrc!crrcd Io as
"condiIioncdsIimuIi. "
NLUKL NL1b N NbLL1 N1LLLLNLL
OuIsidc IhcIaboraIory Ihis associaIivc capaciIy has survivaI vaIuc
onIyi!IhccondiIioncdsIimuIusisin!acIprcdicIivco!Ihc prcscncc o!
Ihc uncondiIioncd onc. IarasiIic wasps, !or cxampIc, havc hunIin
rctIcxcs Iricrcdby Ihc odor o!IhccaIcrpiIIars onwhichIhcy!ccd
buIIhcy canIcarnIo dispIayIhcsamcbchavior whcnsIimuIaIcd by
Ihc odor o! Ihc pIanI Ihc caIcrpiIIar !ccds on. In oIhcr words, Ihc
wasps canIcarnIoprcdicIIhcprcscncco!Ihcirprcy Ihrouhacondi-
IioncdsIiuIus. Thc cvoIuIionaryrcIcvancc o! condiIioncdsIimuIiis
somcIimcs inorcd by psychoIoisIs who arc inIcrcsIcd onIy in Ihc
bcIwccn a sIimuIus and a rcsponsc. BuI bchavioraI ccoIoisIs
kDowIhaI cvoIuIion can buiId biascs as Io Ihc kinds o! sIimuIi IhaI
mayor maynoIbc succcss!uIIy associaIcd, and IhaI ccoIoicaIIy scn-
sibIc cxpIanaIionscanbcivcn!orIhoscbiascs.Foodscarchinhoncy
bccs, !orcxampIc,impIicsIcarninIorcconizcdi!!crcnItIowcrs, IhaI
is, ob|ccIsIhaIarccoIor!uI, havc a spcciaI aroma, andhavc comcIric
arrancmcnIso!pcIaIs. ThissucsIsIhaIIhcyshouIdbcabIcIoasso-
ciaIcnoIsoundsbuIodors, coIors,andmany-poinIcdshapcswiIhIhc
prcscncco!!ood. WccanaIsocxpccIanordcrino!Ihcsc sIimuIircIa-
ivc Io Ihcir usc!uIncss Io makc prcdicIions. tIorI odors shouId bc
casicr Io IcarnIhanIhc shapc o! tIowcrs sinccIhc IaIIcrvarics wiIh
IhcanIc o!obscrvaIionandisIhus Icss prcdicIivc o!Ihcprcscncco!
!ood. xpcrimcnIs wiIh honcybccs conhrm Ihcsc Iwo hypoIhcscs.`
CIassicaI condiIionin in insccIs iIIusIraIcs Ihc bcinnn o! aIruc
inIcrnaImodcI.whiIcIhcbchavioro!ahoncybccmaynoIbca!!ccIcd
by a tIowcr as a whoIc buI onIy bya scI o! sIimuIiIhaI Icnd Io co-
occur in Ihc prcscncc o! tIowcrs, Ihc !acI IhaI Ihcrcis a causaIIink
cIwccn cacho!Ihcsc sIimuIiandIhcir sourcc sucsIsIhaIasimpIi-
hcd modcI o! Ihc tIowcr [a scI o!Iinkcd sIimuIi) is indccd crcaIcd in
Ihc bccsbrain.
UnIikcIhc hydraand Ihc|cIIyhshinsccIs posscss a morc cIaboraIc
ncrvous sysIcminwhichmanyindividuaIncurons !ormIhccompo-
ncnIparIs o!Iarcr whoIcs, Iikca primiIivcbrainorIhcanIiaIhaI
coordinaIc Ihcir arIicuIaIcdImb movcmcnIs. NcvcrIhcIcss, wc nccd
noIconsidcrIhcircnIircncrvoussysIcmIo cxpIainIhcircapaciIy!or
cIassicaIcondiIionin.HiswcIIkDown,!orcxampIc,IhaImanynsccIs
can Icarn Io avoid ccrIan Iimb posiIions by associaIin Ihcm wiIh
an cIccIricaI shock buI Ihc samc associaIion can bc achicvcd by
isoIaIcdinsccIIcIocIhcrwiIhiIsanIion.Thus, iIisIhcpropcrIics
dJ
d+
NlLbN N bMUL1N
o!!oca!asscmb!ics o!ncurons, noI cnIircncrvous sysIcms, IhaIsus
IainIhis capaciIy, propcrIics!ikcIhcirconnccIiviIy andIhc sIrcnIh
o!IhcirconnccIions. Thcsc Iwo propcrIics arc rc!aIcdbccausc a con
nccIion or synapsc bcIwccn Iwoncurons bccomcs sIroncr Ihc morc
o!Icn Ihc ncurons on ciIhcr sidc o! iI arc acIivc simu!Iancous!y. Or
morccxacI!y, simu!IancousacIiviIyint!ucnccs aconnccIionssIrcnIh
a!IhouhnoIncccssari!yinaconsIanIway. C!assica!condiIionin,!or
cxamp!c, is characIcrizcd by a !carnin curvc IhaI disp!ays a c!car
dccc!craIion.whi!caIIhcbcinnino!IhcIraininIhc|oinIprcscnIa
Iiono!Ihc condiIioncdanduncondiIioncdsIimu!ushas a!arcc!!ccI
on bchavior Ihc c!!ccI bccomcs !css and !css pronounccd wiIh Iimc.
This sucsIs Ihc cxisIcncc o! a sIaIc o! cxpccIaIion in Ihc anima!s
mind so IhaI Ihc morc surprisin Ihc sIimu!us Ihc !arcr Ihc condi-
Iionin c!!ccI'U addiIion Io Ihis sub|ccIivc radicnI a ncw kind o!
cnIiIymaybcnccdcdIo!u!!ycxp!ainc!assica!condiIionin.amcnIa!
rcprcscnIaIion. ThcorisIs havc IradiIiona!!y bccn ambiva!cnI abouI
whaI cxacI!y bccomcs associaIcd whcn an anima! is Iraincd. is Ihc
anima!nkin Ihc condiIioncd sIiu!us wiIh a paIIcm o! bchavior
[Ihc rct!cxivc rcsponsc) oris iI associaIin R wiIh a rcprcscnIaIion o!
Ihc uncondiIioncd sIimu!us? Whi!c bchaviorisI psycho!oisIs !avor
Ihc !ormcrcxp!anaIion!aboraIorycxpcrimcnIssucsIIhaIIhc!aIIcr
isIhc corrccI onc. i!ananima!is arIihcia!!ypara!yzcddurinIraiDin,
soIhaI Ihc rct!cxivc bchavior cannoI occur, an associaIion sIi!!!orms
and a!!ccIs Ihc anma!s bchavior oncc iIs abi!iIy Io movc has bccn
rccovcrcd.
Givcn IhaI wc donI know whaI a rcprcscnIaIion in insccIs
mindis !ikc, oIhcrIhanIhc !acI IhaI iI cannoIpossib!ybc a symbo!ic
rcprcscnIaIion, iIwi!!bcusc!u!Iocxp!orcIhc qucsIiono!insccIinIc!
!icncc aIIhcmcchanism-indcpcndcnI!cvc!.ThaIis,wcnccdsimu!a
IionsIhaIdisp!ayancmcrcnIcapaciIy!orc!assica! condiIionincvcn
u Ihc mcchanism o! cmcrcncc is cnIirc!y di!!crcnI !romhaI o! rca!
ncurons. On Ihc oIhcr hand, Io makc surc IhaI wc arc cxp!orin
a spacc o! possibi!iIics IhaI ovcr!aps IhaI o! rca! ncurons wc nccd
simu!aIions inhich Ihc compuIin uniIs inIcracI wiIh oncanoIhcr
Ihrouh cxciIaIion and inhibiIion: in which somc !orm o! !carnin
Iakcs p!acc as Ihcsc inIcracIionsmodi!yIhc sIrcnIho!IhcirinIcrcon
nccIions: and in whih non-symbo!ic rcprcscnIaIions o! rc!aIivc!y
simp!c paIIcrns cmcrc sponIancous!y !rom Ihc dynamics. Thc Iypc
NLUKL NL1b N lNbLL1 N1LLLLNLL
o! simu!aIion IhaI saIishcs Ihcsc rcquircmcnIs is ca!Jcd an urri]ciul
ncurul ncr. A Iypica! ncura! ncI consisIs o! many compuIin uniIs
connccIcd Io cach oIhcr in a ccrIain paIIcm, cach compuIin uniI
bcin much !css sophisIicaIcdIhan Ihc ccnIra! proccssin uniI insidc
adcskIopcompuIcr.a!JiImusIbcab!cIodoisca!cu!aIciIsowndcrcc
o! acIivaIion ivcn Ihc cxciIaIory or inhibiIory sina!s iI rcccivcs
!rom oIhcr uniIs. Thc compuIinuniIs inIcracIIhrouhconnccIions
IhaI can vary B Ihcir abi!iIy Io IransmiI cxciIaIion or inhibiIion,
incrcasinB capaciIyi!Ihc inIcrconnccIcduniIs arc acIivc IocIhcr
ordccrcasini!Ihcy arc simu!Iancous!yinacIivc. Thc variab!c capac
iIyIoIransmiIcxciIaIionorinhibiIionis ca!!cdIhcsIrcnIhorwcihI
o!Ihc connccIion.
Thcsimp!csIncura!ncIhas compuIinuniIsarrancdinIwo!aycrs,
IhcinpuI!aycrandIhcouIpuI!aycr,adcsinknownasapcrccpIron. "
Simp!ypuI, a pcrccpIronisa dcvicc IhaI maps aninpuI paIIcminIo
an ouIpuI paIIcm." Onc paIIcm may bc, !or cxamp!c, a simp!c scn-
sory sIimu!us whi!c Ihc oIhcr may bc a simp!c moIor rcsponsc,
a!Ihouh in pracIicc ncura! ncIs arc sIudicd in a discmbodicd sIaIc
inorin Ihc dcIai!s o! Ihc scnsors producin Ihc inpuI paIIcm and
Ihosc o! Ihc acIuaIors IhaI convcrIIhc ouIpuI paIIcrninIo a moIor
acIion.Thc associaIion bcIwccninpuIandouIpuIpaIIcmsis crcaIcd
by rruininjIhcncura!ncI.Thc IraininproccssconsisIsinrcpcaIcd!y
prcscnIin apaIIcmIo IhcinpuI!aycr, acIivaIin somcuniIsbuI noI
oIhcrs, whi!chxin a dcsircd acIivaIion paIIcm aI Ihc ouIpuI !aycr.
Or in morc cmbodicd Icrms, Ihc Irainin invo!vcs sIimu!aIin Ihc
ncura! ncIs scnsors whi!c simu!Iancous!y showin iI Ihc dcsircd
moIorrcsponsc. Oncc boIhacIivaIionpaIIcms arc scIIhc compuIin
uniIs B cach !aycr bcin Io inIcracI wiIh cach oIhcr, incrcasin or
dccrcasin IhcwcihIo!IhcirconnccIions dcpcndinonwhcIhcrIhc
uniIs aIboIh cnds o!a onnccIion arc acIivc orinacIivc. Thc simu!Ia
ncous prcscnIaIion o! inpuI and ouIpuI paIIcms is pcr!ormcd many
Iimcs unIi! a sIab!c conhuraIion o! wcihIs in Ihc connccIions is
produccd. A!Icr Ihc Irainin is ovcr Ihc hxcd ouIpuI paIIcm is
rcmovcd and Ihc ncura! ncI is IcsIcd Io chcckwhcIhcriI R ab!c Io
rcproducc iIwhcncvcriIisprcscnIcdwiIhIhcoriina!inpuIpaIIcm.
ThiscmcrcnIcapaciIy is cxp!aincdby Ihc !acIIhaIIhc connccIions
havcnowspccihcwcihIs andwi!!IransmiIacIivaIion!romIhcinpuI
!aycr Io Ihc ouIpuI !aycr in |usI Ihc way nccdcd Io rcproducc Ihc
db
d
NlLbN N blMUL1lN
ouIpuI paIIcm. U morc cmbodicd Icrms Ihc hna! producI o! Ihc
Irainin proccss is Ihc abi!iIy Io rcconizc a sIimu!us, a rcconiIion
cxprcsscdbyIhc producIion o!Ihc corrccI moIor rcsponsc.
c!orcdiscussinIhc morc comp!cxncurancI dcsins nccdcdIo
simu!aIc c!assica! condiIionin iIwi!!bc usc!u!Ioivcancxamp!c o!
howsimp!c pcrccpIrons canbc uscd Io modc!IhcinhcriIcdrct!cxcs
IhaI arc Ihc basis o! IhaI kind o! associaIivc !carnin. By dchniIion
innaIc rcBcxivc bchavior is !carncd by a spccics ovcrmany cncra-
Iions noI by oranisms durin Ihcir !i!cIimc. This siIuaIion can bc
simu!aIcdbycoup!in ncura! ncIs Io cncIica!oriIhms a!!owin Ihc
conhuraIion o!wcihIs IhaI consIiIuIcs Ihc producI o! Ihc !camin
proccss Io bc discovcrcd noI by Irainin buIby cvo!uIionary scarch.
In onc simu!aIion Ihc cmcrcncc o!innaIc !ocomoIivc bchavior in
insccI-!ikc crcaIurcs was capIurcdwiIh pcrccpIrons p!ayinIhc ro!c
o!Ihcan!iaIhaIconIro!Ihcdi!!crcnIarIicu!aIcd!imbso!rca!insccIs.
Thcbodicso!Ihc crcaIurcswcrcmadc o!scvcra!parIs [Iorsos, !imbs,
hcads) arIicu!aIcd in a varicIy o! ways. somc arIicu!aIions acIcd !ikc
hincs consIrainin roIaIiona! moIion in onc dirccIion: oIhcrs wcrc
!ikc aba!! and a sockcIa!!owin roIaIion in a!l dirccIions. achbody
parI was spccihcd by a sin!c cnc" codin in!ormaIion abouI iIs
sizc, Ihc Iypc o!| oinIIhaIiIhadwiIh anoIhcr body parI, andhc con-
sIrainIs onIhccxIcnIIowhichIhcsc|oinIscou!dmovcorroIaIcThc
imu!aIcd cnca!sospccihcdIhcpropcrIics o!hcpcrccpIronnccdcd
Io conIro! cach arIicu!aIion. Ihc numbcr o! uniIs in iIs inpuI and
ouIpuI !aycrs, Ihcir connccIions, andIhc cvo!vab!c wcihIs o! Ihosc
connccIions. Thc simu!aIion, in !acI, did noI usc cncIic a!oriIhms
buIcncIicprorammin, soIhcchromosomcwasnoIasimp!c sIrin
o!oncs andzcrocsbuIancxccuIab!cproram spcci!yinhowIobui!d
a bodyparI andiIs ncura! ncI. This wayi!a body parIwasbui!IIwicc
asarcsu!Io!cncIicvariaIion cach copy cou!d bc rown wiIhiIs own
inIcrna!ncura!ncI and wou!dD immcdiaIc!y!uncIiona!.
ThcinpuI!aycro!cachncura!ncIrcccivcd scnsoryin!ormaIionnoI
!rom Ihc cxIcrna!cnvironmcnIbuI!rom Ihc insccIs ownbody, IhaI
is, proprioccpIivc in!ormaIion. Simu!aIcd scnsors incacharIicu!aIion
providcd in!ormaIion abouI|oinI-an!cs and abouI whcIhcr conIacI
had bccn madc bcIwccn onc body parI and anoIhcr, or bcIwccn a
body parI and a porIion o! Ihc cnvironmcnI. Thc ouIpuI !aycr, in
Iun, conIro!!cd Ihc movcmcnI o! Ihc | oinIs Ihcmsc!vcs [pushin,
NLUKL NL1b N lNbLL1 lN1LLLlLNLL
pu!!in, roIaIin, bcndin) Ihrouhsimu!aIcdacIuaIors. ' U addiIion
Iobcin cmbodicd Ihc crcaIurcs wcrc siIuaIcd in a spacc IhaI mim-
ickcd Ihc physica! characIcrisIics o! ciIhcr waIcr or dry !and. This
maIIcrcd bccausc cvo!uIion a!ways Iakcs p!acc in physica! cnviron-
mcnIs IhaI providc oranisms WH opporIuniIics and risks, !ikc a
c!uIIcrcd spacc IhaI a!!ords Ic opporIuniIyIo wa!k in somc dircc-
IionsbuInoI oIhcrs or a c!i!!IhaIa!!ords Ihc risk o!!a!!in. AIIhc
sIarIo!Ihc simu!aIion apopu!aIiono!random!y asscmb!cdcrcaIurcs
incapab!c o! cohcrcnI moIion was un!cashcd in Ihc compuIcr. Thc
Iypc o! !ocomoIivc bchavior Io bc cvo!vcd was Ihcn dccidcd. !or
swimminbchaviorraviIywasIumcdo!!,waIcrviscosiIyIumcdon,
and Ihc hIncss !uncIion was madc Iorcward rcaIcr spccd in boIh
horizonIa! and vcrIica! dirccIions: i!wa!kin bchavior was Ihc IarcI
asIaIicroundwiIh!ricIionwasaddcd, raviIywas Iumcd on, viscos-
iIyo!!, andhorizonIa! spccdwasuscdIocva!uaIchIncss. A!Icrmany
cncraIionsIhcrcsu!IwasIhcsponIancouscmcrcncco!a!arcvari
cIy o! succcss!u!!ocomoIion sIraIcics. snakc!ikc sinusoida! moIion:
Ihc usc o!addlcs, symmcIrica!t!ippcrs, orcvcnscIso!manyt!ippcrs
!orswimmin: avaricIyo!IcrrcsIria!aiIs bascd on!c-!ikc appcnd-
acs. Somc o! Ihc !ocomoIivc sIy!cs rcscmb!cd Ihosc uscd by rca
insccIswhi!coIhcrssccmcdcnIirc!ya!icn. ' '
Thc ouIcomc o!Ihis simu!aIion showcd IhaI comp!cx coordinaIcd
movcmcnIs cou!d cmcrc wiIhouI Ihc nccd !or ccnIra! commands
!rom Ihc brain. ThaI is, cach body parI and iIs arIicu!aIion [wiIh iIs
ownncura!ncI, scnsors, and acIuaIors) p!aycd Ihc ro!c o! arcusab!c
bui!din b!ock a!!owin ovcra!! body moIion Io cmcrc in a com-
p!cIc!y dcccnIra!izcd way as bui!din b!ocks cocvo!vcd IocIhcr. As
mcnIiocdabovc,insccI!imbs andIhciran!ia sccmIohavca simi-
Iar dcrcc o! auIonomy. OnIhc oIhcrhand, insccIs a!so havc brains
IhaIcan cxcrcisc ccnIra!izcdcommandovcrIhcbchavioro!IhccnIirc
body, and a!Ihouh iso!aIcd !imbs canbc condiIioncdIo!camsimp!c
associaIions a comp!cIc simu!aIion o! c!assica! condiIionin nccds
ncura!ncIsIhaIdisp!ayIhc abi!iIicso!a simp!cbrain.TodoIhisIwo
improvcmcnIs musI bc madc Io Ihc basic dcsin. incrcasin iIs
compuIaIiona!powcrandiviniIIhccapaciIyIocncra!izc.ThchrsI
chanc is nccdcd bccausc anima!smaybc condiIioncdIoprcdicI an
c!ccIric shock !rom Iwo di!!crcnI sIimu!i [a !ihI and a sound) whcn
prcscnIcd individua!!ybuInoIwhcn prcscnIcd IocIhcr. ThaIis, Ihcy
d
dd
NLbN N b MUL1N
wiIIproduccIhcrcLcxivcbchaviorwhcncxposcdIociIhcrIhcIihI
or Ihc sound buI noI Io boIh. This abiIiIy is isomorphic WU IhaI
nccdcdIo Icam somc cIcmcnIary IoicaI !uncIions [Ihc IoicaI !unc-
IioncxcIusivc or") IhaI carIy criIics o! ncuraI ncIs corrccIIy poinIcd
ouI was bcyond Ihc compuIaIionaI powcr o! simpIc pcrccpIrons
Thc sccondchanc is ncccssary bccausc whcninsccIs Icarnhrouh
cIassicaI condiIionin iI is vcry unIikcIy IhaI Ihcy wiII hndIhc cxacI
samc condiIioncd sIimuIus on cvcry occasion. This impIics IhaI, in
howcvcrsimpIca!orm, insccIsmusIbc abIc Iooranizc Ihcir cxpcri-
cnccIhrouhIhc usc o! cncraI caIcorics" o!whichsimiIarsIimuIi
arc parIicuIarinsIanccs.
BoIh improvcmcnIs can bc achicvcd by suppIyin a pcrccpIron
wiIh onc ormorc inIcrmcdiaIc Iaycs bcIwccn iIs inpuI and ouIpuI
Iaycrs. Bccausc Ihcsc cxIra Iaycrsarc noI acccssbIc !rom Ihc ouIsidc
Ihcyarc rc!crrcdIo ashiddnluyrs. A ncuraI ncIwiIhhiddcn Iaycrs
is caIIcd a muIIiIaycrpcrccpIron." AIIhouhIhis chanc may sccm
simpIciIIookycarsIobcpropcrIyimpIcmcnIcdbccauscnoonckncw
howIoIrainancuraIncIwiIhmuIIipIc Iaycrs.Thc corco!IhcIrain-
inproccssisIhc Icarnin ruIc, a proccdurc Ioad|usIIhc connccIion
wcihIs Iocnsurc a prorcssivc convcrcncc onIhc sIabIc conhura-
IionIhaIcanproducc Ihc IarcIouIpuIpaIIcrn. Anc!!ccIivcIcarnin
ruIc!ormuIIiIaycrpcrccpIrons isbascdonasimpIcbuIpowcr!uIidca.
insIcado! onIy aIIowin acIivaIion IoIIow!orward!romIhcinpuI Io
IhcouIpuIIaycrs,in!ormaIionabouIIhc dcrcco!mismaIchbcIwccn
Ihc currcnIIyproduccdouIpuIpaIIcmandIhc dcsircdhnaIpaIIcrnis
aIIowcd Io IIow backward. U oIhcr words, Ihc ruIc aIIows Icam
Ihrouh Ihc backpropaaIion o! crror cnsurin Ihc convcrcncc on
IhcIarcIpaIIcmIhrouhIhcprorcssivcminimizaIiono!Ihiscrror. '
AIraininproccssuidcdbyback-propaaIionconsIiIuIcsascarchin
Ihc spacc o!possibIc wcihI conhuraIions, a scarchproccss known
as radicnI dcsccnI" !or iIs rcscmbIancc Io Ihc dynamic Ihrouh
whichaphysicaI radicnI canccIs iIscI!. ' `
As in Ihc casc o! simpIc pcrccpIrons Irainin a muIIiIaycr onc
invoIvcs suppIyn iI wiIh a scnsory sIimuIus and a dcsircd moIor
rcsponsc.AsscvcraIdi!!crcnIbuIsimiIarinpuIacIivaIionpaIIcmsarc
prcscnIcdIoiI,andasin!ormaIionabouIIhc corrccIncsso!iIscurrcnI
ouIpuIpaIIcmIIowsbackwards,IhcuniIsinIhchiddcnIaycrdcvcIop
IhcirownpaIIcmo!acIivaIion. UnIikcIhcinpuIandouIpuIpaIIcrns
NLUKL NL1b N lNbLL1 N1LLLlLNLL
Ihc paIIcrn in Ihc hiddcn Iaycr is cmcrcnI andiIs composiIion is
inIIucnccdbysimiIariIicsinIhcdi!!crcnIscnsorypaIIcrnsincIudcdin
IhcIrainin scI. asIhcinpuIIaycrissIimuIaIcdby di!!crcnIpaIIcrns
and as Ihis acIivaIion IravcIs Io Ihc hiddcn uniIs Ihosc parIs o! Ihc
paIIcmsIhaIrcscmbIccachoIhcrhavc aIarcrc!!ccIonIhc shapc o!
Ihc cmcrcnIpaIIcrn Ihan Ihosc IhaI arc di!!crcnI. AIIhc cnd o!Ihc
IraininIhc connccIions bcIwccnIhcinpuIandIhc hiddcnIaycrhavc
acquircda conhuraIion o! sIrcnIhs IhaI can producc Ihc cmcrcnI
acIivaIionpaIIcrnwhccvcrIhcncuraIncIisprcscnIcdwiIhasimiIar
sIimuIus cvcni!docs noIbcIon IoIhcIraininscI. ThchiddcnuniIs,
H Iurn, wiII causc Ihc rihImoIor rcsponsc usin Ihc sIorcd wcihI
conhuraIion in Ihc connccIions bcIwccn Ihc hiddcn and ouIpI
Iaycrs.ThisimpIicsIhaIaIraincdmuIIiIaycrpcrccpIronhasIhccapac-
iIy Io cncraIizc !rom Ihc scnsorysIimuIiiI rcccivcddurinIrainin
Io many oIhcr simiIar sIimuIi. And Ihis, in Ium, sucsIs IhaI Ihc
cmcrcnI acIivaIionpaIIcrnbchavcsIiRc anon-symboIicrcprcscnIa-
Iiono!scnsorysIimuIi,arcprcscnIaIionIhaIcanbc uscdIorcconizc
rcIaIcd sIimuIi. OrIo puIIhis di!!crcnIIy, durinIrainin Ihc hiddcn
IaycrsIowIy xrruc urrcrcnr!rom Ihc di!!crcnIpaIIcrns conIaincd
in Ihc Irainin scI, a proIoIypc IhaI iI can usc Io producc Ihc rihI
moIorrcsponsc a!Icr Irainin.' Thcsc cmcrcnInon-symboIicrcprc-
scnIaIions andIhc wayIhcycapIurc simiIariIics inwhaIIhcy rcpc-
scnI arc cxacIIy whaI wc nccd Io undcrsIand in whaI scnsc insccI
brainscanbuiIdinIcrnaImodcIs o!IhcircnvironmcnI.
Wc arc nowinaposiIionIo showhowmuIIiIaycrpcrccpIrons can
bc uscd Io simuIaIc Icarnin Ihrouh cIassicaI condiIionin. xisIin
dcsins can bc quiIc compIcx bccausc Ihcy aIIcmpI Io rcproducc
aII Ihc di!!crcnIcharacIcrisIics o!Iiskindo!associaIivc Icam. Ihc
shapc o! Ihc Icarnin curvc: Ihc capaciIy Io associaIc many condi-
IioncdsIimuIi[ odors, coIors, IIowcrshapcs): andIhcincrcascinasso-
ciaIivc sIrcnIh !or condiIioncd sIimuIi IhaI havc morc prcdicIivc
powcr. ''BuIIhcbasicidcacanbc cxpIaincdinasIraihI!orwardway.
Wc nccd a dcsin consisIin o! Iwo muIIiIaycr pcrccpIrons, onc Io
cncraIc a non-symboIicrcprcscnIaIion o!Ihcunc

ndiIioncdsIimu-
Ius and Ihc oIhcr Io cncraIc onc o! Ihc condiIioncd sIimuIus. Thc
hrsI ncuraI ncI pIays Ihc roIc o! an inhcriIcd rcLcx so iIs conhura-
Iion o! wcihIs musI bc riidIy hxcd asi! iI had bccn !oundIhrouh
cvoIuIionary scarch, whiIcIhc sccond onc musIbc abIcIoIcam !rom
dV
V
NLbN N bMUL1N
cxpcricncc, IhaI is, Ihc wcihIs o! iIs connccIions musI bc !ound
IhrouhradicnIdcsccnI. FinaIIy, IhchiddcnuniIso!cachncuraIncI
shouId bc connccIcd Io cach oIhcr IaIcraIIy in such a wayIhaIIhcir
non-symboIcrcprcscnIaIions can inIcracIwiIhonc anoIhcr. urin
Irainin, as cach ncuraI ncI is sIimuIaIcd wiIh Ihc paIIcms corrc-
spondinIoIhcIwo sIimuIi, Ihc cmcrcnI rcprcscnIaIionsIhaI!orm
inIhchiddcnuniIswiIIincrcascIhcwcihIo!somco!IhcIaIcraIcon-
nccIionsanddccrcascIhcwcihIo! oIhcrs. A!IcrIrainin, IhcIaIcraI
connccIions wI posscss a conhuraIiono!wcihIs abIc Io producc
IhcrcprcscnIaIiono!IhcuncondiIioncd sIimuIus i!ivcnIhaIo!Ihc
condiIionconcandviccvcrsa.ThismcansIhaIwhcnIhcinpuIIaycr
o!IhcsccondncuraIncI isprcscnIcdwiIhIhc condiIioncd sIimuIus
iI wiII crcaIc in iIs hiddcn uniIs a paIIcm o! acIivaIion capabIc o!
producin Ihc cxIracIcdproIoIypc o Ihc uncondiIioncd sIimuIus in
Ihc hrsI ncuraI ncI which, in Ium, wiII Iricr Ihc inhcriIcd moIor
rcsponsc.
Thc conccpI o! aproIoIypccxIracIcd !rom cxpcricncc and sIorcd
as a non-symboIic rcprcscnIaIionwiIIpIay sucha cruciaIroIcin Ihc
cxpIanaIion o!animaI bchavior nIhc !oIIowinIwo chapIcrs IhaIiIs
naIurc shouId bc madc vcry cIcar. FirsI o!aII, an cmcrcnI rcprcscn-
IaIion is noI cxpIiciIIy sIorcd as such, Ihc producI o! Ihc Icamin
proccssbcinaconhuraIiono!connccIionwcihIsIhaIcanrccrcaIc
iIwhcnprcscnIcdwiIhIhcrihIinpuI.U oIhcrwords, whaIissIorcd
isnoIasIaIicrcprcscnIaIionbuIIhcmcansIo dynamicaIIyrcproducc
iI. Sccond,unIikcaphoIoraphIhcscrcprcscnIaIionsarcdispcrscdor
disIribuIcdB aIIIhchiddcnuniIsand arc Ihus cIoscrIo a hoIoram.
ThismcansIhaIIhcycanbcsupcrimposcdononcanoIhcrsoIhaIIhc
samc conhuraIion o!wcihIs can scrvc Io rcproducc scvcraI rcprc-
scnIaIions dcpcndin oniIsinpuI, simuIaIinIhc abiIiIy o!insccIsIo
associaIc scvcraI coIors or odors wiIhIhc prcscncc o! !ood. Thc dis-
pcrscdwayinwhichcxIracIcdproIoIypcsarcrcprcscnIcdissoimpor-
IanI!orIhccapaciIyo!ancuraI ncIIo cncraIizcIhaIIhcsc cmcrcnI
rcprcscnIaIionsarcusuaIIyrc!crrcdIoasdisIribuIcdrcprcscnIaIions.'
FinaIIy, unIikchc convcnIionaImbcIwccn a symboIandwhaIIhc
symboI sIands !or, disIribuIcd rcprcscnIaIions arc connccIcd Io Ihc
worIdinanon-arbiIrarywaybccauscIhcproccssIhrouhwhichIhcy
cmcrcisadirccI accommodaIionoradapIaIionIoIhcdcmands o!an
cxIcrnaIrcaIiIy. Thus, muIIiIaycrpcrccpIronso!!crapIausibIc accounI
NLUKL NL1b N NbLL1 N1LLLLNLL
o!IhcinIcnIionaIiIyo!mcnIaI sIaIcs. ' "WhcnncuraIncIsarcsIudicd
in a discmbodicd way, IhaI is, whcn Ihcir inpuI is prcscIccIcd and
prcsIrucIurcdbyIhccxpcrimcnIcr andIhcirouIpuIissimpIyanarbi-
IrarypaIIcrn o!acIivaIionIhaIhas no c!!ccI onan cxIcrnaI cnviron-
mcnI, Ihis cmcrcnIinIcnIionaIiIyisnoI dispIaycd. BuIIhcmomcnI
wc cmbody a ncuraI ncI and siIuaIc hc simuIaIcd body in a spacc
IhaI cana!!ccI iI andbc a!!ccIcdbyiI, Ihc crcaIurcs bchavcin a way
IhaIonc!ccIs compcIIcd!o characIcrizc as inIcnIionaI, IhaIis, asori-
cnIcdIowardcxIcmaIopporIuniIics andrisks.
Thcrcis onc morc qucsIionIhaInccds Io bc addrcsscdwhcn con-
sidcrinIhcappIicabiIiIyo!ncuraIncIsIosimuIaIcinsccIinIcIIicncc.
Whcn a simpIc pcrccpIron is Iraincd Ihc dcsircd moIor bchavior is
ivcnIoIhcmbyIhccxpcrimcnIcr.ThisimpIicsIhaI,unIikcinsccIsin
Ihcir naIuraI cnvironmcnI, ncuraI ncIs do oI Irain IhcmscIvcs.This
cxocnous Irainin is cvcn morc probIcmaIic in Ihc muIIiIaycr casc
bccausc Ihc ncuraIncI is suppIicdnoI onIywiIhIhc dcsircd ouIpuI
paIIcrnbuI aIso wiIhIhc dcrcc Io which iIs currcnI ouIpuI !aiIs Io
maIch Ihc IarcI paIIcrn aI any poinI durin Irainin. Norcovcr, in
boIhcascsiIisIhccxpcrimcnIcrwhodccidcswhaIpaIIcrnsIoincIudc
inIhcIraininscIandIhismayinIroduccsIaIisIicaIbiascsinIhcinpuI
IhaI arIihciaIIy!aciIiIaIc a Icam Iask. Any o! Ihcsc probIcms can
casI dubI onIhc cxIcnIIowhichIhc ncuraI ncIsIhcmscIvcs arc crc-
aIinaBD\bcIwccn IhcirinIcrnaI rcprcscnIaIions and Ihc cxIcrnaI
worId. ThcsiIuaIionissimiIarIoIhaIo!cxocnoushIncsscvaIuaIions
in Ihc casc o! cncIic aIoriIhms. In IhaI casc Ioo Ihc !acI IhaI Ihc
probIcm IhaI cvoIuIion musI soIvc is poscd by Ihc cxpcrimcnIcr
[IhrouhIhcdcsino!Ihc hIncss !uncon) mayinIroduccunwanIcd
biascs inIoIhcscarchproccss.WhcnncuraIncIsorcncIicaIoriIhms
arc uscdinindusIriaI appIicaIions Ihcsc biascs nccd noIbc aprobIcm
andmaycvcnbcdcsirabIci!IhcyshorIcnIhcamounIo!Iimcnccdcd
Io pcr!orm Ihc Irainin or Io conducI a scarch. BuI whcn uscd as
bioIoicaImodcIs cxocnousIrainin and cxocnous hIncss can bc
probIcmaIic. AwayouI o!Ihis diIcmma has aIrcadybccn sucsIcd.
cmbodyIhcimuIaIcdcncs orncurons andsiIuaIcIhcmin a spacc
IhaIprovidcshcmwiIhopporIuniIics andrisks.
Onco!IhcIwosimuIaIionsdiscusscdinIhcprcviouschapIcr, IaIcnI
cncrycnvironmcnIs [I) , iIIusIraIcs howhumans can bc rcpIaccd
noIonIyasanimaIbrccdcrsbuIaIsoasanimaIIraincrs.Whcndcscribin
VI
VZ
NlLbN N bMUL1N
Ls simuIaIions o! popuIaIion ccoIoy wc !ocuscd on onIy onc o!
Ihc Iwo ways inwhich iI can usc ncuraI ncIs. IinkIhcm Io cncIic
aIoriIhms and IcI cvoIuIion hnd Ihc conhuraIion o! wcihIs IhaI
improvcs!orainbchavior. BuI Ls ncuraIncIs canaIsobcconh-
urcd Io aIIow iIs simuIaIcd oranisms Io Icam Io !orac !or ood
durinIhcirIi!cIimcs. L cndocnizcs Ihc Iraininproccss Ihrouh
a cIcvcr adapIaIiono!back-propaaIion. IhcouIpuI Iaycr o!Ihcncu-
raIncIsproduccs noI onIy moIion insIrucIions [movc !orward, Ium
Ic!I,IumrihI) buIaIsoprcdicIionsrcardiDIhcpossibIcIocaIion o!
!ood. ThoscprcdicIions arc comparcdwiIhIhcrcsuIIsobIaincd!rom
!urIhcrcxpIoraIiono!IhccnvironmcnIandIhcdcrcc o!mismaIchis
propaaIcdbackIoIhcoIhcrIaycrs.'IdcaIIy, IhcscIwouscs o!ncuraI
ncIs-simuIaIin Icaminby Ihc spccics andIcamby Ihc oran-
ism-shouId bc uscd IocIhcr so IhaI somcIhin Iikc Ihc cIassicaI
condiIionindcsinouIIincdabovccouId cmcrc. ThchrsIo!IhcIwo
ncuraI ncIs, Ihc onc IhaI crcaIcs a disIribuIcdrcprcscnIaIion o! Ihc
uncondiIioncdsIimuIus, shouIdbcsIowIycvoIvcdwiIhinancnviron-
mcnI IhaI can consIraiD and cnabIc acIion Io yicId ctIcxcs Iinkin
scnsory sIimuIi Io adcquaIc moIor rcsponscs.Thc sccond ncuraI ncI,
Ihc onc wiIh disIribuIcdrcprcscnIaIions o! Ihc condiIioncd sImuIi,
musI arrivc aIa hnaI wcihI conhuraIion durin IhcIi!cImc o!Ihc
crcaIurcs, buI Ihc rcsI o!iIs archiIccIurc couId bc cvoIvcd usin prc-
dicIivc succcss as parI o! Ihc cndocnous cvaIuaIion o! hcss. This
wayIhcbiascsbuiIIbycvoIuIiononwhaI canpIay Ihc roIc o!acon-
diIioncd sImuIus-coIor, tIoraI aroma, pcIaI arrancmcnI, buI noI
sound iDIhc casc o! honcy bccs-couId cmcrc sonIancousIy in a
simuIaIion.
Novin bcyondIhc Icamin capaciIics o! insccIs dcmands a morc
radicaI sIraIcy IhaI simpIy cmbodyin muIIiIaycr pcrccpIrons and
siIuaIin Ihcm in spacc. In parIicuIar, Iarcr animaIs wiIh morc
cIaboraIc ncrvoussysIcms arc capabIc o! IcaminIo dcvcIop novcI
bchaviors. UnIikccIassicaIcondiIionininwhichIhc cxisIcncc o!an
inhcriIcd rctIcx Ioanchor an associaIion is ncccssary, ncw Icamcd
bchaviorscanbcdcvcIopcdIhrouhIhcirassociaIionwiIhrcwardsor
punishmcnIs. Somc insccIs may, in !acI, havc acccss Io Ims morc
sophisIicaIcdIcarnincapaciIy. Honcybccs, !orcxampIc,usccIassicaI
condiIionin Io Icam Io prcdicI Ihc prcscncc o! !ood buI Ihcy musI
aIso Icam by IriaIand crror how Io cxpIoiI a tIowcr bIossom oncciI
NLUKL NL1b N lNbLL1 N1LLLLNLL
hasbccnidcnIihcdNcvcrIhcIcss,IhccapaciIyIoIcarnnovcIbchav-
iors is rcaIIy ampIihcd in animaIs Iikcbirds andmammaIs bccausc
in Ihcir cascpcrccpIion and mcmory o bcyond Ihc worId o! mcrc
sIimuIi. Irovidin cvidcncc!orIhis cIaim andcxaminin Ihckindo!
ncuraIncIsIhaIcandispIay IhcscpcrccpIuaIandmcmorycapaciIics
wi!IbcIhc Iask o!Ihc !oIIowinchapIcr.
VJ
V+
LHPJL bLVLN
NOJ|d| NO|5 d0 Md00d|I d MO0O|V
Aspopu!aIionso!ncuronsrcwandproIi!craIcdinsidcIivincrcaIurcs,
aIhcrin inIo !aycrs and !o!din inIo c!aboraIc Ihrcc-dimcnsiona!
sIrucIurcs, Ihcy providcd an cvcr richcr subsIraIum !or Ihc rowIh
and proIi!craIion o!morc cIhcrca! cnIiIics. mcmorics. Thc mcmorics
IhaIbirds andmamma!s!ormo!acIuaIIyIivcdcpisodcs, !or cxamp!c,
arc morc or !css vividrc-cxpcricnccs o! cvcnIs or siIuaIionsin which
ob|ccIs p!ay spccihc ro!cs [such as acnIs or paIicnIs) and in which
Ihcir inIcracIions makc scnsc IoIhc anima!. Thc conIcnI o!auIobio-
raphica!mcmoricsin animaIsmusIbcIhouhIo! as cndowcd wiIh
sinihcancc noI wiIh sinihcaIion, which is a !inuisIic noIion. Thc
sinihcancc o! a sccnc or cvcnI is rc!aIcd Io iIs capaciIy Io makc a
di!!crcncc in ananima!sIi!c, Io iIs capaciIyIo a!!ccI andbc a!!ccIcd
byIhc anima!sacIions, whi!csinihcaIionisascmanIicnoIionrc!cr-
rin Io Ihc mcanin o! words or scnIcnccs. Birds and non-human
mamma!s may bc incapab!c o! dca!in wiIh sinihcaIion buI Ihcy
surc!y can aIIribuIc sincancc Io Ihc opporIuniIics and risks IhaI
Ihcir cnvirocnI a!!ordsIhcm.
vidcncc !or Ihc cxisIcncc o! di!!crcnI Iypcs o! mcmory comcs
mosI!y!romIhcirdissociaIion in amncsichumanpaIicnIs or in ani-
maIs IhaI havc undcronc surcryIo producc spccihc !csions in Ihc
brain.ThcIypc

o!mcmoryinvo!vcdinc!assica!condiIionin, whcIhcr
in insccIs, birds, ormamma!s, is rc!crrcd Io as proccdura!" bccausc
o!iIsconIcnIs!acko!acccssibi!iIyouIsidco!Ihc scnsory-moIorasso-
ciaIions IhaI iI cnab!cs. Ncmorics whosc conIcnI is acccssib!c arc
rc!crrcd Io as dcc!araIivc, " a caIcory!urIhcr subdividcd dcpcndin
NLUKL NL1b N MMMLN MLMK
onwhcIhcrIhc conIcnI o!Ihcmcmoryis cvcnIs andsiIuaIions IhaI
havc acIua!!y bccn cxpcricnccd, a Iypc o! mcmory rc!crrcd Io as
criscdicmcmcr, orwhcIhcriI consisIs o!IinuisIicaI!y cxprcsscd!acIs,
IhaIis, scmanIic mcmory. 'amain ccrIain arcas in anima!s brains
can scrious!y disab!c Ihcir capaciIy !or dcc!araIivc mcmory whi!c
!cavin proccdura! mcmory inIacI. SimiIar!y,

umans su!!crin !rom

ccrIain kinds o! amncsia can rcIain Ihcir capaciIy Io rcmcmbcr Iin-
uisIic in!ormaIion buI bc incapab!c o! rccaI!in auIobioraphica!
cvcnIs, and vicc vcrsa, Ihcy can rccaI! acIuaI!y Iivcd sccncs buI arc
unab!c Io rcmcmbcr whaI has bccn said IoIhcm.

Thc vcry !acI IhaI

cpisodicnd scmanIic mcmory can bccomc dissociaIcd sucsIs IhaI
human scnsorycxpcricncc andIhc mcmoricswc !orm o!iI arc indc-
pcndcnIo!!anuac andsIron!yrc!aIcdIo onc anoIhcr. BuI !or Ihc
purposc o!Ihis chapIcrwc donoIhavc Io scII!c Ihc conIrovcrsyovcr
IhcIhcsiso! Ihc !inuisIicaIiIy o! cxpcricncc.non-humanmamma!s
and birds do noI posscs !anuac so Ihcir scnsory cxpcricncc and
auIobioraphica!mcmorics cannoIpossib!ybc sIrucIurcdbyiI.
Un!ikc Ihc pcrccpIion o! aIomizcd sIimu!iin insccIs Ihc pcrccpIion
o! ob| ccIs wiIh an cndurin idcnIiIyis a comp!cx Iask, invo!vin Ihc
synIhcsiso!!caIurcs!romdi!!crcnIcxIcrna!aswcI!asinIcrna!scnscs.
IrcdaIory mamma!s, !orcxamp!c, musIuidcIhcir scarchinbchav-
iorby Ihc in!ormaIionIhcycI!romdi!!crcnI scnscs,usinsmcI!Io
brinIhcminIoc!oscproximiIyIoIhcirIarcIIhcnswiIchinIo sihI
or sound. And Ihcy musI consIanI!y caIibraIc Ihc in!ormaIion Ihcy
rcccivc!romIhc cxIcrna!wor!dIoIhIcncraIcdbyIhcirownprop-
rioccpIivc scnscs so Ihcy can disIinuish bcIwccn a chanc in Ihcir
visua!hc!dproduccdbyaprcysmovcmcnIsandIhcapparcnImoIion
crcaIcdby Ihc movcmcnI o!Ihcir own hcad or body. Thc synIhcsis
o! in!ormaIion !rom di!!crcnI scnsory modaIiIics is IhouhI Io bc
pcr!ormcdin an anima!s hippocampus andrc!aIcdarcas, an oran
a!so invo!vcd in Ihc !ormaIion o! cpisodic mcmorics. amac Io Ihc
hippocampus, !orcxamp!c, ipairspcr!ormancconIasksIhaIinvo!vc
comparin onc mcmory Io anoIhcror usin Ihc conIcnI o!amcmory
in a novc! conIcxI. UnIi! Ihc mcchanisms bchind Ihcsc capaciIics
arc !uI!y undcrsIood a phi!osophica! cxp!oraIion o! avian and mam-
ma!ian mcmory is bcsI conducIcd aI Ihc mcchanism-indcpcndcnI
!cvc!. Nuch as Ihc mu!Ii!aycr pcrccpIrons discusscd in Ihc prcvious
chapIcrhc!pcdusdcvc!opabcIIcridcao!whaIarcprcscnIaIioninan
Vb
V
N LbN N bML1N
insccIs brain couId bc Iikc-dcspiIc Ihc !acI IhaI Ihc simuIaIcd
mcchanisms, Iikc Ihc back-propaaIion o! crror, havcnoanaIomicaI
counIcrparI in rcaI brains-Ihc capaciIy o! ncuraI ncIs Io rcconizc
cndurinob|ccIsandmakc scnsc o!inIcracIions bcIwccn ob|ccIs can
bca rcaIaidin hnc-Iunin ourphiIosophicaIinIuiIionsabouIwhaI
ocsoninIhc minds o!Iarcr animaIs.
GivcnIhccompIcxiIyo!Ihcsub|ccIiIwiIIbcusc!uIIobrcakiIdown
inIo Iwo di!!crcnI capaciIics, ob|ccI rcconiIion and sccnc anaIysis,
ivc cxampIcs o! Ihc cvidcncc IhaI animaI psychoIoisIs havc
aIhcrcd !or IhcircxisIcncc, and Ihcn discuss Ihc currcnIIycxisIin
ncuraI ncI dcsins IhaI can approximaIc Ihcm. Thc carcsI Iabora-
Iory cvidcncc !or Ihc cxisIcncc o! Ihc hrsI o! Ihcsc Iwo capaciIics
comcs !rom cxpcrimcnIs caIIcd "Iwo-way cIassihcaIions. " o-way
cIassihcaIioncxpcrimcnIswcrchrsIconducIcdwIhpconsbuIwcrc
IaIcrcxIcndcdIomonkcys andoIhcrmammaIs.Iiconswcrcshown
pro|ccIcdphoIoraphsadIraincdIopcck aI a swiIchIhaIacIivaIcd
a !ood dispcnscr i! ccrIain kinds o! ob|ccIs appcarcd inIhc picIurcs.
Inonc cxpcrimcnI, !orcxampIc, ocra Ihousand sIidcs o!urbanand
counIrysidcsccncswcrcuscd,haI!o! whichconIaincdhumansindi!-
!crcnIposiIions[ siIIin,sIandin),wiIhdi!!crcnIappcaranccs[ cIoIhcd,
nakcd), and shoI !rom di!!crcnI anIcs. This varicIy cIiminaIcd Ihc
possibiIiIy IhaI Ihc picons wcrc rcacIin Io simpIc sIimuIi, such as
coIor orbrihIncss. A!Icr a !cw wccks Ihc picons hadacquircdIhc
capaciIy Io cIassi!y picIurcs inIo Ihosc IhaI conIaincd onc or morc
humans and Ihosc IhaI did nI. This rcsuII sIronIy sucsIs IhaI
Ihc picons wcrc abIc Io cxIracI a proIoIypc !rom Ihcir pcrccpIion
o! human hurcs and Ihcn usc iI Io asscss Ihc dcrcc o! IypicaIiIy
IhaIdi!!crcnI imacs had rcIaIivc Io IhcproIoIypc. Iicons wcrc aIso
shown capabIc o! pcr!ormin Iwo-way cIassihcaIions o! picIurcs
wiIhpiconsvcrsuspicIurcswiIhbirdsoIhcrIhanpicons: o!picIurcs
wiIh andwiIhouI Irccs: and o!picIurcs wiIh and wiIhouI a body o!
waIcr.`
Bc!orc showin how muIIiIaycr pcrccpIrons can dispIay simiIar
ob|ccI rcconiIon capaciIics onc aspccI o! Ihcir dcsin shouId bc
cIarihcd. UnIikcIhc ncuraIncIs discusscdinIhcprcviouschapIcr in
whichIhcinpuI IaycrrcccivcdscnsorysIimuIaIionwhiIcIhc ouIpuI
Iaycr conIroIIcd moIor bchavior, wc arc dcaIin hcrc wiIh parIiaI
modcIs o!amorc compIcxncrvous sysIcm.ThisimpIicsIhaIa ncuraI
NLUKL NL1b N MMMLN MLMK
ncIs inpuImaynoI comcdirccIIy !rom scnsorsbuI !rom oIhcrncuraI
ncIs,whiIciIs ouIpuImaybcsub|ccIIo!urIhcrproccssinbc!orciIis
convcrIcdinIo abodiIy acIionbyacIuaIors.U many cascs IhcncuraI
ncIsdirccIIyconnccIcdIociIhcr scnsors oracIuaIorsarcnoIincorpo-
raIcd inIo Ihc dcsinbuI|usI assumcd Io bc Ihcrc. U Ihc ncuraI ncI
IhaI rcccivcs scnsory in!ormaIion is noI incIudcd, !orcxampIc, Ihcn
Ihconc whosc paIIcm rcconiIion bchaviorisbcin sIudicd canhavc
iIs inpuI uniIs IabcIcd wiIh words, words IhaI sIand !or Ihc ob|ccIs
IhaI Ihc IaIIcr wouId rcccivc i! iIwcrcacIuaIIyconnccIcdIo Ihc !or-
mcr. SimiIarIy,i!IhcncuraI ncIIhaIconIroIsmovcmcnIisnoIparIo!
Ihc dcsin Ihc ouIpuI uniIs o!Ihc onc bcin sIudicd can bc IabcIcd
wiIhwords!ormoIoracIions.U IhcprcscnIcascusinwordsIoIabcI
IhcinpuIuniIswouId dc!caIIhcpurposco!IhccxpcrimcnIivcnIhaI
whaIwc nccdIosiuIaIcisabirdscapaciIyIorcconizc ob| ccIs!rom
Ihc visuaI in!ormaIion providcd by imacs. Thc ouIpuI uniIs, onIhc
oIhcrhand, canbcIabcIcdIinuisIicaIIysinccinarcaIbirdIhisouIpuI
wouId bc Ihc inpuI Io many oIhcr asscmbIics o! ncurons bc!orc
bccominamoIoracIion.
A od iIIusIraIion o! Ihc usc o! ncuraI ncIs !or Ihis purposc is a
simuIaIion dcsincd Io rcconizc human !accs. Thc inpuIIaycr con-
sisIcd o!aIwo-dimcnsionaI arrayo!uniIs, 64uniIs oncach sidc, inIo
whichanimaccouIdbcpro|ccIcd. achinpuIuniIwas connccIcdIo
a onc-dimcnsionaIhiddcn Iaycr consisIin o! 80 uniIsIhaI, in Iurn,
wcrc connccIcdIoanouIpuIIaycrconIainncihIuniIs IabcIcdwiIh
words. urin IraininIhcinpuIIaycr wasprcscnIcdwiIhicIurcso!
I I di!!crcnI human !accs [as wcII as o! ob| ccIs IhaI wcrc noI !accs)
whiIciIsouIpuIIaycrwasivcnanacIivaIionpaIIcrnIhcmcanino!
which was ivcn by Ihc IabcIs. onc uniI was IabcIcd !acc, bcin
acIivc i! Ihc inpuI was indccd a !acc and inacIivc i! iI was noI: Iwo
uniIswcrcIabcIcd!cmaIcandmaIc,IhciracIiviIysini!yincndcr
rcconiIion: and Ihc rcmaininhvc uniIswcrc IabcIcdwiIha namc
and a numbcr ssincd Io cach !acc. A!Icr bcin Iraincd Ihc ncuraI
ncIwasnoI onIyabIIopcr!orm a Iwo-way cIassihcaIiono!picIurcs
inIo Ihosc IhaI conIaincd and did noI conIain!accs, buI aIso Io cor-
rccIIy assin !acc picIurcs Io Ihc rihI cndcr caIcory and cvcn Io
idcnIi!y!accs bcIoninIospccihcpcrsons.ToIcsIiIsabiIiIyIo cncr-
aIizcIhc ncuraI ncI was IcsIcdwiIhpicIurcs o!Ihc samcpcopIcIhaI
wcrc noIparI o!IhcIraininscI andiI pcr!ormcd aImosI tIawIcssIy.
V
Vd
NLbN N bMUL1N
Norcovcr, iIcou!dcorrccI!yidcnIi!y!accsinwhich 2UpcrccnIo!Ihc
conIcnIwas dcIibcraIc!y obscurcdbycomp!cIinIhc!aa!paIIcrnwiIh
in!ormaIion!romIhc disIribuIcdrcprcscnIaIionsiniIshiddcnuniIs.
hc prcscncc o! !abc!s in Ihc ouIpuI uniIs dcsinaIin !inuisIic
caIcorics !ikc ma!c" and !cma!c"hadnoc!!ccI onIhcabiIiIyo!Ihc
ncra! ncI Io rcconizc !accs. Thc cxp!anaIion !or Ihis capaciIy Iics
insIcad in Ihc 80 uniIs o! Ihc hiddcn!aycr. Bc!orc Irainin aI! Ihc
pssib!c paIIcrns o! acIiviIy IhaI cou!d occur in Ihis !aycr !ormcd an
sIrucIurcdspacc o!80 dimcnsions. AsIhc ncuraIncIwasIraincdiI
rsI subdividcd Ihis inIcrna! spacc o! possibi!iIics inIo Iwo rcions,
onc corrcspondin Io!accs aDdIhc oIhcrIonon-!accs. FurIhcrIrain-
in!cdIhcncura!ncIIoparIiIionIhc !accrcioninIoIwosubrcions,
onc !or!cma!cIhc oIhcr!orma!c !accs, whichwcrc Ihcn!urIhcrsub-
dividcdinIo cvcnsmaI!crrcions!orindividua!!accs. Thc ccnIcr" o!
cach rcion corrcspondcd Io a proIoIypc-Ihc casicsI Io rcconizc
!accs inIhcIraiininpuIorIhc casicsIIorcconizcvicwso!anindi-
vidua! !acc-whiIc Ihc bordcrs bcIwccn rcions corrcspondcd Io
ambiuousinpuIs, suchaspicIurcsinwhichIhcconIcnIcou!dnoIbc
dcIcrmincd Io bc a !cc, or in which a !acc was disp!aycd in such
a wayIhaI iI was impossib!c Io dccidc whcIhcr iI bc!oncdIo ama!c
or a !cma!c. This simu!aIion providcs a powcr!u! insihI inIo how
an ob|ccIivc caIcory can bc capIurcd wihouI usin any IinuisIic
rcsourccs. Thc sccrcI is Ihc mappin o! rcluricnsc]similurin inrc rclu-
ricns c]rrcximinin Ihc possibiIiIy spacc o! acIivaIion paIIcms o! Ihc
hiddcn!aycr. ThaIis, ob|ccIs IhaI rcscmb!c cach oIhcrbccomc ncih-
borinpoinIsinIhc inIcrna!possibi!iIyspacc, andvicc vcrsa, ob|ccIs
wiIh a hih dcrcc o! dissimi!ariIy [!accs and non-!accs) cnd up as
poinIsIhaIarc !araway!rom cachoIhcrinIhc spacc o!possib!c acIi-
vaIionpaIIcrns.
Onc !imiIaIiono!Ihissimu!aIionis IhaI cvcnIhouhIhc!inuisIic
!abc!s o!Ihc ouIpuIuniIs donoIp!ayanycp!anaIoryro!c Ihcirprcs-
cncc indicaIcsIhaIwc arc dcaIin wiIh aparI!y discmbodicdncura!
ncI. Wc cou!d c!iminaIc Ihc nccd !or !abc!s y addin oIhcr ncura!
ncIsusinIhcoIpuI!aycrsacIiviIypaIIcrnIocncraIcmoIorbcav-
ior.hisbchaviorcou!dbcrc!aIivc!ysimp!c,Ihcsimu!aIcdcquiva!cnI
o! pcckin on a !ood dispcnscr. In !aboraIory anima!s Ihis kind o!
bchavior is noI achicvcdIhrouh c!assica!condiIioninbuI Ihrouh
a di!!crcnIIra\Din proccss caI!cd insrrumcnrulccndiricninj. InsIcad o!
NLUKL NL1b N MMMLN MLMK
usin a rcucx as iIs sIarIin poinI insIrumcnIa! condiIionin sIarIs
wiIh a bchavior IhaI Icnds Io occur sponIancous!y usin rcwads Io
incrcasc iIs !rcqucncy orpunishmcnIs Io dccrcasc iI. Thc posiIivc or
ncaIivcrcin!orccmcnImusIbcconIincnIonchancso!bchavior.i!
Ihc!rcqucncyo!occurrcncc docsnoIchancIhcrcin!orccmcnIis noI
ivcn. In addIion, !aboraIory cxpcrimcnIs musI inc!udc a sina!, a
simp!c sound orIihI, Iohc!pIhcanima!IcI!whichonc o!iIspossib!y
many bchaviors shou!d bc associaIcd wiIh Ihc posiIivc or ncaIivc
rcin!orccmcnI. This sina! is ca!!cd a discrminaIivc sIimu!us. " OuI-
sidc o!Ihc!aboraIoryrcin!orccmcnI!carnincan!cadIoIhcdcvc!op-
mcnI o! roIinc bchavior IhaI hc!ps an aDima! copc wiIh Ihc
comp!cxiIyo!asiIuaIionbuIiIcana!so!cadIoIhc!ormaIiono!supcr-
sIiIioushabiIsbcarinnorc!aIionIorca!opporIuiIics andrisks.Thc
ro!co!discriminaIivc sIim!iisprccisc!yIoa!!owananima!Io disIin-
uish Ihc bchaviorIhaI is bcin rcwardcd !rom co-occurrin bchav-
iorsIhaIarc irrc!canI.
Thc Iypica! !aboraIory ouIcomc o! insIrumcnIaI condiIionin, an
anima! IhaI!cs Io prcss a !cvcr Io obIaina pcI!cI o! !ood, ivcs a
vcry poor idca o!Ihc Iypcs o!bchaviorIhaI caDbc !carncd Ihrouh
Ihiskindo!Irainin.Thissip!c bchavioris uscdbccausc o!Ihc casc
wiIhwhichincrcascs ordccrcascsinIhc!rcqucncy o!!cvcr-prcssin
can bc mcasurcd. BuI wiIh carc!u! manipu!aIion o! rcin!orccmcnI
schcdu!cs aDd chaDcs in discriminaIivc sIimu!i comp!cx paIIcrns
o! bchavior can bc cncraIcd as cxcmpIihcd by Ihc c!aboraIc Iricks
IhaI caD bc IauhIIo circus anima!s. Thc IarcI bchaviormay D, !or
cxamp!c, a raI sIandin oniIs hind !cs ina ivcn comcr o!Ihc cac.
Thc cxpcrimcnIcr musIhrsIrcwardoinIoIhaI comcr, a bchavior
IhaIIakcsp!acc sponIancous!yoncc inawhi!c. U Ihischanccbchav-
ior is rcwardcd and a discriminaIivc sIimu!us is uscd Io !cI Ihc raI
knowwhaI is bcinrcwardcd, iIwiI!incrcasc in !rcqucncy. AI IhaI
poinIiI sIos bcin rcwardcdandadi!!crcnI!ow-!rcqucncybchavior,
such as movin Ihc hcad Io an uprihI posiIion, is rcwardcd aDd
IinkcdIoanoIhcrdiscriminaIivcsIimu!usunIiIiIs!rcqucncyincrcascs.
Thcproccdurc conIinucsoncbchaviora!paIIcma!IcrIhcncxI[raisin
Ihc !c!Ipaw, raisin Ihc rihIpa) unIi!Ihc dcsircd hna!bchavioris
obIaincd. This proccss is ca!!cd succcssivc approximaIion. "'
ThcmainobsIac!cIoIhc dcsino!ncra!ncIs ab!cIo!camIhisway
is noIcapIurinIhc capaciIyo!rcin!orccrsIoa!!ccIbuIIhc capaciIyo!
VV
I
NlLbN N bMUL1N
ananima!Iobc a!!ccIcdbyIhcm.Usin!oodasarcin!orccr,!orcxa-
p!c,imp!icsIhaIananima!ishunryor aI!casInoI!u!!ysaIiaIcd. BuI
simu!aIinhuncr wou!d sccmo invo!vc rcproducin aspccIs o! iIs
phcnomcno!oy, suchasIhcpanso!ancmpIy sIomach. ar!ycxp!a-
naIionso

huncr, in!acI,IooksIomachconIracIionsIobc

arIo!Ihc
mcchanismIhrouhwhichhuncr moIivaIcs!oodscckinbchavior.
This hypoIhcsis was shownIo bc !a!sc by cxpcrimcnIs inwhichIhc
sIomachs o!raIswcrc rcmovcd, aIIachinIhcircsophaiIoIhcirsma!!
inIcsIincs, !cavinIhc bchaviorassociaIcdwiIhhuncrinIacI. 'U Ihc
prcvious chapIcr wc arucd IhaI Ihc dcrcc Io which an anima! is
surpriscd or sIarI!cd bya novc! sIimuIus is a sub|ccIivc radicnI IhaI
can bc dissipaIcd by habiIuaIion buI iIs cxisIcncc prcsupposcs IhaI
o! an ob|ccIivc radicnI, a conccnIraIion o! cxciIaIory subsIanccs in
Ihc ncura! subsIraIc. And simi!ar!y !or huncr. conIracIions o! Ihc
sIomachandoIhcr componcnIs o!Ihc cxpcricncc consIiIIc asub|cc-
Iivc radinI buIoncIhaI isa!so undcrpinncd by anob|ccIivc onc,
such as conccnIraIions o! !ucosc in Ihc anima!s b!ood sIrcam.
Kcmovin a raIs sIomach c!imiaIcs Ihc !ormcr buI noI Ihc !aIIcr.
This mcans IhaI a!! a ncura! ncI musI capIurc is Ihc moIivaIiona!
c!!ccIo!huncrnoIanyIhinabouIiIs phcnomcno!oy.
Imp!cmcnIin Ihc capaciIy !or insIrumcnIa! condiIionin can bc
doncbyaddinIoIhcncura!ncIshand!inscnsoryandmoIoracIivi-
Iics oIhcrs acIin as a sourcc o!moIivaIion !or bchavior. U onc rc!a-
Iivc!ysimp!cimp!cmcnIaIion !ourncura!ncIs wcrc !inkcdIocIhcr,
Iwo !or inIcrna! rcin!orccmcnI [p!casurc and pain) and Iwo !or
moIorbcavior [ approachandavoid) . IaIcra!hibiIoryconnccIions
cnsurcdIhaI on!yonc mcmbcr o! cachpair was acIivaIcdaI any onc
Iimc.xIcrna!rcin!orccmcnIwasconnccIcdIoIhcinIcrna!rcin!orcc-
mcnI ncura! ncIsIhrouhhxcdconnccIionsIo capIurcIhc !acIIhaI
anima!s do noI havc Io !cam Io rcspond Io cxIcma! sourccs o!p!ca
surc and pain. Simu!aIcd anima!s cquippcdwiIh Ihcsc !our ncuraI
ncIs wcrc ab!c Io masIcr somc Iasks, such as naviaIion Ihrouh a
mazc, IhaInorma!!yrcquircIraininIhrouhinsIrmcnIa!condiIion-
inin!aboraIoranima!s. ' U oIcrsimu!aIionsIhcposiIivcrcin!orcc-
mcnIccnIcrswcrc!urIhcrdi!!crcnIiaIcdsoIhaIIhcirouIpuIacIivaIion
paIIcms dcnoIcd a ivcn dcrcc o! huncr or IhirsI. This incrcscd
Ihc dcrcc Io which Ihc bchavior o! Ihc simu!aIcd oranisms was
NLUKL NL1b N MMMLlN MLMK
oa!-oricnIcd, scckin noI|usI posiIivc rcin!orccmcnI m cncra!buI
!oodwhcnIhcywcrchunryorwaIcrwhcnIhcywcrcIhirsIy.'
Oncc wc cmbody and siIuaIc ncura! ncIs wiIh Ihc capaciIy Io
rcconizc cndurin cnIiIics Ihc ncxI sIcp wou!d bc ivin Ihcm Ihc
abiiIy Io ana!yzc sccncs inwhich Ihosc cnIiIics inIcracI p!ayin Ihc
ro!cs o!acnIorpaIicnI. IaboraIory cvidcncc !orIhc capaciIyo!birds
and mamma!s Io pcr!orm sccnc ana!ysis is morc di!hcu!I Io obIain
bccausc o! our comp!cIc !ack o! acccss Io Ihcir phcnomcno!oy. U
somc cascs wc can compcnsaIc !or Iis !imiIaIion by Icachin an
anima! how Io cxprcss iIs cxpcricncc Ihrouh cxIcma! mcans. II is
wc!! known, !or insIancc, IhaI chimpanzccs can bc IauhI Io usc a
rc!aIivc!y!arc vocabu!aryin sin !anuac. Thcycan a!sobc IauhI
Io uscnoIhand csIurcsbuIco!orcdp!asIicob|ccIso!di!!crcnIshapcs
arbiIrari!y paircd wiIh rc!crcnIs, IhaI is, paircd in such a way IhaI
nciIhcr Ihc shapc nor Ihc co!or o!Ihc p!asIic ob|ccIbcars any rcscm-
b!ancc IoiIsrc!crcnI-abucIrian!c,!orinsIancc,associaIcdwiIhan
app!c. Chimpanzccs arc noI on!yab!c !o associaIc Ic p!asIic ob|ccIs
wiIhrca!cnIiIicsandacIionsbuIcana!soarrancIhcmin scqucnccs
IhaIivcusin!ormaIionabouIhowIhcymakcscnsco!asccnc. Givcn
Ihc p!asIic ob|ccIs associaIcd wiIh Ihc Iraincr, wiIh iIsc!!, wiIh Ihc
acIion Ioivc,andwiIhIhccnIiIyapp!c,!orcxamp!c, achimpan-
zcc canarranc IhcminwhaIsccmsIobc rcqucsIs!ikcTraincr Givc
App!c Chimp. Thc chimpanzcc can a!so bc Iraincd Io undcrsIand
rcqucsIs!ikcChimp GivcApp!cTraincr. ' Thcsc rcsu!IsdonoIshow
IhaIchimpanzccscan !cam Io usc!anuac bccausc Ihc scqucnccso!
p!asIicob|ccIsdonoIhavc a rca!synIax, IhaIis, IhcydonoIhavcIhc
combinaIoria! producIiviIy o! human words. BuI Ihc ouIcomc docs
showIhaIIhcycanp!acccnIiIics andacIionsinrc!aIionsIhaI arcsi-
nihcanIIoIhcm.IhcycanundcrsIandandmcmorizcsccncsinwhich
onc parIicipanI acIs as an acnI, anoIhcr as a paIicnI, andm which
ccrIain acIions !ink Ihc Iwo IocIhcr.
Givcn Ihc scarciIy o! cvidcncc abouI Ihc ncura! subsIraIum and
phcnomcno!oy undcr!yin Ihc capaciIy !or sccnc ana!ysis in ani-
ma!s, and abouI Ihc rc!aIcdabi!iIyIomcmorizc acIua!!y cxpcricnccd
sccncs, Ihcsub|ccImusIbcapproachcdindirccI!y.Wcwi!!hrsIdiscuss
Ihckindso!modc!sIhaIhavc bccncrcaIcd!orhumancpisodicmcm
ory, a casc inwhich wc do havc acccss Io Ihc phcnomcno!oy, and
II
IZ
N LbN N b MUL1N
Ihcnwork ourwaybackIo Ihc animaI casc. CurrcnI simuIaIions oI
human cpisodicmcmoryusc proposiIions, Ihcmcanin oIscnIcnccs
IhaI cxprcss [IruIy orIaIscIy) IacIs abouI Ihc worId. IroposiIionsarc
uscd ascomponcnIparIsoImorccompIcxwhoIcscaIIcdscrirrs, sIruc-
IurcdrcpcscnIaIionscapIurinIhcrcuIariIicsoIrouIincsiIuaIions.'
ScripIs canbc uscd, IorcxampIc, IorcprcscnI IhcrcuIariIicsinacIiv-
iIicsIikconIoarcsIauranI. Thcspacc oIpossibIc acIionsinrcsIau
ranI sccncs has a sIrucIurc-wiIh somc acIions bcin causaI
prccondiIionsIorIhc occurrcnccoIoIhcrs-asIrucIurcIhaIaIIowsus
IomakcinIcrcnccsIikc U IhccusIomcrdocsnoIIowwhaIIoodis
avaiIabIc shc musI askIorIhcmcnu or IIIhc cusIomcris hnishcd
caIin hc musI pay Ihc chcck. Thc hypoIhcsis bchind scripIs is IhaI
Ihis sIrucIurc musIbc capIurcdbyourmindsiIwcarcIosucccssIuIIy
and rouIincIy anIicipaIc acIion possibiIiIics usin our cpisodic mcm-
ory oIpasI visiIs Io rcsIauranIs.Thc rcsIauranI-oin scripI was crc-
aIcdby hrsIsubdividin Ihc possibiIiIy spaccinIoIypcs oIrcsIauranI
[IancyrcsIauranI, caIcIcria, IasI-Iood|oinI) sincc Ihc acIion opporIu-
niIicsin cachoIIhcsc IypcsisquiIcdiIcrcnI.ThcnIhc diIIcrcnIrcs-
IauranIIypcs[ caIIcdIracks)wcrcsubdividcdinIosccncsnIcrin,
Ordcrin, aIin, xiIin, cach oI which was dchncd by a
scqucncc oIsimpIc acIions. achIrackwas aIso ivcna scIoIcharac-
IcrisIicob|ccIs [IabIcs, mcnus, Iood) androIcs [ cusIomcr,waiIcr,cook,
cashicr) . ThcIypicaIcondiIions oIcnIryIoIhcsiIuaIion[hunry cus
Iomcr wiI moncy) aswcII as iIs IypicaI ouIcomc [ saIiaIcdcusIomcr
wiIhIcssmoncy) wcrc aIsoincIudcd
ScripIsandoIhcr sIrucIurcd rcprcscnIaIionsoI sccncs wcrc crcaIcd
inIhc symboicschooI oIAtihaIInIcIhcncc so iI is noI surprisin
IhaI Ihcrc is aIinuisIicbias buiII inIo Ihcm. SpccihcaIIy, Ihc conIcnI
oI cpisodic mcmory was assucd IobcIanuacIikc, an assumpIion
IhaI may bc dcIcndcd on a varicIy oI rounds. Wc may aruc, Ior
insIancc, IhaI human cxpcricncc is sIrucIurcd IinuisIicaIIy, a Ihcsis
IhaIhasIiIIccxpcrimcnIaIsupporIbuIIhaIhasbccnvcryintucnIiaI
in phiIosophicaI circIs in Ihc pasI 200 ycars. A wcakcr buI morc
dcIcnsibIc arumcnI wouId bc IhaI wc havc no acccss Io oIhcr pco-
pIcscpisodicmcmorics cxccpIIhrouhvcrbaI rcporIs. This isindccd
IrucbuIiImaybcarucdIhaIIhoscrcporIsarconIyapaIc shadowoI
Ihc morc or Icss vivid rc-cxpcricnccs consIiIuIin auIobioraphicaI
rccoIIccIions. FinaIIy, Ihis way oImodcIin cpisodic mcmory maybc
NLUKL NL1b N MMMLN MLMK
dcIcndcd inIcrms oI Ihc oaI oI a parIicuIar simuIaIion. Thc abovc
scripI,Ior cxampIc, was crcaIcd Io ivc simuIaIcd acnIs Ihc capaciIy
Io undcrsIand shorI sIorics Iakin pIacc in rcsIauranIs, Ihc acnIs
dispIayin Ihcir undcrsIandin by answcrin qucsIions abou

, or
paraphrasinpassacsIrom, Ihosc shorIsorics. Bccausc shorI sIorics
IypicaIIy Icavc ouI many dcIaiIs Ihc abiIiIy Ioparaphrasc Ihcm orIo
answcr qucsIionsabouIIhcminvoIvcsinIcrrinIhoscmissinpicccs
oIinIormaIionusinasanaidsomcIhinIiRc a scripI. GivcnIhcoaI
oI cndowin a simuIaIcdacnIwiIhIhiscmcrcnIcapaciIyIhc con-
ccpI oI a scripI maybc acccpIcdas bcinuscIuI Ior cxpIanaIions oI
cpisodicmcmoryaIIhcmcchanism-indcpcndcnIIcvcI.
ThchrsIsIcpIowardadapIinIhiscxpIanaIory sIraIcyIoIhc casc
oIavianormammaIianmcmoryinvoIvcscrcaIinancuraIncIimpIc-
mcnIaIion oI scripIs, onc IhaI dispIays Ihc samc capaciIics Io para-
phrasc and answcr qucsIions abouI shorI sIorics. UnIiRc Ihc scripIs
crcaIcdbyIhc symboIicschooIIhosc usinncuraIncIsdonoIhavcIo
bc handcraIIcdby Ihcir crcaIors. Ihc scripIs arc Icamcd by cxampIc
cxIracIcdIromsIaIisIicaI rcuIariIicsinacIuaI sIorics muchIikc muI-
IiIaycr pcrccpIrons cxIracI proIoIypcs Irom scnsory sIimuIaIion. In
oIhcrwords,unIikcIhcsymboIicvcrsionoIscripIsIhoscimpIcmcnIcd
wiIhncuraIncIarccmcrcnI.'ThcmoduIcsIhaIcomposcIhisimpIc-
mcnaIiondiIIcrinimporIanIwaysIromIhc ncuraIncIs discusscd so
Iar soadcscripIionoIIhcirdcsinwiIIbc ncccssary bcIorc cxpIainin
how Ihc cnIirc asscmbIy works. Thc Iwo novcI archiIccturcs arc
rcurrnr nurul nrs and s/f-crjunizinj murs. KccurrcnI ncuraI ncIs
arc Iikc muIIiIaycr pcrccpIrons aumcnIcd wiIh Iccdback conncc-
Iions. InrcuIarmuItiIaycrpcrccpIronsIhcrcisbackwardmovcmcnI
oIinIormaIionabouI Ihc dcrcc oIdiscrcpancybcIwccncurrcnI and
dcsircd ouIpuI paIIcms buIIhis IccdbackopcraIcs onIy durinIrain-
in. AIIcr Ihc wcihIs oI Ihc connccIions havc convcrcd on Ihcir
hnaI vaIucs Ihc ncuraI ncI ccascs Io usc any Iccdback. KccurrcnI
ncuraIncIs, onIhc oIhcrhand,uscIccdbackdurinacIuaI opcraIion.
Thc simpIcsI vcrsion adds Io a muIIiIaycr pcrccpIion an cxIra scI oI
uniIs, caIIcd conIcxI uniIs, Iinkcd Io Ihc hiddcn Iaycr by conncc
IionswithunchancabIcwcihIs and opcraIin aI IuII sIrcnIh. This
mcans IhaI Ihc conIcxI uniIs rcccivc an cxacI copy oI Ihc acIivaIion
paIIcrnIhaIIhc hiddcn Iaycr hadimmcdiaIcIy bcIorc iIs currcnI onc.
ThiscopyisIhcnIcdbackIoIhchiddcnIaycr [aIonwiIhIhcacIivaIion
IJ
I+
NLbN N bMUL1N
comin IromIhc inpuI uniIs) causin iIsprcscnIsIaIcIo bc rc!aIcd
Io iIs immcdiaIc pasI. Thc rcsu!I is an cmcrcnI capaciIy Io cxIracI
rcmrcrulrcjulurirics Irom scnsory sIimu!aIion, a capaciIy IhaI can bc
uscd, Iorcxamp!c,Ioana!yzc spokcnscnIcnccs andprcdicI Ihc ncxI
wordina scrics
Thc sccondIypc oI ncura! ncI nccdcd Io imp!cmcnI scripIs isIhc
sc!I-oranizin map. U iIs simp!csI vcrsion Ihis dcsin uscs on!y Iwo
!aycrs onc oIwhich, Ihc inpuI !aycr, IuncIions |usI !ikc B mu!Ii!aycr
pcrccpIrons. Thc sccond !aycr, Ihc map iIsc!I, is madc oI compuIin
uniIs arrancd ina spccihc spaIia! paIIcrn, suchasaIwo-dimcnsiona!
rccIanu!ararray. This dcsin is said Io sc!I-oranizc bccausc un!ikc
mu!Ii!aycrpcrccpIrons iIis noI ivcn a IarcI ouIpuI durin Irainin
orIhc dcrcc oI mismaIch IhaI iIs currcnI ouIpuI has rc!aIivc Io Ihc
IarcI. InoIhcr words, IhcIrainin oIa sc!I-oranizin map isunsu-
pcrviscd.ThcwayinwhichIhcIraininprocccdsisasIo!!ows.vcry
uniIinIhcinpuI!aycrisconnccIcdIocvcryuniIinIhcmap!aycr, Ihc
wcihIs oI Ihc connccIions scI Io random va!ucs aI Ihc bcinnin
oI Ihc proccss. An acIivaIion paIIcrn Irom a scI oIIrainin cxamp!cs
is Ihcn prcscnIcd IoIhcinpuI!aycr. Bccausc anacIivaIionpaIIcmis
simp!y a !isI oI inIcnsiIics IhaI can bc cxprcsscd as numbcrs, and a
conhuraIionoIwcihIsincachscIoIconnccIionsisa!isIoIsIrcnIhs
IhaI can a!so bc numcrica!!y cxprcsscd, Ihc acIivaIionpaIIcm oIIhc
inpuI uniIs canbc comparcd IoriIs dcrcc oIsimi!ariIyIoIhcwcihI
conhuraIionoIcachscIoIconnccIions IoIhcmap!aycr.AIIcrsIimu-
!aIin Ihc inpuI !aycr a comparison ispcrIormcdIor cach scI oI con-
nccIionsandIhcwcihIconhuraIionIhaIhappcnsIobcmorc simi!ar
Io Ihc inpuI paIIcrn is scIccIcd. ThaI is, aI Ihc sIarI oI Ihc Irainin
whaIcvcr simi!ariIy Ihcrc may bc is comp!cIc!y by chanccbuIiIncv-
crIhcIcss aI!ows us Io pick a parIicu!ar uniIinIhc map!aycr as Ihc
winncr."
AIIcr sc!ccIin Ihc winncr Ihc wcihI conhuraIion oI Ihc scI oI
connccIions IoIhcwinninmapuniI is chancdsoIhaI iIrcscmb!cs
Ihc inpuI acIivaIionpaIIcrna!iII!cbiI morc. ThcconnccIionsoIIhc
spaIia! ncihbor oIIhaImapuniIarc a!so ad|usIcdIobccomc morc
!ikcIhcinpuI, IhcdcrccoIad|usImcnIdiminishinIhcIurIhcraway
IhcmapuniIsarcIromIhcwinncr.ThcnIhcproccssisrcpcaIcdwiIh
Ihc samc inpuI paIIcrn, Ihc winncr now bcin Ihc samc map uniI as
IhcprcviousiIcraIionsinccIhcwcihIsoIiIsconnccIionshavca!rcady
NLUKL NL1b N MMMLN MLMK
bccnmadcmorcsimi!arIoIhcinpuI.ThcwcihIsoIIhcconnccIions
IoIhisuniIarc madc cvcnmorcsimi!aras arc Ihosc oIiIsncihbors
cxccpI IhaI nowIhc sizc oIIhcncihborhood is dccrcscd. Thispro-
ccssisrcpcaIcdunIi! Ihcsizc oIIhcncihborhood hasbccnrcduccd
Ioasin!cmapuniI.AIIhaIpoinIIhcconhuraIionoIwcihIsassoci-
aIcdwiIhIhcwininmapuniIarca!mosIidcnIica!IoIhcinpuIacIi-
vaIionpaIIcrn,whi!cIhosc oIiIsncihborsarcvcry simi!arIoiI.This
mcansIhaI iIwc acIivaIcdIhc wiDnin uniI and aI!owcdIhis acIiva
Iion Io Iravc! backwards Ihrouh iIs wcihIcd connccIions iI wou!d
rccrcaIc Ihc oriina! paIIcrn aI IhcinpuI!aycr. And iIa ncihborin
uniI was acIivaIcd iI wou!d rccrcaIc a paIIcrn simi!ar Io Ihc oriina!
onc. Throuh Ihs unsupcrviscd Irainin proccdurc Ihc acIivaIion
paIIcrninIhcinpuI!aycr iscIIccIivcIy sIorcd"inarcionoIIhcmap
!aycr, or morc cxacI!y, Ihc mcans Io rcproducc iI arc sIorcd in Ihc
conhuraIionoIwcihIsbcIwccnIhcwinninmapuniIandIhcinpuI
!aycr. ThisIraininproccdurccanbcrcpcaIcdmany IimcsIordiIIcr-
cnIinpuIpaIIcrns cach oncsIorcd" inadiIIcrcnI rcionoIIhcmap.
WhcndiscussinIhcIaccrcconiIionncura!ncIabovcwc sawIhaI
simi!arIaccswcrcmappcdinIoncarbyrcionsinIhcspaccoIpossib!c
acIivaIion paIIcrns oIIhc hiddcn uniIs, and IhaI Ihis IransIormaIion
oIsimi!ariIyrc!aIionsinIorc!aIios oIspaIia!proximiIywascrucia!Io
Ihcabi!iIyoIIhcncura!ncIIorcconizcob| ccIs.Asimi!arcIIccIIakcs
p!acc in sc!I-oranizin maps cxccpI IhaI in Ihis casc Ihc simi!ariIy
bcIwccn inpuIs is mappcd inIo proximIy rc!aIions in Ihc map !aycr
iIsc!I. U oIhcr words, Ihc rc!aIions oI proximiIy arc now bcIwccn
rcionsoaconcrcIcacIua!spaccnoIanabsIracIpossibiIiIyspacc.This
chancs Ihc dcrccIowhichIhc sIorcd"disIribuIcdrcprcscnIaIionis
acccssib!c by oIhcrncuraI ncIs. Thc proIoIypcs IhaI amu!Ii!aycrpcr-
ccpIioncxIracIsarcavai!ab!con!yIoiIscII, a!imiIcdacccssIhaImakcs
Ihcm uscIu! Io modcI proccdura! mcmory buI noI dcc!araIivc mcm-
ory, whcIhcr cpisodic or scmanIic. BuI oncc Ihcsc proIoIypcs arc
madc cxp!iciI!y spaIia! Ihcy bccomc pub!ic!y avai!ab!c. oIhr ncura!
ncIs IhaI can acIivaIc Ihc winnn map uniI in Ihc sc!I-oranizin
map and IhaI can rcad Ihc rcsu!Iin acIivaIion paIIcm aI iIs inpuI
!aycr wi!! bc ab!c Io rcIricvc Ihc disIribuIcd rcprcscnIaIion. U addi-
Iion, IhcsIorcd"disIribuIcdrcprcscnIaIionismadcsimp!crandmorc
compacIbccauscasc!I-oranizinmapcanpcrIormarcducIionoIIhc
numbcroIdimcnsions oIIhcpossibihIy spacc oIacIivaIionpaIIcrns.
Ib
I
NLbN N bMUL1N
U Ihc numbcr o!hiddcnuniIs in a ncura! ncI is, say, cihI, IhcniIs
possibi!iIy spacc has cihIdimcnsions.This numbcr canbc rcduccd Io
on!yIwowhcnIhc acIivaIionpaIcmispro|ccIcdinIoIhc Iwo spaIia!
dimcnsions o!Ihc map. Thc sc!!-oranizinmap iIsc!! sc!ccIs Ihc Iwo
dimcnsions IhaI havc Ihc mosI variabi!iIy, IhaI is, Ihc Iwo IhaI arc
mosI !ikc!y Io bc rc!cvanI !or Ihc abi!iIy Io rcconizc paIIcrns. Thc
oIhcrsixdimcnsionsarcapproximaIcdbyIhcwayinwhichdi!!crcnI
disIribuIcd rcprcscnIaIions arc p!accd in Ihc map, a p!accmcnI IhaI
may dcvc!op an inIcrIwincd !orm rcscmb!in Ihc way in which a
onc-dimcnsiona!!racIacurvcIricsIoapproximaIcaIwo-dimcnsiona!
sur!accyrcpcaIcd!y!o!dinoniIsc!!.'"
Combinin rccurrcnI ncura! ncIs and sc!!-oranizin maps Io
imp!cmcnI scripIs has bccn donc in a simu!aIion ca!!cd isccrn. "
isccm uscs cihI scparaIc modu!cs, six o!which arc uscd !or pro-
ccssin proposiIions andIwo!ormcmorysIorac.ThchrsIproccssin
modu!cuscsarccurrcnIncura!ncIIoparscscnIcnccs,IhaIis,iIinpuIs
a Icmpora! scqucncc o! words [sub|ccI, vcrb, ob|ccI) and ouIpuIs a
scrics o! ro!cs. acnI, paIicnI, acIion pcr!ormcd, insIrumcnI uscd in
Ihc acIion. A sccond rccurrcnI ncura!ncI uscs Ihosc ro!cs as aninpuI
Io parsc sIorics, IhaI is, iI maps Ihc scricso! ro!cs inIo Ihc caIcorics
o!a scripI. ooIhcrproccssinmodu!cspcr!ormIhc rcvcrsc opcra-
Iions. onc Iakcs scripIs as inpuIs and ouIpuIs a scrics o! ro!cs, Ihc
oIhcr inpuIs Ihc ro!cs and ouIpuIs a scnIcncc. Fina!!y, Iwo addiIiona!
proccssin modu!cs arc uscd !or Ihc spccia!izcd Iasks o! answcrin
qucsIions and producin paraphrascs. ' Thc Iwo mcmory modu!cs
arc sc!!-oranizin maps uscd !or Ihc sIorac and sharino!disIrib-
uIcd rcprcscnIaIins, onc corrcspondin Io scmanIic mcmory Ihc
oIhcrIo cpisodicmcmory. Thc!ormcris nccdcdbccausc Ihcabi!iIyIo
undcrsIand shorI sIorics prcsupposcs a prcvious!y acquircd !cxicon
whi!c Ihc !aIIcr is whcrc Ihc sIrucIurc o! scripIs is mcmorizcd. Thc
cpisodic mcmory uscs scvcra! maps arrancdinIhrcc di!!crcnI!aycrs.
onc !aycr !or scripIs [oin Io a rcsIauranI, shoppin, Iravc!in) :
anoIhcr!or Iracks [ di!!crcnI p!accsIocaI orshop, di!!crcnI mcans o!
IransporIaIion

: and onc Io sIorc Ihc paIIcms corrcspondin Io Ihc

incumbcnIs o!ro!cs [whois doinIhc caIin, shoppin, orIravc!in:
whcrc Ihcsc acIiviIics arc pcr!ormcd: whaI is bcin caIcn orbouhI) .
Thcmu!Iip!cmaps !orm apyramida!sIrucIurcin Ihc scnsc IhaI cach
uniI o! Ihc Iop map is connccIcd Io a scparaIc Irack map, and cach
NLUKL NL1b N MMMLN MLMK
uniI o! a Irack map b connccIcd Io a !arcr map in which ro!cs arc
boundIoIhcirincumbcnIs.Thispyramida!sIrucIurcrcaI!y!aci!iIaIcs
Ihc cxIracIion o!Ihc inhcrcnI!y hicrarchica! sIrucIurc o!scripIs !rom
Ihcdi!!crcnIshorI sIoricsinc!udcdinIhcIraininscI.'Thc cndrcsu!I
is amodu!arncura!ncIIhaIcxhibiIsIhc cmcrcnIcapaciIyIoanswcr
qucsIionsabouIshorIsIorics, orparaphrascparIso!Ihcm,byin!crrin
missin picccs o!in!ormaIion usin iIs cpisodic mcmory.
Usin isccrn Io uidc phi!osophica! Ihinkin abouI mamma!ian
andavianmcmory invo!vcs simp!i!yin somc aspccIs o!iIs dcsin as
wc!!as comp!cxi!yin oIhcrs. Ononc hand, Ihc dcsinmusIbc madc
sim!crbccausc Ihcanima!capaciIicswcnccdIorcproduccarcnoIas
sophisIicaIcd as Ihosc IhaIisccrnsimu!aIcs. U wc Ihink o!scripIs as
inIcrna! modc!s IhaI cxp!ain acIua!!y !ivcd sccncs Ihcn Ihc nccdcd
simp!ihcaIionwou!dboi!downIIhis.whcrcashumanbcinsuscas
cxp!anaIoryrcsourccsnoIon!yphysica!causcsbuIa!sorcasons[ socia!
convcnIions, cu!Iura!va!ucs) andmoIivcs [pcrsona! oa!s) , birds and
mamma!s nccd on!y !ramc Ihcir cxp!anaIions in Icrms o! causcs
sincc Ihcy arc noI capab!c o! ascribin mcnIa! sIaIcs Io oIhcrs nor
undcrsIandhowsharcdconvcnIionsorva!ucsshapcbchavior.Whcn,
!or cxamp!c, a chimpanzcc uscs co!orcd shapcs Io makc a rcqucsI
[ aincrGivcApp!c Chimp") iIismcrc!yIryinIocausa!!ya!!ccIIhc
bchavioro!Ihc IraincrnoIIoint!ucncchis orhcr mind. OnIhc oIhcr
hand, wc wou!d havc Io makc isccrn morc comp!cx bccausc iniIs
currcnI!ormiI makcs usc o!!inuisIicin!ormaIionIhaIwou!dnoIbc
avai!ab!cIoanima!s. IIsinIcr!accIoIhccxIcrnaIwor!disimp!cmcnIcd
Ihrouhimacs o!!cIIcrsIrans!ormcdinIoacIivaIionpaIIcrns, IhaIis,
iI cxIracIsscripIs!romprinIcdIcxIsnoI!romscnsory cxpcricncc.This
rcaI!y !aci!iIaIcs Ihc Iask o! assinin Ihc ro!cs o! acnI andpaIicnI
Io ob| ccIs in a sccnc usin Ihc synIacIic and scmanIic sIrucIurc con-
Iaincd in prinIcd scnIcnccs, a sIrucIurc IhaI is noI auIomaIica!!y
providcdbyvisua! cxpcricncc.
ThcchancsncccssaryIobrcakIhc!inuisIicdcpcndcncco!scripIs
shou!d bc casicr Io imp!cmcnI in isccrn Ihan in Ihc oriina! sym-
bo!ic vcrsion bccauscIhc Iask o!makin scnsc o!siIuaIions is carric
ouI mIhc !ormcr wiIh cmcrcnI disIribuIcd rcprcscnIaIions uscd as
buiIdinb!ocksIocomposccqua!!ycmcrcnIhicrarchica!Iaxonomics.
Thosc disIribuIcd rcprcscnIaIions cou!d, !or cxamp!c, bc supp!icdby
avidcocamcra usinIhc !accrcconiIionncura!ncIdiscusscdabovc.
I
Id
NLbN N bMUL1N
BuI Ihc rca!prob!cmis noI so much Ihc kind o! in!ormaIio

uscd
asscnsoryinpuI asIhc!acIIhaIisccrnis cnIirc!y discmbodicd. Thc
!ack o! a body siIuaIcd in spacc !cavcs isccrnwiIhouI Ihc rcsourccs
nccdcd Io disIinuish whaI is rc!cvanI in an imac !or Ihc purposc
o!assininpcrccivcd ob|ccIsIoIhc ro!cs o!acnIandpaIicnI, or Io
undcrsIandIhc connccIionbcIwccnacausc andiIsc!!ccI.Aswc su-
csIcd in Ihcprcvious chapIcr posscssion o! a bodyivcs anima!s a
varicIy o!ways Io h!Icr ouI irrc!cvancics. FirsI o!a!!, !carnin aI Ihc
!cvc! o!Ihc spccics bui!dsinIo ananima!sbody a varicIyo!inhcriIcd
bchaviora! Icndcncics IhaI havc cvo!vcd m connccIion wiIh a rca!
cnvironmcnI and IhaI arc Ihcrc!orc rc!cvanI Io opcraIin m IhaI
cnvironmcnI. HabiIuaIion, inIurn, a!!owsanima!snoI Io spcnd co-
niIivc rcsourccs on sIimu!i IhaI rcquircd an acIivc rcsponsc prior
IocIIin uscd Io iI, in c!!ccIIrans!ormin a sIimu!usIhaIwas rc!c-
vanI inIo an irrc!cvanI onc. C!assica! condiIionin pcrmiIs inhcriIcd
rct!cxcs Iobc IricrcdnoIon!ybya sIimu!usIhaIhassurviva!va!uc
buI a!so by a prcvious!y irrc!cvanI sIimu!us Ihais ncvcrIhc!css prc-
dicIivc o!Ihcprcscncc o!IhchrsI.
Thcscsimp!c!ormso!proccdura!mcmoryrcprcscnIasscssmcnIso!
sinihcancc madc by Ihc spccics as a who!c ovcr manycncraIions,
asscssmcnIs IhaIcanbc!urIhcrmodu!aIcdor adapIcddurinan ani-
ma!s!i!cIimc.ToIhcsc irrc!cvancyh!Icrs a!rcady avai!ab!c Io insccIs
wc musI addIhc habiIsacquircdIhrouh insIrumcnIa! condiIionin
in !arcr anima!s, habiIs IhaI arc !css consIraincd by cncIics. Thc
t!cxibi!iIy o! habiIs comcs aI a pricc sincc as iI was arucd abovc
!carninIhrouhrcin!orccmcnIcan!cadIoIhc!ormaIiono!supcrsIi-
IioushabiIsbcarin norc!aIionIo Ihc capaciIyo! ancnvironmcnIIo
a!!ccI and bc a!!ccIcd by Ihc anima!. NcvcrIhc!css, whcn anima!s
!carn oodhabiIsIhcycannoI on!y auIomaIc a scrics o!acIions Iobc
cxccuIcd wiIhouI any conscious p!annin buI Ihcy can a!so !carn Io
anIicipaIcIhcrouIinc conscqucnccs o!IhciracIions. ConsidcrinIhaI
much o!whaImakcs up a scripI is rouIinc andrcpcIiIivc proccdura!
mcmory m Ihc !orm o! ood habiIs can considcrab!ynarrow down
Ihc ranc o! cxpcricncc IhaI musI bc dc!ibcraIc!y considcrcd by Ihc
anima! Io makc scnsc o! siIuaIions. U oIhcr words, an cmbodicd
acnIcan!cIdi!!crcnI!ormso!proccdura!mcmoryIakccarco!h!Icr-
inouI insinihcanIdcIai!swhi!ciI!ocuscsiIsaIIcnIion onIhc mosI
imporIanIaspccIso!acomp!cxsccnc.
NLUKL NL1b N MMMLlN MLMK
U addiIionIo ivin isccrnabodywcmusIa!sosiIuaIcIhaIbody
in spacc. Whcn simu!aIcd acnIs arc modc!cd wiIhouI a body Ihcy
arcIrcaIcdassmarIp!anncrsbuIdumbcxccuIioncrs,p!annin cvcry-
Ihin m Ihcir hcads prior Io pcr!ormin an acIion. Whcn Ihcy arc
simu!aIcd wiIhouI siIuaIin Ihcm in spacc Ihcy arc IrcaIcd as i!
Ihcyhadno in!ormaIionabouIIhcirimmcdiaIcsurroundins.'' This
in!ormaIionisrc!crrcdIo as indcxiculkncwlcdjc. Know!cdc abouI an
cxIcrna! ob|ccIs posiIion, !or cxamp!c, can bc cxprcsscdindcxica!!y,
Ihc ob|ccI is m !ronI o! Ihc acnI and Io iIs !c!I, or in a conIcxI-
indcpcndcnI!ormbyiviniIs!aIiIudcand!oniIudc.Whi!cindcxica!
know!cdc comcs !rom an acnIs siIuaIcdpoinI o!vicw-a round-
!cvc! vicw o! rca!iIy, as iI wcrc-Ic sourcc o! conIcxI-indcpcndcnI
know!cdc in simu!aIions is Ihc od!ikc acria! poinI o! vicw o! Ihc
dcsincr.'U IhcparIicu!arcasc o!scripIsindcxica!know!cdc isncc-
cssarybccausc aIIhc!owcsI!cvc!o!IhcIaxonomyIhcrcarc spccihc
ro!cs IhaI musIbcbound Io IhcirincumbcnIs, IhcincumbcnIsbcin
m mosI cascs cnIiIics known on!y indcxicaI!y. Apossib!c obsIac!c in
Ihisrcard is IhaIncura! ncIs arc noI ood aI so!vin bindin prob-
!cms,whcIhcrIhcprob!cmisbindinpronounsIopropcrnamcsma
!inuisIicconIcxI, variab!csIoIhcirva!ucsinamaIhcmaIica!conIcxI,
or ro!cs IoIhcirincumbcnIs masccnc-ana!ysis prob!cm. Onc wayo!
ovcrcominIhis !imiIaIion is sucsIcdbyIhc cxpcrimcnIa! obscrva-
Iion IhaI roups o! ncurons m rca! brains, cach o! which cxIracIs
di!!crcnI!caIurcs!romIhcsamcob|ccI,IcndIohrcinsynchronywiIh
onc anoIhcr whcn pcrccivin IhaI ob|ccI. U oIhcr words, Icmpora!
simu!IanciIy inIhc hrinbchavior o! ncurons can acI as a dynamic
bindcr. This insihI has a!rcady bccn succcss!u!!y cxp!oiIcd m somc
modu!arncura!ncIdcsinsIhaIpcr!ormscripI-!ikc in!crcnccs.'
vcn Ihouh wc arc sIi!! !ar !rom undcrsIandin how avian and
mamma!ianmcmory andpcrccpIionworkIhc idcas wc havc dcrivcd
!romsimu!aIionssucsIIhaIIhckcyinrcdicnImIhccxp!anaIiono!
Ihosc capaciIics conIinucs Io bc Ihc cxIracIion o! proIoIypcs !rom
scnsory cxposurc andIhcsIorac"o!Ihosc proIoIypcs mIhc!ormo!
disIribuIcd rcprcscnIaIions. U Ihc casc o! sccnc ana!ysis Ihc proIo-
IypcsmusIbc cxIracIcdnoIon!y!romsimi!ariIics inIhc spaIia!con-
huraIion o! Ihc dchnin !caIurcs o! scnscd ob| ccIs buI a!so !rom
paIIcrnsinIhcirIcmpora!bchavior, suchasIhcrcu!arco-occurrcncc
o!!incar causcs andIhcir c!!ccIs. Thcmorc cndurin!ormo!mcmory
IV
II
NLbN N b MUL1N
nccdcd Ior auIobioraphica! rcco!!ccIions, in Iurn, musI bc imp!c-
mcnIcdIhrouhIhcusc oIsc!I-oranizinmapsorsimi!ar dcsins in
whichIhcnumbcroIdimcnsionsoIIhcmapisnoIhxcdinadvancc.

`
Fina!!y, wcmusIprovidc Ihcsc ncura! ncIswiIhIhcncccssary irrc!c-
vancyh!IcrswiIhwhichcvo!uIionsupp!ics cmbodicdanima!s. Thcsc
arc parIicu!ar!y ncccssary Ior Ihc opcraIion oI sc!I-oranizin maps
bccausc Ihc mcchanism Ihrouh which Ihcy achicvc Ihc sIorac
oI disIribuIcd rcprcscnIaIions prcsupposcs a criIcrion oI simi!ariIy
providcd cxocnous!y, and simi!ariIy is a vcry prob!cmaIic rc!aIion
Iwoob|ccIsIhaIarcdissimi!arIocachoIhcrvisua!!ymaybcsimi!arin
IhciraudiIory, o!IacIory, orhapIicpropcrIics. vcnmorcprob!cmaIic
is Ihc IacI IhaI any Iwo cnIiIics may bc simiIar Io cach oIhcr in an
inhniIcnumbcroIpcrccpIua!!yirrc!cvanIways. incininhabiIanIs
oIp!ancI arIh, Ior insIancc, or inmovin s!owcr Ihan Ihc spccd oI
IhI.

vo!uIionhash!IcrcdouImanyirrc!cvanIdimcnsions oIsimi-
!ariIy Irom Ihc ncura! machincry uscd by anima! brains buI discm-
bodicdncura!ncIs donoIhavcacccssIoIhcscauIomaIich!Icrs.
II is Iimc now Io movc bcyond simu!aIions oI individua! acnIs
andIhcirindividua!mindsIoconsidcrcnIirccommuniIics oIacnIs.
Whi!c in Ihc Iormcr casc wc arc conccmcdwiIhhow an cmbodicd
and siIuaIcd acnI makcs scnsc oI Ihc opporIuniIics and risks pro-
vidcdIoiIbyiIs cnviromcnI, inIhc!aIIcrwcmusIaddIhc opporIu-
niIics and risks supp!icd by iIs inIcracIions wiIh mcmbcrs oI Ihc
communiIy oIwhichiI iIsc!Iisa componcnI parI. Sinccwcwi!!now
bcconccmcdwiIh!arcrwho!cs wiIhIhcirowncmcrcnIpropcrIics
Ihc dcIai!s oI howindividua! acnIs pcrccivc and mcmorizc can bc
IakcnIor ranIcd. Thismcans IhaIsimu!aIionscan uscInuisIica!!y
cxprcsscd ru!cs and proccdurcs Io cncraIc acnI bchavior wiIhouI
bcin Ihc qucsIion. Thc Io!!owin chapIcr wi!! Iack!c Ihis sub|ccI
sIarIin wiIh mu!IiacnI simu!aIions oI Ihc soca! bchavior oI our
c!oscsI rc!aIivcs inIhcanima! wor!d and movin onIo considcr how
Ihc morc comp!cx soca! bchavior oI our hunIcr-aIhcrcr anccsIors
cou!dhavc cmcrcd.
LHPJLLHJ
MJ|||dO|5 d0 ||I0d|O b||d|OIO5
AsIhcpsycho!oica!!iIcoIanima!sbccamcrichcr andIhcirbchavior
incrcasin!y dcIcrmincd by IhcirmcmbcrshipinacommuniIy Ihcir
inIcractionsprovidcdIhcmcdiumIorIhcriscoIancwkindoIcnIiIy,
soca! sIraIcics, and cncraIcd ncw possibi!iIy spaccs in which Ihc
sinu!ariIics dchncd scIs oI muIua!!y sIabi!izin sIraIcics. A!Ihouh
somc insccI spcccs havc an c!aboraIc soca! !iIc wiIhin co!onics or
hivcsIhcirinIcracIionsdonoIinvo!vcrcconiIionoIindividua!con-
spcchcsandIakcp!accbyindrccI communcaIon, IhaIis, by a!Icra-
IionoIIhcphysica!andchcmica!propcrIics oIasharcdcnviromcnI.
Nonkcys and apcs, on Ihc oIhcr hand, arc capab!c oI rcconizin
cachoIhcrnoIon!yasuniqucindividua!oranismsbuIa!soasbc!on-
int accrIainkinship roupandoccupyinaivcnrankina domi-
nancchicrarchy.Thc!ikc!ihoodIhaIayounmomcyorapcwi!!sIarI
a hhI wiIh anoIhcr, Ior insIancc, dcpcnds noI on!y onmcmorics oI
IhcouIcomcsoIrcviousarcssivccncounIcrsbuIa!so onIhcprcs-
cncc or abscncc oI an o!dcr rc!aIivc acIin as a soca! a!!y. OIhcr
cxamp!cs oI Ihc kind oI soca!prob!cm-so!vin pcrIormcdby Ihcsc
anima!sinc!udc coaIiIionIormaIion-a!owcrrankin anima!maybc
ab!c Io cha!!cnc a riva! iI hc or shc has prcvious!y csIab!ishcd an
a!!ianccwiIhahihcrrankinonc-aswc!!as rcconci!iaIorybchavior
IoundoIhcdamacoIprcviousarcssivc cncounIcrs. ' Thcobscrvcd
soca! capacIics oI conIcmporary non-human primaIcs providc us
wiIhIhcbcsIuidcIo spccu!aIc abouI howour!asIcommonanccs-
Iorscou!dhavcaIIccIcd[andbc aIIccIcdby) oncanoIhcr. Norc spc-
chca!!y,IhcyprovidcuswiIhinsihIinIoIhcmcchanismsbchindIhc
III
IIZ
NLbN N b MUL1N
cmcrcncc oI rcciproca! a!Iruism, a socia! inIcracIion invo!vin Ihc
cxchanc oI Iavors in which Ihcrc is a Icmpora! dc!ay bcIwccn
IhcprovisionoIascrviccandiIsrcciprocaIion.Kcciproca!a!Iruism,in
Iurn, mayhavcbccn Ihc poinI oIdcparIurc Ior a !onproccssIhaI
wou!d cvcnIuaI!y !cad Io Ihc cmcrcncc oI communa! so!idariIy in
ancicnIhunIcr-aIhcrcrcommuniIics.
So!idariIy in conIcmporary human communiIics acIs as a socia!
radicnIIhaIvaricsamon diIIcrcnIroupscndowinIhcmwiIhdiI-
IcrcnI capaciIics Iorpo!iIica!mobi!izaIion and oIhcr Iorms oI coI!cc-
Iivcsocia! acIion. ThcmcchanismoIcmcrcncc oIso!idariIy inIhcsc
communiIicsdcpcndsonIhccxisIcnccoIadcnscncIworkoIconncc-
Iions amon Ihcir mcmbcrs. In a communiIy in which cvcryonc
knows cvcryonc c!sc word oI mouIhIravc!sIasI, parIicu!ar!y iI Ihc
sub|ccIoIconvcrsaIionisIhcIransrcssionoIa!oca!norm. anunIu!-
h!!cd promisc. an unpaid bcI, an unrcciprocaIcd Iavor. This rapid
IransmissionoI!inuisIicinIormaIiona!!owsIhccommuniIyIoacI as
arcpuIaIion-sIoracmcchanism.IIinaddiIionridicu!c andosIracism
arc uscd Io coI!ccIivc!y punish Ihosc who dishonor commiImcnIs
IhcnIhccommuniIyhasIhc capaciIyIoacI asancnIorccmcnImcch-
anism. Thc qucsIionIo bc cxp!orcdinIhischapIcris howIhcIransi-
Iion bcIwccn Ihc rcciproca! a!Iruism oI Ihc common anccsIors oI
humansandchimpanzccsandIhc communa!so!idariIyoIIhccarIicsI
hunIcr-aIhcrcrs cou!d havc occurrcd wiIhouI Ihc usc oI IinuisIic
rcsourccs. Somc cxpcrIs bc!icvc IhaI Ihc hrsI cmcrcncc oI!anuac
[!oIIocncsis) Iookp!accbcIwccnIhirIyandhIIyIhousandycarsao
whcn buria! riIcs, sophisIicaIcd sIonc Ioo!s, and rcu!ar Iadc wcrc
a!rcady wc!I csIabIishcd. OIhcrs bc!icvc IhaI !oIIocncsis occurrcd
in an car!icr pcriod, Ihc midd!c pa!co!iIhic, IhaI bcan abouI Iwo
hundrcd andhIIy Ihousand ycars ao

iIhrway, by Ihc Iimc !an-

uac appcarcd Ihc anccsIors oI modcn humans had a!rcady spcnI
hundrcds oIIhousands oIycarsasmcmbcrsoIcommuniIicsinwhich
a dua! cconomy oIhunIin and aIhcrin invo!vcd ncw ro!c assin
mcnIs, pubIic moniIorin oI rc!aIions, and co!IccIivc cva!uaIion oI
rcsourcc a!IocaIions. U oIhcr words, a comp!cx socia! !iIc prccxisIcd
IhccmcrcnccoI!anuac.
Kcciproca! a!Iruism amon non-humanprimaIcsinvoIvcs sIraIcic
inIcracIions bcIwccn Iwo anima!s supp!yin onc anoIhcr wiIh an
MUL1LN1b N KM1L b1K1LLb
opporIuniIy Io coopcraIc-Io roomparIsoIIhcbodyIhaIarcinac-
ccssib!c wiIhouI hc!p, Ior cxamp!c-buI a!so wiIh Ihc risk oI bcin
chcaIcd, sincc Ihc dc!aybcIorcrcciprocaIionmcansIhaI Ihcrc isno
immcdiaIc assurancc Ihc scrvicc providcd wi!! bc rcpaid. This is an
cxamp!c oI ascciuldilcmmua socia!inIcracIionIhaI can havc co!!cc-
IivcbcnchIsbuIIhaI is cndancrcdbyIhcpossibi!iIyoIrcaIcrindi-
vidua! ains. U Ihc simp!csI socia! di!cmmas inIcracIions bcIwccn
pairsoIacnIsarcsIrucIurcdbyIwoopporIuniIics anIworisks. Ihc
IcmpIin opporIuniIy Io chcaI a coopcraIor ainin aI iIs cxpcnsc
[ ca!!cd IcmpIaIion" or simp!y T"): Ihc opporIuniIy oI muIua! ain
by coopcraIors [a rcward" or K"): Ihc risk oI muIua! chcaIin [ a
punishmcnI" orI") : andIhcrisk Ior a coopcraIoroIbcinchcaIcd
[bcinmadca suckcr" orS" ) . Thc hc!d oImaIhcmaIicsIhaIsIudics
sIraIcic inIcracIions modc!s Ihcm as amcs so Ihcsc opporIuniIics
and risks arc rcIcrrcd Io as Ihc payoIIs" oI Ihc amc. WhaI makcs
a sIraIcicinIcracIiona socia!di!cmmaisIhc ordcrin oIIhcpayoIIs.
U Ihc siIuaIionrcIcrrcd Io as Ihc Priscncr'sdilcmmuIhc payoIIs arc
arrancd !ikc Ihis. T > K > I > S. II K > T Ihcn Ihcrc is no conticI
bcIwccn individua! and coI!ccIivc ains so a!! di!cmmas musI havc
T > K. BuI Ihis parIicu!ar arrancmcnI a!so imp!ics IhaI Ihc worsI
IhinIhaI canhappcnisIobca coopcraIor IhaIis chcaIcd [I > S) . A
diIIcrcnIdi!cmmacanbc cncraIcdbyIhcarrancmcnIT>K> S>I.
This P caI!cd Chickcn"-aIIcr c amc in which Iwo drivcrs on a
co!Iisioncourscp!ayIo sccwhoswcrvcsouIoIIhcwayhrsI-bccausc
un!ikc Ihc prcvious casc Ihc worsI ouIcomc is whcnnciIhcr drivcr
swcrvcs and boIh crash [ S > I) . Iikc a!! opporIuniIics and risks Ihc
payoIIs oI sIraIcic inIcracIions sIrucIurc a spacc oI possibiIiIics, an
ob|ccIivcpossibiIiIy spacc sharcdby IhcinIcracIin acnIs.
InaddiIionIoIhisob|ccIivc spacc wcmusI a!so modc!Ihc sub| cc-
IivcspaccoIpossibi!iIics, IhaIis,IhcsIraIcicchoiccsavai!ab!c IoIhc
acnIs. U Ihc simp!csIcasc Ihcsc arc simp!yIhcchoiccs Io coopcraIc
or chcaI buI in morc comp!cx siIuaIions Ihc sub|ccIivc possibi!iIy
spacc conIains noI on!y choiccs buI sIraIcics. Bccausc in Ihc !aIIcr
casc Ihc va!uc oI onc acnIs sIraIcy dcpcnds on iIs rc!aIion Io Ihc
sIraIcics oIoIhcr acnIswhaI wc nccd Iocxp!orc isIhc srucIurc oI
aninIcrsub|ccIivcpossibi!iIy spacc. ThisinIcrsub| ccIivc spacc is sIruc-
Iurcdby sinu!ariIics caI!cdNushcuuilibriu dchnincombinaIions oI
IIJ
II+
NLbN N b MUL1N
sIraIcicsIhaIsIabi!izc onc anoIhcrbccausc noacnIcancIabcIIcr
payo!! by uni!aIcra!!y chanin Io a di!!crcnI sIraIcy. Whcn Ihc
opporIuniIics andrisksarcarrancdasinIhcIrisoncrsdi!cmmaand
IhcinIcracIionoccurs|usIonccIhcon!yNashcqui!ibriumis!orboIh
acnIsIochcaI.Ihcywou!dbcbcIIcro!!i!IhcycoopcraIcd,sinccK>
I, buInciIhcronccanchoosccoopcraIionuni!aIcra!!ywiIhouIriskin
bcin chcaIcd andcIIin S. Ihc inIcracIionis rcpcaIcdmanyIimcs
Ihcn muIua! coopcraIionbccomcs a Nash cqui!ibrium cxccpI in Ihc
vcry !asI cncounIcr in which muIua! chcaIin pays bcsI. BuI u Ihc
acnIs knowwhcnIhc !asI inIcracIionwi!! occur IhcniIpaysIochcaI
in Ihc prcvious onc as wc!!, and !or IhaI rcason in Ihc onc prior Io
IhaI onc, andsoona!!IhcwayIoIhcvcryhrsIonc. SoIhcinIcrcsIin
casc !rom Ihc poinI o! vicw o! modc!in Ihc cmcrcncc o! coopcra-
IioninprimaIccommuniIicsisarcpcaIcd [oriIcraIcd)sIraIcicinIcr-
acIion in which nciIhcr acnI knows whcn Ihc !asI cncounIcr wi!!
Iakc p!acc. This is ca!!cd Ihc indc]nirclyircrurcdIrisoncrs di!cmma, a
vcrsioncharacIcrizcdbyIhc !acIIhaIIhc shadowo!Ihc !uIurc" [Ihc
probabi!iIy o! !uIurc inIcracIions) hc!ps Io sIabi!izc coopcraIivc ouI-
comcs. Whi!c socia! di!cmmas invo!vin on!y a sin!c inIcracIion
rcprcscnI Ihc conBicI bcIwccn individua!and co!!ccIivc inIcrcsIs, Ihc
iIcraIcd vcrsion adds a Icmpora! dimcnsionIhaI capIurcs Ihc Icnsion
bcwccn shorI- and!on-Icrm inIcrcsIs.
Thc simp!csI sIraIcics arc Ihosc IhaI do noI Iakc hisIory inIo
accounI.ovcIhc sIraIcy o!a!ways coopcraIinandHawkIhcsIraI-
cyo!a!wayschcaIin.NcxIin comp!cxiIyarc sIraIcicsinwhichan
acnIs choiccsIakcinIoaccounIon!yIhc prcvious acIiono!Ihc oIhcr
acnI, coopcraIini!iIcoopcraIcdorchcaIini!iIchcaIcd, !orcxam-
p!c.I!wc addIoIhisIhaIinIhcvcryhrsIcncounIcr,whcnIhcrcis no
hisIoryonwhichIobasc a choicc, anacnIa!waycoopcraIcswc cI
Ihc!amoussIraIcyknownas 1ir-]cr-rur,winncro!IhchrsIsimu!aIcd
compcIiIion bcIwccn sIraIcics. NcxI in comp!cxiIy arc sIraIcics
IhaIinvo!vcrcmcbcrinnoI on!yIhc oIhcr acnIsprcvious choicc
buI a!so acnIs own prcvious choicc. A cxamp!c wou!d bc a
sIraIcyinsIrucin an acnI IocoopcraIcinIhc ncxIcncounIcruiIs
prcvious copcraIionwas rcciprocaIcd: Io shi!Iback Io coopcraIc i!
boIh prcvious!y chcaIcd: Io chcaI aain u iI chcaIcdwhiIc Ihc oIhcr
onc coopcraIcd: and hna!!y, Io shi!I back Io chcaI i! iI coopcraIcd
whi!c Ihc oIhcr onc chcaIcd. This sIraIcy is rc!crrcd Io as Puvlcv
MUL1LN1b N KM1L b1K1LLb
[orWinSIay, Ioosc-Shi!I) anddi!!crs !rom TiI!or-IaI in IhaI iIIakcs
advanIaco!uncondiIiona!coopcraIors[ovcs) . Norccomp!cxsIraI-
cics can bc cncraIcd by incrcasin Ihc amounI o! mcmory IhaI
acnIs musI brin Io bcar rcmcmbcrin, !or insIancc, noI |usI Ihc
prcviousacIionbuIIhcprcviousIwoacIions. Acxamp!c wou!dbca
morc !orivin" vcrsion o! TI-!or-IaI [ ca!!cd TiI-!or-Iwo-IaIs") in
whichanacnImusIbc chcaIcdIwicc inarowbc!orc chcaIinback.
ThcscidcascanbcuscdIomodc!Ihcsocia!sIraIcicso!non-human
primaIcsinIhc casc o! dyadicinIcracIions, suchas IhcrcciprocaIion
o!roominor!oodsharinbcIwccnIwoanima!s,andon!ywhcnIhc
inIcracIinpair is noI rc!aIcd bykinship, sincc kindrcd anima!s may
bccncIia!!yprcdisposcd Iowrd coopcraIion. Somc Iriadic inIcrac
Iions, !iRc sIarIina hhI in Ihc prcscncc o! an a!!y, maybc brokcn
downinIo Iwo dyadic oncs. Ihc inIcracIion wiIhIhc a!!y, who may
havcbccn roomcd inadvancc andiIsbcinprcscnI aI Ihc hhIisiIs
way o!rcciprocaIin, andIhc aonisIicinIcracIioniIsc!!, a riIua!izcd
conIcsI in which opponcnIs asscss cach oIhcrs sIrcnIh. In Ihis con-
IcsI usin Ihc Hawk sIraIcymcans Io csca!aIc Ihc !cvc! o!vio!cncc
unIi! in|urcd whi!c usin ovc mcans Io makc arcssivc disp!ays
[ such as bcarin oncs IccIh) unIi! csca!aIion sIarIs Ihcn wiIhdraw.
U addiIion Io rcsIricIin Ihc app!icaIion o! Ihc modc! Io pairs o!
unrcIaIcd anima!s Ihcrc is Ihc qucsIion o!whcIhcr non-human pri-
maIcs can in!acI ana!yzc cach oIhcrs bchavior and basc Ihcir own
rcsponscs on Ihc rcsu!Is o! Ihosc ana!yscs. vidcncc in !avor o! Ihis
hypoIhcsiscomcs!romcxpcrimcnIsusindyadso!unrc!aIcdIamarin
monkcys p!accd ina siIuaIioninwhichonc o!Ihcmhas an opporIu-
niIyIo providc !ood !or Ihc oIhcrwiIhouIiIsc!!cIIinany rcward.
InonccxpcrimcnIsomcIamarinmonkcyswcrcprcvious!yIraincdIo
usc Ihc Hawk and ovc sIraIcics-somc wou!d ncvcr pu!!Ihc !cvcr
IhaI providcd Ihc oIhcr onc wiIh !ood whi!c oIhcrs a!ways pu!!cd
iI-and Ihcn Ihcy wcrc paircd wiIhunIraincd monkcys. Thc rcsu!Is
wcrc c!car. Ihc unIraincdmonRcys pu!Icd sinihcanI!y morc !orIhc
ovcs Ihan!orIhc Hawks.
Thisrcsu!I, howcvcr, docsnoIncccssari!yimp!yIhaIIhc unIraincd
monkcys wcrc rcciprocaIin pasI bchavior IhaI was mcnIa!!y ana-
!yzcd and dcIcrmincd Iobc a!IruisIic, so a di!!crcnI cxpcrimcnI was
carricd ouI Io ru!c ouI Ihc possibi!iIy IhaI mcrc saIiaIion was Ihc
moIivaIin !acIor. o unIraincd monkcys wcrc p!accd in Ihc abovc
IIb
II
N LbN N bMUL1N
situation but aItcrnatin in roIcs, cach havin scvcraI chanccs to
dcIivcr Iood Ior thc othcr without immcdatc bcnchts to itscII. Thc
ratcatwhichthcycoopcratcdin thcscintcractionswasrccordcdand
thcn thcy wcrc madctointcractin asituationinwhichcachpuII oI
thcIcvcr rcsuItcd in Iood Iorboth oIthcm. FinaIIy, thcycnacd m
anothcr scssion oI aItcmatin aItruistic Iood cxchanc. U thcir ratc
oI coopcration dcpcndcd on mcrc satiation thcn thc intcrvcnin
situation in which both ot Iood shouId havc incrcascd thc ratc oI
coopcration. U onthc contrary, thc ratc dcpcndcd onthcirmcmory
oI thc othcrs past bchavior thcn thc intcrvcnin situation shouId
havc had no cIIcct. Thc rcsuIts wcrc compatibIc with thc Iattcr
hypothcsis thcmonkcyssccmcdtobcrcspondntopriorintcntionaI
acts oIIood ivinbythcirconspccihcs.'FinaIIy, thcrc isthc qucstion
oI whcthcr monkcys and apcs havc thc conitivc capacity to usc
bchavioraI stratcics IkcTit-Ior-tatorIavIov aspart oIa Iarcr com-
munity in which cach animaI must rcmcmbcr thc Iast outcomc oI
cvcry intcraction with othcr mcmbcrs, IitcraIIy kccpin scorc oI aII
its stratcic intcractions. Iaboratory studcs indcatc that onIy chim-
panzccs havc thc conitivc sophistication to do this whiIc othcr
spccics maybcusin simpIcrbchavioraI stratcics, such asmirrorin
cach othcrs bchavior orIavorin thosc with whom thcy spcnd timc
in cIosc proximity.
Givcn that aII wc nccd in this chaptcr is a way oI modcIin thc
sociaI stratcics oI thc common anccstors oI humans and chimpan-
zccs this cvidcncc is cnouh to aIIow us to assumc that thcy couId
mcmorizcpartncr-spccihccontincncics,punishchcatcrs,andpcrIorm
scorc kccpin. Thcncxt stcpistohndawaytocxpIorcthccmcrcnt
conscqucnccs Ior ancntirc communityoImany indchnitcIyitcratcd
dyadcintcractions.U particuIar, wcnccdtobcabIctotrackachan-
in composition oI stratcics ovcr many cncrations. U somc stratc-
ics comc to dominatc thc community in thc Ion run and iI thcsc
stratcicsinvoIvcsomcIormoIrcciprocaIaItruism [IikcTit-Ior-tator
IavIov) thcnsociaI coopcration can bc considcrcd a stabIc outcomc
and thus oncIkcIytohavc actuaIIyoccurrcdamon our anccstors.
Thc simpIcst way oI cxpIorin this qucstion is by coupIin modcIs
oIstratcicintcractiontocncticaIorithms. odinaivcnstratcy
into thc simuIatcd chromosomc oI a cnctic aIorithm is rcIativcIy
straihtIorwardbccauscthcchoiccsthatmakcupastratcyarcbinary
MUL1LN1b N K M1L b1K1LLb
[ coopcratc orchcat) asarcthcbtsthat composcthccncs. Iourbts
can odc Iorthc bnary rcsponscs to an acnts prcvous movc whIc
an addcdtwobtscantcIIthcacntwhattodonthcvcryhrstmovc."
Thc outcomc oI thc sImuIatons, on thc othcr hand, can bc qutc
compIcx bccausc thc htncss oI a partcuIar stratcy s Ircqucncy-
dcpcndcnt n thc scnscthat as a stratcybccomcs morc Ircqucnt n
a popuIatont can crcatc thc condtons Ior ts cvcntuaI dcmsc. U
othcrwords, thcrcIatvc succcssoIastratcydcpcndsnotonIyonts
ntrnscpropcrtcsbutonthcproportonoIacntsthatarcusniI as
wcII as onthc proporton oIacnts usn othcr stratccs.
Frcqucncy-dcpcndcncy crcatcs such compIcx dynamcs that thc
rcsuIts Irom onc smuIatonmay chanc whcn onc oI thc paramc-
tcrs-ntaIproportonsoIstratccs, ntaImxoIstratccs,numbcr
oIcncratons, mutaton ratc-s chancd. Ths s cIcarIyIIustratcd
bythcIatcoIthcvcryhrstsImuIatonoIthskndnwhchTt-Ior-tat
camctodomnatcthcpopuIatonaItcr50cncratonsnaIarcnum-
bcr oI runs. Ths rcsuIt was vcwcd as a watcrshcd n thc study oI
rccprocaI aItrusm andaIsocarrcdsomcmoraIcomIorbccausc, to
usc psychoIocaI mctaphors, thcwnnIn stratcy was ncc sncc
t aIways coopcratcd n thc hrst movc: t was Iorvn rcncwn
coopcratonIaIormcrchcatcrcoopcratcd: andt wasnotIabIctobc
auscdbccausc t was rctaIatory, punshn chcatcrsbychcatn.
'

U thc ornaIsmuIatonthcntaImxcontancdonIyHawk, ovc,

adTt-Ior-tatasaItcrnatvcs.HawkspcrIormcdwcIIathrstbytakn
advantac oI ovcs, but as thc ovcs bcan to dsappcar Irom thc
popuIaton so dd thc Hawks, Icavn onIy Tt-Ior-tat. But whcn thc
samc sImuIaton was aIIowcd to run Ioncr than 50 cncratons
Ircqucncy-dcpcndcnt tcndcnccs bccamcmanIcst. amutatontrans-
Iormcd onc copy oI Tt-Ior-tat nto ovc and bccausc aII Hawk
stratccswcrc onc ovcs wcrc abIctosIowIysprcadasmorc muta-
tonsaccumuIatcdatrandom:andonccthcovcstrtcyhadacqurcd
a crtcaI mass a mutaton producn a copy oI Hawk had aan a
chancc to ct a IoothoId and thcn nvadc thc popuIaton. 1 s now
wcIIknownthatIavIovwII do bcttcr nths sccnaro bccauscunIkc
Tt-Ior-tat iI can takc advantac oI ovcs and prcvcnt thcm Irom
crcatn thc condtons that Icad tothc sprcad oI Hawks. IavIov, on
thcothcrhand, docs not dowcIIaanstHawkssotosprcadthrouh
thcpopuIatont nccds to uscTt-Ior-tatasastcppnstonc. ' '
II
IId
NLbN N bMUL1lN
U Ihcsc simu!aIions Ihc bchaviora! sIraIcics wcrc modc!cd as i!
Ihcy wcrc inhcriIcd in a riid way, IhaI is, as i!Ihc acnIs did noI
!carndurinIhcir!i!cIimcs. Whcn!carnin is pcr!ormcdbyIhc spc-
cicsovcrmany cncraIions aNashcqui!ibriumrc!crs Io apopu!aIion
o! sIraIcics IhaI is sIab!c aainsI invasion !rom muIanI sIraIcics.
Thcsc arc rc!crrcd Io as cvcluricnur srublc srrurcjics, sIraIcics IhaI
pcr!orm succcss!u!!y whcn inIcracIin wiIh oIhcrs as wc!! as whcn
inIcracIinwiIhcopicso!Ihcmsc!vcs.U somc cascsiIis casyIocsIab-
!ishwhcIhcrasIrIcyis cvo!uIionarysIab!c. IhcHawksIraIcy, !or
insIancc,isc!car!ynoIsIab!cbccausc cvcnIhouhiIisoodaIcxp!oiI-
in oIhcrs iI docs noI do wc!! aainsI iIsc!!. BuI as Ihc casc o! TiI-
!or-IaI shows a sIraIcy prcvious!y IhouhI Iobc cvo!uIionary sIab!c
mayIunouInoIIobc soinIhc!onrunandIhiscomp!cxiIycaI!s!or
a comp!cmcnIary approach usin boIh compuIcr simu!aIions and
maIhcmaIica!ana!ysis. 'Whi!cmaIhcmaIica!ana!ysisrcvca!sIhcsin-
u!ariIicsIhaIsIrucIurc IhcinIcrsub|ccIivcpossibi!iIy spacccompuIcr
simu!aIions canacIua!!y!oI!owIhcpaIhIowardIhc sinu!ariIics. Wc
havca!rcadycncounIcrcdIhis comp!cmcnIariIybcIwccnIhcIwodi!-
!crcnIapproachcsincar!icrchapIcrsbuIinIhc casc o!simu!aIions o!
inIcracIin acnIs bcin ab!c Io Irack a hisIorica! Ira| ccIory is cvcn
morc imporIanI bccausc manyinIcrcsIin cvcnIs occurdurin rrun-
sicnrs, IhaIis,durinIhcpcriodo!Iimcbc!orcasinu!ariIyisrcachd.
U somc cascsa!rcqucncy-dcpcndcnIdynamicmayspcndmosIo!iIs
Iimcin!onIransicnIs.This imp!icsIhaIivin ancxp!anaIiono!Ihc
!on-Icrm conscqucnccs o!sIraIcicinIcracIioninvo!vcsshowinnoI
on!yIhaIaNashcqui!ibriumcxisIsandIhaIiIcanbcrcachcdbuIa!so
providinancsIimaIc o!Ihc Iimc sca!c ncccssaryIo rcachiI. '
Simu!aIions cana!socomp!cmcnImaIhcmaIica!ana!ysisbyhc!pin
Ioc!ari!ysomc conccpIua! assumpIions. Thcrc arc Iwo aspccIs o!Ihc
conccpI o! a Nash cqui!ibrium IhaI nccd c!arihcaIion. Onc has Io
dowiIhIhcmainIcnancc o!cqui!ibrium, IhaIis,wiIhIhcmcchanism
IhaI rcsIorcs sIabi!iIy a!Icran cxIcrna! shock, whi!c Ihc oIhcr rc!aIcs
IoIhc sc!ccIion o! onc Nashcqui!ibriumovcranoIhcrin Ihc casc o!
possibi!iIy spacs sIrucIurcd by mu!Iip!c sinu!ariIics. Thc sc!ccIion
prob!cmcan bc so!vcdIhrouha mcchanismIhaIpcr!orms a scarch
o! Ihc possibi!iIy spacc Io convcrc on onc o! Ihc scvcra! avai!ab!c
Nash cqu!ibria. Incar!icrchapIcrswchavc sccn IhaIbio!oica!cvo!u-
Iion consIiIuIcs a powcr!u! scarch mcchanism and Ihc simu!aIions
MUL1LN1b N KlM1L b1K1LLb
|usI discusscd shows how usc!u! Ihcy canbc Io hnc-Iunc our inIu-
iIionsabouIIhc sc!ccIionproccss. ' ThcmainIcnanccprob!cm, onIhc
oIhcr hand, nccds a di!!crcnI approach. In IradiIiona! maIhcmaIica!
modc!s Ihc prob!cm is so!vcd via assumpIion o! !u!! common
know!cdc on Ihc parI o! a!! acnIs. iI is noI cnouhIhaI a sIraIcy
is such IhaI no onc can swiIchuni!aIcra!!yIo a di!!crcnIonc andbc
bcIIcro!!buI,inaddiIion, cvcryacnImusIknowIhaIaI!oIhcracnIs
knowIhis.ThcassumpIiono!commonknow!cdc,howcvcr,dcmands
posIu!aIin coniIivc capaciIics IhaI Ihc anccsIors o! humans and
chimpanzccs cou!dnoIhavchad. SoIhcso!uIionIoIhcmainIcnancc
prob!cm musI bc souhI in Ihc dcvc!opmcnI o! ood habiIs IhaI
achicvc Ihc samc rcsu!I In Ihc casc o!ourprimaIc anccsIors Ihosc
ood habiIsmay havc bccn dcvc!opcd Ihrouhrcin!orccmcnI !carn-
ininIimcso!scarciIy, IimcsinwhichiIwasincvcryoncssc!!-inIcrcsI
Io coopcraIc by sharin !ood, buIbcinhabiIs as opposcd Io raIiona!
dccisions Ihcy cou!dhavc survivcdinIoIimcs o!p!cnIy. '
Usin compuIcr simu!aIions Io c!ari!y Ihc mainIcnancc prob!cm
imp!ics Ihc usc o!acnIsIhaI canacquirc habiIs inIhcir!i!cIimc and
Ihis, in Iurn, invo!vcs makin scvcra! chancs in Ihc assumpIions
bui!I inIo Ihc modc!s. InparIicu!ar, un!ikc Ihc casc o! discmbodicd
sIraIcicsinwhichIhcpayo!!scansimp!ybc assumcdIoacIasinccn-
Iivcs,rcin!orccmcnI!carninimpIicsrc!aIinIoIhc opporIuniIicsand
risks o! sIraIcicinIcracIion as rcwards and punishmcnIs. Thus, Ihc
hrsI chanc IhaI nccds Io c madc is Io inIroducc a cuIo!! poinI in
Ihc scrics o! payo!!s Io makc somc unambiuous!y rcwardin and
Ihc oIhcrs punishin. I!acin Ihc cuIo!!poinI in Ihc midd!c, soIhaI,
T< K < 0 <I < S, yic!dsIhc mosI inIcrcsIin rcsu!Is.'NcxI wcnccd
Io crcaIc ncura!ncIs IhaI can rc!aIc IoIhcsc payo!!s as posiIivc and
ncaIivc rcin!orccmcnI. As wc saw in Ihc prcvious chapIcr Ihis
invo!vcs a ccrIain dcrcc o! cmbodimcnI in Ihc !orm o! ncura! ncIs
pcr!orminmoIivaIiona! ro!cs, suchas simu!aIcdhuncrorIhirsI,buI
in Ihc simp!csI casc a!! wc nccdIo assumc is IhaI Ihc acnIs havc a
[non-cmcrcnI) Icndcncy Io avoid Ihc risk o! unrcciprocaIcd coop-
craIion.Thisboi!sdownIoIhc unprob!cmaIicassumpIionIhaIacnIs
dis!ikc bcin chcaIcd buI iI a!so carrics Ihc imp!icaIion IhaI cvcry
mcmbcr o! a dyad has an inccnIivc Io Iry Io cI Ihc oIhcr mcmbcr
Io coopcraIc. Thus whaI is nccdcd is a popuIaIion o! ncura! ncIs
IhaIIrain onc anoIhcronaconIinuousbasis asIhcyaIIcmpI Io cIiciI
IIV
IZ
NlLbN N b MUL1N
coopcraIion!romIhcirparIncrs. Ihc inpuI unIsrcccivc in!ormaIion
abouIIhcprcvious ouIcomc [or ouIcomcs) whi!casin!c ouIpuIuniI
yic!ds a binary choicc Io coopcraIc or chcaI in Ihc ncxI cncounIcr.
Thc wcihIs o! Ihc connccIions arc chancd on!y i! Ihc prcvious
inIcracIionwas ncaIivc, IhaI is, i!Ihcpayo!!wasIor S, andin pro-
porIionIohowncaIivc iIwas, morc!or S Ihan!orI. '
U onc simu!aIion an iniIia! popu!aIion o! ! 00 ncura! ncIs was
paircd aI random and Ihc rcpcaIcd inIcracIion bcIwccn dyads !o!-
!owcdIoobscrvcIhcscqucncco!chancsIhaI!cdIomuIua!coopcra-
Iion. cpcndin on Ihc sIraIcy d!!crcnI scqucnccs wcrc !o!!owcd.
Iwo Iav!ovsIraIcics, !orcxamp!c, cvcnIua!!y!ockcdinIo coopcra-
Iion !o!!owin Ihc scqucncc uni!aIcra! dc!ccIion-bi!aIcra! dc!ccIion-
bi!aIcra! coopcraIion. U wc imainc IhaI bio!oica! cvo!uIion had
a!rcady brouhI a communIyo!sIraIcisIs ncara Nash cqui!ibrium,
IhcaddIiono!!carninmaybcsccnas amcans !orIhccommunIyIo
rcIurn Io Ihc sinu!ariIy a!Icr a !orccd dcparIurc !rom iI. U oIhcr
words, Ihc habiI o!IryinIo c!iciI coopcraIion!rom aparIncr wou!d
bc a way o! so!vin Ihc mainIcnancc prob!cm. On thc oIhcr hand,
wiIhouIcoup!inncura!ncIsIocncIica!oriIhmsIhcabi!iIyIo!carn
by iIsc!! is noI a so!uIion Io Ihc prob!cm. This is i!!usIraIcd by Ihc
c!!ccIo!misIakcsonIhcco!!ccIivcouIcomc. rrorscanbcmadcwhcn
onc acnI incorrccI!y pcrccivcs" Ihc oIhcrs !asI movc or whcn iI
misIakcn!yrcspondsIoiIWhcnTI-!or-IaIcon!ronIsiIsc!!,!orcxam-
p!c, and onc o! Ihc dyadmcmbcrsmsIakcn!yrcsponds by chcaIin
a!Icr a coopcraIivc acI, Ihc sccond mcmbcr wi!!rcIa!iaIc bychcaIn
andIhc Iwowi!!!ock Ihcmsc!vcs inIo a !cudIhaI can on!ybc cndcd
byanoIhcrcrror.Iav!ov,onIhcoIhcrhand,isnoIon!yab!cIorccovcr
!roman crrora!Icr a sin!c round o! punishmcnI, buIiI can usc iIs
owncrrorsIo spoIovc-!ikcsIraIcicsoncc iIdscovcrsIhaIiI nccd
noI!carrcIa!iaIion.WhcnIhcsIraIcicsarc!u!!yinhcriIcdcrrorshc!p
sIabi!izc coopcraIion by a!!own pramaIic rcciprocaIors [Iav!ov) Io
sprcadIhrouhIhc popu!aIion.
'
' BuIwiIhouIinhcriIancc Ihisc!!ccI
dsappcars. Thc rcason is IhaI a popu!aIion o! !carnin dyads is, in
c!!ccI, scarchinIhcspacc o!possib!c sIraIcics byiIsc!!.U Ihccasc o!
ncura! ncIs IhaI Iakc inIo accounI Ihc !asI Iwo prcvious movcs Ihcrc
arc 2! possib!c sIraIcics and hcncc 2! squarcd [44! ) possib!c pairs.
Thc c!!ccI o! misIakcs is Io comp!icaIc Ihc scarch o! Ihis !arc spacc
andIomakciI hardcr!or coopcraIionIosprcad.

MUL1LN1b N KM1L b1K1LLb

ThcrcarcIwomorc!acIorsIoconsidcrwhcncva!uaIinIhcchanccs
IhaIrcciproca!a!Iruismcanbccomc csIab!ishcd n a popu!aIion. Ihc
ro!c o! spaIia! rc!aIions and o! kinship. Thc !ormcr is imporIanI
bccausc opporIuniIics !or nIcracIion arc a!!ccIcd by proximiIy, an
c!!ccIinorcd ina!!Ihc prcvious simu!aIions inwhichpairs o!sIraIc-
ics wcrc pickcd aI random Io nIcracI. Thc!aIIcr musI bc inc!udcd
dcspiIc Ihc !acI IhaI cncrosiIy amon rc!aIivcs is noI Ihc rcsu!I o!
a sIraIcicinIcracIion bccausc iI canacI as an cxocnous sourcc o!
coopcraIivc bchavior and int!ucncc Ihcovcra!!ouIcomc. Thc c!!ccI
o!boIh!acIors is Io crcaIc roups o!coopcraIorswiIhina!arcr com-
munIy wiIhin which rcciproca! a!Iruism can bccomc sIab!c. Onc
rc!aIivc!y simp!c way o! dcvc!opin inIuiIions abouI spaIia! c!!ccIs is
Io coup!c modc!s o! sIraIcic inIcracIion wiIhcc!!u!ar auIomaIa. As
discusscdinChapIcr2 Ihcscsimu!aIionsinvo!vcpopu!aIions o!hniIc
sIaIcauIomaIa IhaIinIcracIon!ywiIhIhcirncihborscapIurinprox-
imiIyc!!c

IsdirccI!y. ThcIask o!proramminanyo!IhcabovcsIraI-

cicsinIo ahnIc sIaIcauIomaIonisquiIcsIraihI!orward. Ihc inpuI
Io Ihc auIomaIon is Ihc ouIcomc o! Ihc prcvious cncounIcr [or
cncounIcrs) whi!ciIsouIpuIisIhcchoiccIo coopcraIc orchcaIinIhc
ncxI inIcracIion.'' Un!ikc Ihc c!assic modc! o! iIcraIcd Irisoncrs
i!cmmainwhichon!yIhcordcrin o!Ihcpayo!!s maIIcrs [T<K<I
< S) a crucia!!acIor in Ihc cc!!u!arauIomaIonvcrsionis howmuch
!arcr Ihcpayo!!!oruni!aIcra! dc!ccIion is rc!aIivcIoIhaI!ormuIua!
coopcraIion. For !ow cnouh va!ucs coopcraIion prcvai!s whi!c !or
!arc va!ucs chcaIin a!ways docs so iI is in Ihc inIcrmcdiaIc ranc
IhaI Ihc inIcrcsIin rcsu!Is !ic. Onc o! Ihcsc rcsu!Is is IhaI whi!c a
popu!aIiono!coopcraIinncihbors canbcinvadcduasin!cchcaIcr
ariscs, apopu!aIiono!chcaIcrs nccds a!arc cnouhc!usIcro!ncih-
borincoopcraIors Iobc invadcd.''
A!cssabsIracIwayo!cxp!orinIhcc!!ccIo!proximiIycanbcimp!c-
mcnIcd by makin Ihc choicc Io coopcraIc or chcaI Io bc abouI Ihc
usco!rcsourccs !ocaIcdinspacc. U oncsimu!aIioniI was!oundIhaI
roups o! coopcraIors can !orm i! Ihc disIribuIion o! !ood is paIchy
andu Ihcrc arc cosIs associaIcdwiIhmovin bcIwccn paIchcs, such
as an incrcascd risk o!prcdaIion. ThcinhcriIab!c coopcraIivc bchav-
ior in Ihis simu!aIion was rcsIrainI in Ihc consumpIion o! !ood Io
prcvcnIIhcovcrcxp!oiIaIionandcxhausIiono!Ihcrcsourcc, andIhc
ouIcomcwasIhaIroupso!coopcraIors cmcrcdbuIon!yas!onas
IZI
IZZ
N LbN N bMUL1N
Ihc!oodpaIchcs wcrc sma!! andwdc!y spaccdKnshp s anoIhcr
way o!cncraIncoopcraInroups!orIwod!!crcnIrcasons.FrsI,
bo!oca!cvo!uIoncanvcrscIoa!IrusIcbchavorbcIwccnc!osc!y
rc!aIcdanma!sby a mcchansm known as inclusivc]rncss. any cnc
prcdsposn an anima!Ioward a!Irusm canpropaaIc! Ihc anma!
Isc!!rcproduccs or ! c!osc rc!aIvcswIh whch I sharcs Ihosc cncs
rcproducc, so a!IrusIc csIurcs Ioward rc!aIvcs can havc sm!ar
cvo!uIonary c!!ccIs as sc!!-nIcrcsIcdbchavor. Sccond, knshp can
nt!ucnccIhc!ormaIono!roupsbccauscIhcbcnchIs o!kin-drccIcd
a!Irusmcanon!ybc cn| oycd!rc!aIvcssIaync!osc spaIa!proxim-
Iy.BoIho!Ihcscpossb!cc!!ccIshavcbccnIcsIcdnsmu!aIonsusn
cncIc a!orIhms coup!cd Io ncura! ncIs n whch cvo!uIon !u!!y
dcIcrmncdIhchna!wchIconhuraIon.
In onc smu!aIonapopu!aIon o! 00mu!IIaycrpcrccpIrons was
cvo!vcd,usnn!ormaIonabouIknshpasannpuIandIhcdccson
Iosharcorhoard!oodasanouIpuI. Thcpopu!aIonwasbrokcndown
nIo 20 roups o!rc!aIvcs andhIncss wasmcasurcdbyIhc amounI
o! !ood aIhcrcd durn a !!cImc. Thc ouIcomc con!ormcd Io Ihc
nc!usvc hIncss hypoIhcss. wh!c a!IrusIc bchavorIoward sIran-
crs waswccdcdouIa!IrusmbcIwccn rc!aIvcs was noI. Thcn a scc-
ondsimu!aIon waspcr!ormcdnwhchIhc!oodwasvcn a spaIa!
dsIrbuIon andIhc ncura! ncIs wcrc provdcdwIh addIona! unIs
Io a!!owIhcrbchavorIo bc shapcd byspaIa! posIon. !or youn
acnIs Iwo npuI unIs avc Ihc posIon o! Ihcr parcnIs | onc !or
an!c, Ihc oIhcr!or dsIancc)wh!c!or adu!I acnIs IhcnpuIwas
IhcposIono!!oodIcms.ForboIhacroupsaddIona!ouIpuIunIs
a!!owcdIhc acnIsIopcr!ormsmp!cmovcmcnIdccsons |Io movc
!orward, IoIurnrhI, orIoIurn!c!I) . IarcnIs hadIodccdcwhcIhcr
Io !ccd Ihcmsc!vcs, !ccd Ihcr cxsIn o!!sprn, or savc !ood !or
!uIurco!!sprnsnccIhcycou!drcproduccscvcra!IimcsdurnIhcr
!!c.Thcrcsu!IswcrcIhaIn youn acnIsaIcndcncyIosIayincvcr
cIoscrproximIyIoIhcrpaIIcrnscvo!vcd,andwIhIIhccmcrcncc
o! c!usIcrs o! coopcraIors. ForaduIIs Ihc rcsu!Iswcrc morc comp!cx
bccausc Ihcr choccs nvo!vcd compromscs. U parIcu!ar, ! Ihcy
wcrc Ioo a!IrusIc and dd noI su!hccnI!y !ccd Ihcmsc!vcs Ihcy had
shortcr !!c spans and !css ovcra!! o!!sprn: ! on Ihc conIrary, Ihcy
wcrcnoIa!IrusIccnouhIhcro!!sprnIcndcdIodc. Sorcproduc-
Ivc succcssbc!oncdIoIhoscIhaIsIrokcIhcrhIcompromsc.'
MULlLN1b N KlM1L b1K1LLb
IcIs pausc Io consdcr whaI has bccn arucd up Io Ihs ponI.
sIn!aboraIory cvdcnccsucsIsIhaIchmpanzccscannoI on!y
rcmcmbcrIhc ouIcomcs o!pasI cncounIcrs wIh ndvdua! mcmbcrs
o!IhcrcommunIy buI a!soana!yzc sccncsnvo!vn soca!nIcrac-
Ions Io dcIccI Icndcnccs Io coopcraIc or chcaI. Thc dscusson o!
cpsodcmcmorynIhcprcvouschapIcrmakcsIhccxsIcncco!Ihcsc
capacIcsraIhcrunsurprsnbuIancwIwsIhasbccnaddcdnIhs
onc. Ihc rcsponscs o! chmpanzccs n !uIurc nIcractons Icnd Io bc
corrc!aIcdwIh Ihcrcsu!Is o!Ihosc ana!yscsmakn Ihcrbchavoran
nsIancc o!asoca!sIraIcy. Usn Ihcsc capacIcs!orsoca!prob!cm-
so!vn as a udc Ihc qucsIon wc scI ouI Io answcr was whcIhcr
somc !orm o! rccproca! a!Irusm cou!d havc bccomc csIab!shcd n
IhccommunIcs o!Ihc !asIcommonanccsIors o!humansandchm-
panzccs.A!IhouhIhcconc!usonwc rcachcdwasnoIunambuous
I sccms p!ausb!c Io assumc IhaI scIs o! coopcraIvc sIraIccs o!
chanincomposIon cou!d havc bccomc csIab!shcd nrchuman
communIcsIhrouhaninIcracIono!scvcra!!acIors.cncIcprcds-
posIons coup!cdIooodhabIs:!ormaIon o!c!usIcrso! coopcraIors
!vinin c!oscproximIy: andIhcc!!ccIs o!nc!usvc hIncss. U oIhcr
words, sIab!c so!uIons Io soca! d!cmmas invo!vn ndchnIc!y
rcpcaIcd dyadc nIcractons cou!d havc bccn arrvcdaI co!!ccIvc!y
andIhcnmanIancdnacommunIy. ThcncxIsIcpsIouscIhcdcas
dcvc!opcd so!arIomodc!Ihc IransIon !rom ourprmaIcanccsIors
Io prc-!nusIc hunIcr-aIhcrcrs as a hsIorca! cvcnI IhaInvo!vcd
arrvinaIso!uIonsIoncwsoca!d!cmmas.
Oncway o!approachnIhc cmcrcncc o!coopcraIvchunIin |or
aIhcrin)sIomodc!IasaIransIon!romIhcIwo-pcrson Irsoncrs
d!cmma-in whch a who!c communIy maybcnvo!vcdbuI inIcr-
actons arc dyadc-IoIhc mu!I-pcrsonIrsoncrs d!cmma in whch
cach mcmbcr sIraIccaI!y nIcracIs wIh Ihc rcsI o!Ihc communIy.
AnoIhcr namc !or soca! d!cmmas nvo!vin a p!ura!Iy o! acnIs s
pub!c ood d!cmmas. U Ihc casc o! coopcraIvc hunIin Ihc pub!c
oodsIhc mcaI obIancdaIIhc cndo!IhchunI. apub!coodnvo!v-
in a cosI Io oopcraIors buI noI Io Ihosc who rc!rain !rom hunIin
buI cn|oyIs rcwards whcn Ihc mcaI s rcdsIrbuIcd. U Ihs casc Ihc
opportunIcs and rsks sIrucIurnIhcob|ccIvc spacc o!possb!Ics
bccomc morc comp!cx Io ca!cu!aIc sncc Ihcy musIbc a !uncIon o!
Ihcnumbcro!coopcraIors,IhccosIso!coopcraIn,andIhcdcrccIo
IZJ
IZ+
NLbN N b MUL1N
whichcoopcraIion amp!ihcs [ormu!Iip!ics) individua! c!!orIs. Thcsc
Ihrcc!acIorscanbcuscdIoca!cu!aIcIhcpub!icoodIhaIisproduccd,
a numbcrIhaI musI Ihcn bc dividcdbyIhc numbcr o! pcop!c inIo
which Ihc spoi!s arc rcdisIribuIcd. Sincc a chcaIcr cIs Ihc samc
amounIo!mcaIas somconcwhoparIicipaIcsinIhchunIiIspayo!!is
simp!yIhc rcsu!I o!IhaIdivision, whi!c Ihc payo!! !ora coopcraIor is
IhaIamounIminusIhccosIso!coopcraIin.ForIhcrcIobcadi!cmma
Ihc mu!Iip!icaIion!acIor musI bc !arcr Ihanonc -c!sc Ihcrc arc no
bcnchIsIo co!!aboraIivchunIinovcrand abovc Ihosc o!individua!s
hunIin a!onc-buI iI musI a!so bc !css IhanIhc numbcr o!pcrsons
inIo which Ihc spoi!s wi!I bc rcdisIribuIcd. Thc rcason is IhaI whi!c
chcaIcrs a!ways dobcIIcr IhancoopcraIorsbccauscIhcirpayo!!ncvcr
inc!udcscosIs,i!Ihcmu!Iip!icaIion!acIoris!arccnouhchcaIcrscan
improvcIhcirpayo!!byswiIchinIo coopcraIion. '`
A imporIanI di!!crcncc bcIwccn dyadic and communa! inIcrac-
Iions is IhaIsIraIcics !ikc TI!or-TaI arc noI parI o!Ihc inIcrsub| cc-
Iivc possibi!iIy spacc bccausc o!Ihc di!hcu!Iy o!!ocusin rcIa!iaIion.
A ood i!!usIraIion o! Ihis di!hcu!Iy is Ihc so-ca!!cd rrujcdy c]rhc
ccmmcns. a siIuaIion invo!vin a communiIy o! acnIs cxpoiIin a
commonrcsourccIhaIisvu!ncrab!cIodcp!cIionbyovcrc!oiIaIion.'
I! a sin!c chcaIcr ovcrcxp!oiIs and a !ormcr coopcraIor aIIcmpIs Io
rcIa!iaIcbya!so ovcrcxp!oiIin, asI-!or-IaIwou!ddo, IhisnoIon!y
un!air!ypcna!izcs Ihosc who rc!raincd !rom sc!hshbchaviorbuI iI
may a!so Iricr a wavc o! !urIhcr rcIa!iaIions in which cvcryonc
ovcrcxp!oiIs Ihc rcsourcc wiIh a prcdicIab!y Iraic ouIcomc. Thus,
viab!csIraIcicsmusI hnda wayo!rcp!acinIhc individua! punish
mcnI o! chcaIcrs by Ihc co!!ccIivc cn!orccmcnI o! coopcraIion. In
addiIion,whi!cinIhc casc o!dyadicinIcracIions iI is up IoIhc mcm-
bcrs o!a dyad Io dcIccIand rcmcmbcr dcparIurcs !rom coopcraIion,
whcn an cnIirc communiIy is invo!vcd such individua! mcmorics
arc noI cnouh. This mcans IhaI wc nccd a wayIo rcp!acc Ihc indi-
vidua!rcconiIiono!chcaIcrsbyIhc co!!ccIivc sIorac o!rcpuIaIions.
As mcnIioncd abovc Ihcsc Iwo mcchanisms arc imp!cmcnIcd in
conIcmporarycommuniIics by Ihc rapid sprcad o! word o!mouIh
abouIvio!aIions o!oca!norms andbyIhc usc o!osIracismandridi-
cu!c Iopunishvio!aIors. BuI in Ihc prcscnI conIcxI wc musI modc!
Ihc cmcrcncc o!boIh co!!ccIivc mcchanisms wiIhouIprcsupposin
Ihc cxisIcncc o!!anuac.
MUL1LN1b N K M1L b1K1LLb
Thcprob!cm o!sIorinrcpuIaIionscanbcso!vcdbyrcp!acinword
o!mouIhwiIhIhcsharcdcxpcricncc o!vio!aIionso!coopcraIion. For
cnIirccommuniIicsiIwou!dbcunrca!isIicIo assumc IhaIcvcryvio!a-
Iioncanbc wiIncsscdbya!!mcmbcrs. BuI i! a communiIyIcndcd Io
sorIiIsc!!ouIinIodisIincIroupsandi!mcmbcrs o!Ihcsc sma!!roups
Icndcd Io sIay in c!osc proximiIy Io onc anoIhcr Ihcn Ihc cpisodic
mcmorics o! roup mcmbcrs wou!d su!hcc Io kccp a rccord o! Ihc
pasIbchavior o!oIhcrroup mcmbcrs.As wc|usIsaw,boIhcco!oi-
ca!andcncIic!acIors canbrin abouI Ihc !ormaIiono! suchroups
so wc canIakc Ihosc rcsu!Is as indicaIions IhaIIhcrc is a so!uIionIo
Ihis prob!cm. Thcprob!cm o! co!!ccIivc cn!orccmcnI, on Ihc oIhcr
hand, is noI so casy Io so!vc bccausc o! Ihc cxisIcncc o! !rcqucncy-
dcpcndcnI c!!ccIs. Thiscanbci!!usIraIcdwiIhasimp!c cxamp!c. cIs
assumcIhccxisIcncco!ccrIaininhcriIcdIcndcncicsIhaIwou!dinuu-
cncc Ihc ouIcomc o! a pub!ic oods di!cmma, bchaviora!Icndcncics
!ikcvcnc!u!ncss, Ihc IcndcncyIopunishIhoscIhaIarc cauhI, and
bo!dncss,IhcIcndcncyIochcaIrc!aIivcIoIhcchancco!bcincauhI.
Wccancodc dcrccs o!bo!dncss andvcnc!u!ncssinIoIhcsimu!aIcd
chromosomcso!acncIica!oriIhandIrackIhcrcsu!Iinco!!ccIivc
dynamic. In onc simu!aIion a popu!aIion o! acnIs sIarIcd wiIh
cncsdcIcrminininIcrmcdiaIc !cvc!s o!boIhIcndcncics. A!Icra!cw
cncraIionsIhc dcrcc o!bo!dncss bcanIo!a!!bccausc cvcnwiIhouI
vcry hih vcnc!u!ncss bcin cauhI chaIin was cosI!y. As Ihc
proporIiono!bo!dacnIsdccrcascdcncs !orvcnc!u!ncssa!sobcan
Io disappcar bccausc wiIh on!y a !cw chcaIcrs cn!orccmcnI cosIs
bccomc a !iabi!iIy. BuI as Ihc avcrac dcrcc o!vcnc!u!ncss in Ihc
communiIy !c!! and wiIh iI Ihc cosIs o!bcin cauhI, bo!dncss madc
a comcback.'
Simu!aIions !ikc Ihis showIhaI Ihc cxisIcncc o! ccrIain pcrsona!
characIcrisIics is noI cnouh Io yic!d Ihc dcsircd ouIcomc bccausc
Ihc acIiviIy o!runishinj chcurcrsisirsc/furublicjccd.i!somc coopcra-
Iors rc!rain !rom punishin bccausc o! Ihc cosIs associaIcd wiIh iI
Ihcn Ihcy arc chcaIin Ihosc IhaI do punish. In oIhcr words, any
bchavior IhaIpromoIcs apub!icood is iIsc!! a sccond ordcrpub!ic
ood. This sucsIs IhaI Ihc so!uIion Io pub!ic ood d!cmmas musI
invo!vcboIhprimiIivcnorms-inhcriIcdorhabiIua!paIIcrnso!bchav-
ior !cadin Io Ihc punishcnI o! chcaIcrs-as wc!! as mcruncrms, Ihc
Icndcncy Io punish Ihosc IhaI donoIpunish.' This possib!c so!uIion
IZb
IZ
NlLbN N b MUL1lN
musIbccoup!cdwiIh IhaI IorchcaIcrdcIccIion,IhaIis,wiIhIhcIor-
maIionoIsma!!roupswiIhina!arcrcommuniIy. Inoncsimu!aIion,
Ior cxamp!c, acnIs wcrc maIchcd inIo roups usin Ihc norm do
noI choosc chcaIcrs asmcmbcrs" andIhc mcIanorm do noI choosc
Ihosc IhaI choosc chcaIcrs. " BoIh dcrcc oI coopcraIivcncss and
dcrcc oI vcncIu!ncss wcrc codcd inIo a cncIic a!oriIhm and a
popu!aIion was a!!owcd Io cvo!vc diIIcrcnI sIraIcics as Ihc acnIs
madc dccisions abouI roup IormaIion and choiccs Io coopcraIc or
chcaI. Thc ouIcomc dcpcndcd on Ihrcc condiIions aIIccIin roup
IormaIion. Thc hrsI condiIion rc!aIcd Io Ihc way inwmchmcmbcr-
shipwasdccidcd,IhaIis,whcIhcrauni!aIcra!choiccby anacnIwho
wanIcd Io |oin was cnouh or whcIhcr acccpIancc or rc|ccIion oI
mcmbcrship dcpcndcd a!so onIhcroup [muIua! choicc) . WiIh uni-
!aIcra! choicc Ihcrc wasnoroomIormcIanorms so Ihc ouIcomc was
prcdicIab!y IhaI !ow coopcraIors chcaIcd on hih coopcraIors. Thc
sccond condiIion invo!vcd Ihc possibi!iIy Ior vcncIu! mcmbcrs Io
sp!iI Irom a roup Io cxprcss Ihcir dissaIisIacIion wiIh Ihosc who
rcIuscd Io punish. WiIhouI Ihis possibi!iIy cncs Ior mh vcncIu!-
ncss cou!dnoIcIcsIab!ishcdand!owcoopcraIorsconIinucdIochcaI.
Fina!!y, IhcIhird condiIioninvo!vcd Ihc possibi!iIy IhaIacnIs who
may havc dcparIcd Irom a prcvous roup cou!d rccnIcr anoIhcr
roupa!!owin rcsi!icnI roups oIcoopcraIorsIo rccruiIncw mcm-
bcrsandrowaIIhccxpcnscoIoIhcrroups.ThcsIraIcycombinin
muIua! choicc, sp!iIIin, andrc|oinnpcrIormcdbcsI as a so!uIion Io
Ihcmu!Ii-pcrson Irisoncrs i!cmma.
"
Wcmayconc!udcIhaIwhcnIhccmcrcnccoImcIanorms a!!owcd
communiIicsIoso!vcIhcpub!icoodsdi!cmmainvo!vcdinco!!ccIivc
hunIinandaIhcrinIhcIransiIionIromanima!Iohumancommu-
niIics was comp!cIc. As mcnIioncd aI Ihc sIarI oIIhis chapIcr Ihcsc
novc! communiIics cmbodicda ncw rcsourcc radicnI, aradicnIoI
so!idariIy. From IhaI poinI on Ihc IurIhcr dcvc!opmcnI oI human
socia!iIybcanIo dcpcndnoIon!yonIhcinIcracIionsoIsma!! roups
wiIhin a cmmuniIy buI on Ihc inIcracIions bcIwccn disIincI com-
muniIics. Ihc ouIcomc oIwhichwas dcIcrmincd inparIby Ihc so!i-
dariIy radicnIs IhaI cach oI Ihcm cou!d cxp!oiI. Norc spccihca!!y,
whcn Ihc pracIiccs oI car!yhumancommuniIicsbcanIo inc!udcIhc
producIion oI sIonc Ioo!s, chanin Ihc ways inwmchhunIinand
aIhcrinwcrcconduccd,ncwNashcqui!ibriaIhaIwcrcimprovcmcnIs
MUL1LN1b N KlM1L b1K1LLb
ovcrIhc!doncsappcarcdinIhcirpossibi!iIyspaccs.ArcaIcrdcrcc
oI so!idariIy ina communiIy cou!dhavc madc Ihc IransiIion Io Ihc
ncwsinu!ariIysmooIhcrbyincrcasinIhc dcrccoIsoa!coordina-
Iion nccdcd Io movc Ihrouh Ihc possibi!iIy spacc whi!c sIayin in
Ihc coopcraIivc payoII rcion oIIhaI spacc. CommuniIics IhaI wcrc
ab!cIo succcssIu!!y makcIhaImovcwou!dhavc IhcrcIorc incrcascd
IhcirchanccoIsurviva!rc!aIivcIocommuniIicsIhaIIai!cdIo doso.
Thus, whaI IuIurc simu!aIions musI a!!ow us Io do is Io Irackmany
communiIics oI acnIs as Ihcy movc Irom onc Nash cqui!ibrium Io
anoIhcr in an cvcr chanin spacc oI possibi!iIics. In Ihc Io!!owin
chapIcr wc wi!! cxaminc onc oIIhc sourccs oIncw Nash cqui!ibria,
sIonc Ioo! Icchno!oy, as wc!! as a ncw socia! radicnI IhaI cmcrcs
IromIhcinIcracIionsbcIwccncommuniIics.IhcradicnIrcprcscnIcd
by IhcainsIromIrac.
IZ
IZd
LHPJL NNL
MJ|||dO|5 d0 bOO O LCOO0IC5
As communiIics o! car!y humans bcan Io sprcad ovcr Ihc p!ancI
IhciracIiviIics addcd Io iIs sur!acc onc morc !aycr o!comp!cxiIy. On
onc hand, Ihcirmcmbcrs, onwhosc bodicsancicnIcommuna!habiIs
hada!rcadyscdimcnIcd,acquircdIhccapaciIyIoIrans!orm sIonc inIo
manu!acIurcdIoo!s. OnIhc oIhcr, Ihc Ioo!sIhcmsc!vcsandIhc dcbris
produccd durin Ihcir manu!acIurc s!ow!y accumu!aIcd !cavin
bchind a rccord noI un!ikc IhaI o! !ossi!s cmbcddcd in co!oica!
sIraIa. NosI o! Ihc know!cdc wc havc abouI Ihc cu!Iurc o! Ihosc
car!ycommuniIics comcs !rom Ihosc dcposiIs o!sIonc arIi!acts. NcxI
IoIhcmarcco!oisIshavc a!so !oundIhc boncs o!anima!sIhaI dicd
aIdi!!crcnIacs, ivinus in!ormaIionabouIhunIin Icchniqucs, as
wc!! as ashcs !rom wood or bonc, providin us WU cvidcncc!orIhc
rouIinc usc o! hrc. NcvcrIhc!css, iI is Ihc sIonc arIi!acIs Ihcmsc!vcs
IhaIarc our richcsIsourcc o!daIa abouIIhc wayo!!i!c o!ouranccs-
Iors. Onc o!Ihcsa!csIconc!usionsIhaIcanbc drawn !rom Ihc sIraIa
Bwhich Ihcsc dcposiIs arc arrancd is Ihcir rcaI !oncviIy. Ihc o!d-
csIsIraIiraphicscqucncc (Ihc O!duvaiscqucncc) is abouI onc and a
ha!!mi!!ionycarso!d. ' Nucho!IhcconIcnIo!IhisandoIhcrscqucnccs
is rc!aIivc!y uni!orm buI as wc approachmorc rcccnIIimcsIhc Ioo!
and dcbris dcposiIs bcin Io divcrsi!y. Ihc variabi!iIy in Ioo! Iypcs
incrcascsinIhcmidd!cpa!co!iIhic (Iwohundrcdandh!IccnIhousand
Io!orIyIhousand ycars ao) and rcachcs a pcak in Ihc uppcrpa!co-
!iIhic (!orIyIhousandIoIcnIhousandycarsao) .
Too!s !rom Ihcsc morc rcccnI pcriods disp!ay a comp!cxiIy IhaI
bcars wiIncssIoIhcincrcascdsophisIicaIiono!Ihc manua! opcraIions
MUL1LN1b N b1NL L LLNMLb
uscd Io produccdIhcm. !rom Ihc o!dcsImcIhod o! pcrcusson, Ihc
dc!bcraIc brcakng o!asIoncinIopcccsbya!cwsIrongb!ows: IoIhc
morc rcccnIIcchniquco!rcIouchng, usng!ghIcrbuImorc !rcucnI
b!ows: Io Ihc usc o! prcssurc!!akng, rsI !or Ihc crcaIon o! sharp
b!adcs, Ihcn !or Ihc prccson shapng o! a Ioo! by conIro!Iing Ihc
Ihckncss o! Is d!!crcnI parIs, Ihc sharpcning o! Is cnds, and Ihc
c!mnaIon o! curvcs. U habIs promoIng coopcraIon consIIuIcd
Ihc rsI !orm o! human cu!Iura! conIcnI IransmIIcd across gcncra-
Ions, Ihcmanua!sk!!snvo!vcd n Ihc manu!acIurc o! sIonc Ioo!s
grcaI!ycnrchcdIhscu!Iura!hcrIagc.ThcdsIncIonbcIwccnhabIs
and sk!Is s !kc IhaI bcIwccn Icndcnccs and capacIcs. habIs arc
rcpcIIvc and !mIcd n varaIon wh!c sk!!s arc morc !!cxb!c and
opcn cndcd. Ikc a!I capacIcs Io a!!ccI sk!!s musI bc adapIcd Io a
capacIyIobc a!!ccIcd. pcrcussonb!ows musI am!or cracks or !rac-
Iurcs: cdgcs musI bc crcaIcdn argd sIonc capab!c o!ho!dng onIo
ashac:!!akngopcraIosmusIbcapp!cdIonc-graincdsIonc: pro-
ducIon o! !argc Ioo!s musI usc rocks o! Ihc rghI dimcnsons. BoIh
manua!skI!sandIhccomp!cxproccdurcsIowhchIhcygavcrscarc
ccrIanIy o!dcr Ihan spokcn !anguagc suggcsIng IhaI rhc hund muy
huvc ruujhrrhc mcurh rc srcuk, IhaI s, IhaI ordcrcd scrcs o! manua!
opcraIons may havc !ormcd Ihc background agansIwhichordcrcd
scrcs o!voca!zaIonsrsIcmcrgcd.
Varab!Iy n Ioo! and dcbrs dcposIs s cvdcncc noI onIy o! Ihc
progrcssvc d!!crcnIaIon o! sk!!s buI a!so o! cu!Iura! cnrchmcnI
wiIhncommunIcsandcvcno!conIacIbcIwccnd!!crcnIcommun-
Ics. Thc !aIIcr !o!!ows !rom Ihc !acI IhaI good qua!Iy sIonc was noI
a!waysavai!ab!c !oca!Iy so IhaIIhad Iobc mporIcd. Kc!ab!c mcIh-
ods Io dcIcrminc Ihc provcnancc o! d!!crcnI maIcra!s havc bccn
dcvc!opcdby archco!ogsIs whohavc shownIhaIagrcaIamounI o!
movcmcnI o! hgh-qua!iIy !!nI Iookp!acc n Ihc mdd!cpa!co!Ihc.
Kough!y, !a gvcnmaIcra!hasIravc!cd !css Ihanvc kiIomcIcrsIs
assumcd Io bc o! !oca! orgn: ! Ihc dsIancc s bcIwccn vc and
IwcnIy k!omcIcrs Is sadIobco!rcgona!orgn: and!IhcmaIcra!
has bccn IransporIcd ovcr disIanccs grcaIcr Ihan 30 kiIomcIcrs I s
dccmcd cxoIc or mporIcd.Wh!c n Ihc mdd!c pa!co!IhcIhc !on-
gcsI movcmcnIs sccmIobc o!Ihc ordcr o! a hundrcd kIomcIcrs by
Ihc uppcrpa!co!iIhcIhcyhavcncrcascd Io 400 k!omcIcrs.WhaIs
noI c!car swhcIhcrIhc !ong-disIancc movcmcnIo!!!nIand oIhcr
IZV
IJ
NLbN N bMUL1N
maIcria!s was cIIccIcd Ihrough somc Iorm oI Iradc or by anoIhcr
mcchanism. Oncpossibi!iIyisIhaI IhcmovcmcnIbcganwiIhaIrav-
cIng dc!cgaIion carryiDg an oIIcring IhaI was Ihcn cnIrusIcd Io a
varicIyoIinIcrmcdiarycommuniIics nonc oIwhichcvcr!cII iIs own
IcrriIory.TLs wou!dp!y IhaI somc Iorm oI coopcraIion bcIwccn
diIIcrcnI communiIics had bccomc sIab!c cnough Io supporI such a
mu!IisIagcdIransporImcchanism buI, as was argucd inIhcprcvious
chapIcr, IhcsIabi!iIy oIcoopcraIionL hardIoachicvcwiIhinasing!c
communiIy!cIa!oncbcIwccnscvcra!diIIcrcnIoncs. SoIradcrcmaiDs
a possibi!iIy cvcn iI iI invo!vcd on!y a vcry primiIivc Iorm oI barIcr
wiIhnoc!carconccpIionoIproIs.
vidcnccIromconIcmporaryhunIcr-gaIhcrcrsindicaIcsIhaIwiILin
communiIics Ihc on!yIypc oIsIab!c cxchangc isiDIhcIorm oIIrans
acIionsinIcrna!IokinsLipgroups.muIua!aidorgiIIsnoIimmcdiaIc!y
rcciprocaIcd.ThcinIcracIionsbcIwccnsuchgroupswiIhinaIribccan
bccomc !css pcrsona!buIiI is on!yinIransacIions bcIwccn diIIcrcnI
IribcsIhaI socia!disIancc iDcrcascs cnough Iorimpcrsona! cxchangc
Io cxisI.` This cvidcncc can scrvc on!y as a rough approximaIion Io
Ihink abouI midd!c pa!co!iIhic communiIics bccausc Iodays hunIcr-
gaIhcrcrsa!rcadyposscss spokcn!anguagc. BuIIhisis!css oIaImiIa-
IionIhaniI may sccm bccausc Ihc cxisIcncc oI a comp!cx !anguagc
hasncvcrbccnaprcrcquisiIcIorIradccvcniDrc!aIivc!yrcccnImcs.
inarcasoIiDIcnsc commcrcia! acIiviIy bcIwccn communiIics oI diI-
IcrcnI !inguisIic backgrounds simpIcd !anguagcs, ca!!cd Iradc pid-
gins, arc oIIcn crcaIcd in a rc!aIivc!y shorI Iimc and Ihcy disappcar
asIhc conIacI siIuaIion ccascs Io cxisI. Ior simp!cr Iorms oI barIcr a
Icw gcsIurcs and unaricu!aIcd sounds cou!d havc suIccd Io carry
ouI a IransacIion. Thcmain qucsIion Ihcn isnoI somuch abouI !in
guisIicsbuIabouIcconomics, IhaIis,abouIsiIuaIionsinwhichchoiccs
musIbcmadc duc IoIhcscarciIyoIrawmaIcria!sandIhca!IcrnaIivc
uscs Io which Ihcmay bc puI. ScarciIy is a conccpI IhaI musI bc
hand!cdcarcIuy bccausc iI is casy Io inIroducc cIhnoccnIric prc|u-
diccsinanana!ysis.JudgingconIcmporaryhunIcr-gaIhcrcrsassuIIcr-
ingIromrcsourccscarciIyas comparcdIoagricu!Iura!isIs,Iorcxamp!c,
ismisIakcn. Thc Iormcrmayhavc!css posscssions IhanIhc!aIIcrbuI
IhaIisacondiIionIhaIpcrIccI!yIsIhcirmigraIoryIIcsIy!c,posscss-
ingon!y asmuch asIhcycancarrywiIhIhcm.
MUL1LN1b N b1NL L LLNMLb
ThcconccpI oIscarciIy, IhcrcIorc, musIa!ways bcuscdrc!aIivc Io
spccicrcsourccs.ThcIypo!ogica!divcrsiIy oIIoo!siDmidd!cpa!co
!iILicdcposiIsandIhcdiIIcrcnIia!inIcnsiIywiIhwhichIhcIoo!swcrc
workcdiscvidcnccIhaIIhcirproduccrshadanawarcncssoIIhc!im-
iIcd avai!abi!iIy oI Ligh-qua!iIy maIcria!s and wcrc Iorccd Io makc
choiccsbcIwccnpossib!cuscs.iIIhcrawmaIcria!wasimporIcdiIwas
uscd Io crcaIc a widcr varicIy oI Ioo!s and Io crcaIc morc comp!cx
Iorms |such as doub!c-cdgcd Ioo!s) Ihan H iI was oI !oca! origin.'
InccnIivcs Io Iradc wcrc a!so crcaIcd by cnvironmcnIa! IacIors IhaI
!imiIcd Ihc mobi!iIy oIcar!y humans consIraining Ihcir acccss Io gco
!ogica!sourccs oIrawmaIcria!s.Thcmidd!c pa!co!iILic!asIcdso!ong
andwiIncsscd suchdramaIicc!imaIic!!ucIuaIions |IromIu!!Icmpcr-
aIc Io Iu!! g!acia!) IhaI in Ihc co!d pcriods such inccnIivcs Io Iradc
musI havc bccncommon. Iina!!y, Ihcrcwcrcp!cnIyoIdisIiDcI com
muniIicsIoprovidcIhcdcgrccoIsocia!disIanccncccssary Iorimpcr
sona!cxchangc. A!Ihougharchco!ogisIs!abc!simi!ardcposiIsoIsIonc
Ioo!s rccovcrcd Irom diIIcrcnI siIcswiIhnamcs!ic NousIcrian or
AurignacianIhcscIcrmsshou!dnoIbcIakcnIorcIcrIosc!I-idcnIicd
cIhnicgroupsoccupyiDg!argc IcrriIorics. KaIhcr, Ihc IransiIionIrom
Ihcmidd!cIoIhcuppcrpa!coIILiciDvo!vcdmanysma!!bandsiDhab-
iIiDgIavorab!c arcas wiIh!argc cmpIy zoncs bcIwccnIhcm, simi!ari-
Iics in Ioo! dcsign bcing Ihc rcsu!I oIcu!Iura! conIacI. II is p!ausib!c
IhaI somc oI Ihcsc conIacIs invo!vcd Iradc buI Ihis hypoIhcsis is
norma!!y rcsisIcd by anIhropo!ogisIs bccausc orIhodox cconomic
modc!s makc ccrIain assumpIions-pcrsona! sc!I-inIcrcsI, opIima!
dccision-making, cosI-Ircc inIormaIion-IhaI arc c!car!y iDadcquaIc
in Ihc conIcxI oI conIcmporary hunIcr-gaIhcrcrs !cI a!onc midd!c
pa!co!iIhichumans."
OnIhcoIhcrhand,scvcra!simu!aIionsoIbarIcrhavcbccncrcaIcd
IhaIdispcnscwiIhIhcscassumpIionsrcmovingaI!casIoncsourccoI
ob|ccIions. To Ihc cxIcnI IhaI Ihsc simu!aIions arc uscd Io modc!
hunIcr-gaIhcrcrsIhcagcnIsdcp!oycdinIhcmshou!dnoIbcconsid
crcdIobcpcrsonsbuIbands orIribcssinccIradc docsnoIIakp!acc
wiIhincommuniIicsbuIbcIwccnIhcm.Simu!aIcdagcnIscanbcuscd
Io modc! cnIirc communiIics bccausc, !ikc Ihcpcrsons composing
Ihcm, Ihcyarcindividua!cnIiIics.ThcIcrm iDdividua! has comcIo
bc uscd as synonymous wiIh pcrson buI IhaI is mis!cading. As an
IJI
IJZ
NLbN N bMUL1N
ad|ccIvc I can!cgImaIc!ybc uscdIo spcako!ndvdua! organza-
Ions, ndvdua! cIcs, ndvdua!naIon-sIaIcs, or any oIhcr cnIIy
IhaIssngu!arandhsIorca!!yunquc.AnoIhcrrcasonwhycommu-
nIcscanbcmodc!cdasagcnIssIhaIonccIhcyhavc acqurcd ccr-
gcnI propcrIcs o! Ihcr own, propcrIcs Ikc so!darIy or nIcma!
cu!Iura! homogcncIy, Ihcy can p!ay an cxp!anaIory ro!c ndcpcn-
dcnI!y o! Ihc pcrsons IhaI composc Ihcm. Whc Iwo counIcs
nIcracI, !or cxamp!c, Ihc acIua! nIcracIons may bc pcr!ormcd by
ndvdua! pcrsons buIIhc dcIa!s o!Ihcrpcrsona!nIcrcsIsandprc!-
crcnccs w!!bc cxp!anaIor!y rrc!cvanI ! any oIhcr Iwo communIy
mcmbcrs cou!d rcp!acc Ihcm and !cavc Ihc ouIcomc nvaranI. In
oIhcr words, H Ihc ouIcomc o! an nIcrcounIy nIcracIon s
noIa!Icrcdbychangcs n Ihc dcnIIy o!Ihc pcrsonsIhaIpcr!ormcd
I Ihcn Ihc dcnng!caIurcs o!Ihcrpcrsona!dcnIIycanbc !c!I ouI
o! ancxp!anaIon. Bccausc wc arc modc!ngndvdua!communIcs
noIons !kc IhaI o! a communIys sc!!-nIcrcsI or o! Is cu!Iura! and
maIcra!prc!crcnccsarcpcr!ccI!yvaId.Thc pcrsonscomposngIhcsc
communIcscansI!!bcconccvcdasposscssngIhcrownsc!!-nIcrcsI
buIoncIhaIs subordnaIcdIoIhaI o!Ihc cnIrcgroupwIhdcnIc
cosIsassocaIcdwIhIhcbrcakngo!IhaIbond.
ThusnwhaI!o!!ows IhcIcrm agcnIw!!rc!crIo ancnIrccom-
munIyandIhcIcrmsub|ccIvcIoIhcsharcdprc!crcnccscharacIcr-
zngIhaIcommunIy. IcIs rsI dcscrbc Ihc ob|ccIvc andsub|ccIvc
possbIIy spaccs nvo!vcd na IragnIcracIonbcIwccnIwo com-
munIcs. Thcsub|ccIvc spaccconIansa!!Ihc possb!c combnaIons
o! quanIIcs o! Ihc ob| ccIs Iobc Iradcd. U Ihc sp!c casc o!pa!co-
IIhc barIcr wc can assumc Ihcrc wou!dbc on!y Iwo such ob|ccIs.
hgh-quaIIy!!nIandparIa!!yworkcdsIoncIooIs.AIwo-dmcnsonaI
possbIIy spacc n whch cvcry ponI s a combnaIon o! a ccrIan
quanIIyo!!!nIandaccrIanquanIIyo!Ioo!scanbc uscdIocxprcss
Ihcprc!crcnccso!cachagcnI,Ihcsub|ccIvcprc!crcncc!oronco!Ihc
ob|ccIs n Icrms o! a numbcr o! unIs o!Ihc oIhcr ob|ccI. onc agcnI
maybcw!!ng Io!cIgo o!oncpccc o!hgh-quaIIy!!nIncxchangc
!orIwoparIa!!yworkcdIoo!s,!orcxamp!c,wh!canoIhcrmayprc!cr
ad!!crcnIcombnaIon.I!anagcnIcannoI choosc bcIwccn, say,Iwo
pcccs o! !!nI and Ihrcc Ioo!s bccausc boIh quanIIcs arc cqua!!y
dcsrabIc IhcnIhaIponInIhc possbIIyspaccsa combnaIonIhaI
makcsnod!!crcnccIoIhc agcnI. Jonng scvcra!suchponIsbya!nc
MUL1LN1b N b1NL L LLNMLb
dcncs an indi]crcncc curvc.
'

ach agcnI mayhavc scvcra! nd!!cr-

cncc curvcs d!!crng n dcsrab!Iy. Wh!c dcsrcs or prc!crcnccs
cannoIbccomparcdquanIIaIvc!y-wccannoI|udgc bycxacI!yhow
muchmorc onc ob|ccI s dcsrcdovcranoIhcr-wc can ncvcrIhc!css
arrangc prc!crcnccs n an ordcr mang rgorous | udgmcnIs abouI
an ob|ccIs grcaIcr or !csscr dcsrabIIy. Usng Ihcsc quaIIaIvc
|udgmcnIs Io rank many nd!!crcncc curvcs c!ds an nd!!crcncc
mapcapIurngIhcsIrucIurco!Ihcsub|ccIvcpossb!Iyspacc!orcach
agcnI.
hc ob|ccIvcpossb!IyspacccanaIsobcdcncdbyd!!crcnIcom-
bnaIons o! Ihc Iwo Iradc ob|ccIs buI now Ihc acIua! amounIs o!
hgh-quaIIy !!n and parIa!!y workcd sIonc Ioo!s IhaI Ih agcnIs
havcava!ab!cIoIhcmmusIbcIakcnnIoaccounI.ThaIs,un!kcIhc
sub|ccIvc spacc s whch on!y dcsrcs arc mappcdn Ihc ob|ccIvc
spacc wc musInc!udc Ihc nIaIcndowmcnIs o!cacho!Ihc Iradcrs,
snccIhcsc dcnc whaIcanacIua!!ybcIradcd. Inaspacco!a!! com-
bnaIons o! Iradc ob| ccIs Ihc na!cndowmcnI o! boIh agcnIs arc
cachasng!cponI. oncagcnImayhavca!!IhcparIa!!yworkcdIoo!s
wh!c Ihc oIhcr has a!! Ihc hgh-quaIIy !!nI, !or cxamp!c, or Ihcy
maycachbrngIoIhcIradngsIuaIonagvcnamounIo!cachob|ccI.
ThcoIhcrob|ccIvc componcnIsIhcprccaIwhchIhcIwoob| ccIs
arc cxchangcd. Sncc Ihcrc s no moncy nvo!vcd Ihcsc prccs arc
sp!y Ihc amounI o! onc ob|ccI IhaI s cusIomar!y cxchangcd by
anamounIo!anoIhcrob|ccI. IromIhccndowmcnIponI |IhcponI
rcprcscnIng Ihc nIa! sIaIc o! Ihc nIcracIon) prcc !ncs may bc
drawnshowng Ihc raIos o!Ihc Iwo goods IhaIcou!dbc Iradcd aI a
gvcn prcc. Ina!!y, bccausc boIh ob|ccIvc and sub|ccIvc spaccs
dcnc possb!c combnaIons o! quanIIcs o! Ihc samc ob| ccIs Ihc
cndowmcnI ponIs and prcc Incs o! Ihc !ormcr can bc nIcgraIcd
wIh Ihc nd!!crcncc maps o! Ihc !aIIcr Io yc!d a sng!c composIc
dagramcaI!cdandgcworIhBox,abcauI!uIgcomcIrcaIconsIruc-
Ion condcnsng nIwo dmcnsonsup Io ! 2d!!crcnIconccpIs!rom
cconomcs
1 1
Bccausc Ihs composIc dagram nc!udcs sub|ccIvc and ob|ccIvc
!acIorsIa!!owsusIodcncIhcIwogradcnIsIhaIdrvcIragnIcr-
acons. On onc hand, Ihc scIs o! nd!!crcncc curvcs arrangcd by
ncrcascd dcsrab!Iy dcnc Iwo sub|ccIvc gradcnIs o! saIs!acIon,
onc!orcachagcnI. AnmporIanI!caIurco!Ihcsc gradcnIs s pars o!
IJJ
IJ+
NLbN N b MUL1N
nd!!crcncc curvcs IhaI |usI Iouch cach oIhcr. AI Ihcsc ponIs o!
IangcncyIhc amounI o! saIs!acIonIhaI cach agcnI wou!dgcI!rom
an addIona!unI o!Ihc Iradcd ob|cI arc dcnIca! andhcncc Ihcrc
s no!urIhcr nccnIvc Io Iradc. Thcsc arc Ihc ponIs aI whch Ihc
sub|ccIvc gradcnIs havc bccn dsspaIcd. On Ihc oIhcr hand, an
ob|ccIvcgradcnIdcnngIhc poIcnIa!gans!romIradcscapIurcd
byIhcdagramas a rcgonconIanngcombnaIonso!IhcIwogoods
IhaI arc morc dcsrab!c Ihan Ihc nIa! cndowmcnIs. U wc cou!d
cxacI!yquanI!ydcsrc wc wou!dbc ab!c Io csIabIshIhaI na gvcn
IransacIonIhc !oscs o! onc agcnI wcrc morc Ihan compcnsaIcd by
Ihcgans o! Ihc oIhcr and IhaIIhc IradngnIcracIonhad achcvcd
unambguousgans. BuI sncc wccaoIquanI!yI Ihc on!y unam-
bguous |udgmcnIs wc can makc abouI nIcracIons IhaI mprovc
saIs!acIon arc ouIcomcs nvo!vnggans !oronc agcnIandno !osscs
!or Ihc oIhcr. Usng Ihcsc qua!IaIvc | udgmcnIs wc can dvdc Ihc
dagramnIoIwo arcas, onc conIannga!!ocaIonsIhaImakc aI!casI
onc agcnI worsc o!! and anoIhcr conIanng Ihosc IhaI do noI and
IhaI Ihcrc!orc acI as an nccnIvc Io Iradc. WIhn Ihc sccond arca
Ihcrcs asngu!arIy, ca!!cda compcIIvc cquIbrum, dcnngIhc
ponI aIwhcha!! poIcnIa! gans !rom Iradc havc bccn cxhausIcd,
IhaIs, IhcponIaIwhchIh ob|ccIvcgradcnIhasdsappcarcdand
cannoI!ongcrdrvc nIcracIons.
Ikc any oIhcr dagram capIurng Ihc mcchansm-ndcpcndcnI
componcnIo!ancxp!anaIonIhcdgcworIhBoxsaysnoIhngabouI
IhcacIua!proccssIhaIwou!ddrvcIwoagcnIsIowardIhc sngu!arIy.
Whcn modcIng Ihs dynamc proccss cconomsIs usua!!y makc Iwo
assumpIons whch arc un|usIcd n Ihc prcscnI conIcxI. IrsI, I s
assumcdIhaIIhc agcnIs havc Ihc ca!cu!aIona! capacIyandIhcncc-
cssaryn!ormaIonImakcopIma!choccsbcIwccna!IcrnaIvc com-
bnaIons o! Ihc Iwo goods. Sccond, Ihc nIcracIons bcIwccn Ihc
IradcrsarcassumcdIobcgudcdby aucIonccrwhodocsnoIa!!ow
Ihc IransacIon Io procccd unI! I can Iakc p!acc aI Ihc sngu!arIy.
IhcaucIonccrroposcs aprcc andIhcIradcrsanswcrbygvngIhc
quanIIcs o! a good Ihcyou!d bc w!Ing Io Iradc aI IhaI prcc: !
Ihcsc quanIIcs!cavc gans o!Iradc unrca!zcd |!Ihcrc arc wasIc!u!
cxccsscsordccIs) Ihcnasccondprccsproposcd, quanIIcso!!crcd,
andIhc proccsssIcraIcd. U Ihs wayIhc auIonccr gropcs Is way
Ioward Ihc compcIIvc cqu!brum ad auIhorzcs Ihc Iradc on!y
MULlLN1b N b1NL L LLNMLb
whcnIhcsngu!arIyhasbccnrcachcd.Thsmcchansmscar!ynoI
rca!sIcnmosIIradng sIuaIons, cvcnn conIcmporary oncs, andI
docs noI show how prccs can cmcrgc cndogcnous!y !rom acIua!
ncgoIaIons. So!romIhc ponI o!vcwo! modcIng pa!co!IhcIradc
wc wou!dhavc Io gcI rd o!Ihc aucIonccr and o! agcnIs posscssng
unrca!sIcca!cu!aIona! capacIcs | opImzngraIona!Iy)
l2
BoIh o! hcsc prob!cmaIc assumpIons havc bccn addrcsscd n
mu!IagcnI compuIcr smu!aIons. In onc o! Ihcsc Ihc assumpIon o!
opImzng raIonaIIy was Iack!cd drccI!y and radca!!y. Ihc agcnIs
hadnoawarcncsso!IhcIradngsIuaIon:nomcmoryo!pasIIransac-
Ions: no noIon o! proIs: and on!y cnough nIc!Igcncc Io avod
!osscs.ThcscagcnIswcrc appropraIc!yca!!cdzcro-nIc!!gcnccIrad-
crs. InsIcad o! an aucIonccr Ihc smu!aIon uscd a doub!c aucIon.
agcnIsddnoI rcspondIo cxogcnousprccsbyprovdngn!ormaIon
abouI Ihc quanIIcs Ihcy wcrc w!!ng Io Iradc buI suppIcd prccs
drccI!y. bds !IhcywanIcdIobuyor asks u IhcywanIcdIo sc!!.
Thc bds andasks wcrc maIchcdconInuous!ycvcn!Ihc sngu!arIy
had noI bccn rcachcd. In oIhcr words, unIkc Ihc proccss drccIcd
by Ihc aucIonccr nwhchIransacIonsIakc p!acc on!yaI Ic com-
pcIIvc cquIbrum n a smu!aIcd doub!c aucIon Ihcy can occur
anywhcrcwIhnIhcarcadcnngIhcgans!romIradc,IhaIs,Irans-
acIonscanIakcp!accaway!romcquIbrum.WhcnzcronIc!Igcncc
Iradcrs wcrc p!accd n an dgcworIh Box Ihcy wcrc ab!c Io ovc
IowardIhc sngu!arIyandcomc osc II usng on!y!oca!n!orma-
Ion Io cImb Ihc gradcnIs o! saIs!acIon. WhaI Ihc ouIcomc o!Ihs
smu!aIonshowcd was IhaI Ihc cruca!ngrcdcnI nIhc mcchansm
was noI Ihc ca!cu!aIona!abIIcs o! Ihc Iradcrs buI Ihc sIrucIurc o!
Ihc nIcracIon, an nIcracIon IhaI camc osc Io Ihc hagg!ng IhaI
Iakcs p!acc nmarkcIp!accsorbazaars
SmIar rcsu!Is wcrc obIancd n anoIhcr smu!aIon ca!!cd Sug-
arscapc n whch Ihc IoIa!!y dscmbodcd zcro-nIc!!gcncc Iradcrs
wcrc rcp!accdbyagcnIsIhaI had a body |IhaI s, a mcIabo!sm) and
IhaIwcrc sIuaIcdna spacc wIha ccrIandsIrbuIono!rcsourccs.
Thcbchavoro! boIh agcnIs andrcsourccs was cnIrc!y dcIcrmncd
by ru!cs and was Ihcrc!orc noI cmcrgcnI, buI Ihc usc o! ru!cs s an
advanIagc nIhs casc bccausc !Ihc acIons o!Ihc agcnIs had bccn
produccdbyancura!ncII wou!dbchardnoI IovcwIhosc acIons
as mp!yng pcrsona! agcncy. WIh ru!cs, on Ihc oIhcr hand, no
IJb
IJ
NLbN N b MUL1N
assumpIionabouIundcrIying mcchanisms ismadcso agcncycanbc
aIIribuIcd Io Iargcr uniIs, Iikc cnIirc houschoIds or communiIics.
NcvcrIhcIcss somc carc musIbc IakcnwhcninIcrprcIing Ihc ruIcs
sincc Ihcsc spccicd Ihc rangc o! vision o! agcnIs-dcncd as Ihc
numbcro!gridsquarcs o!Ihcspacc IhcyinhabiIcdwiIhinwhichIhcy
couId dcIccI Ihc prcscncc o! !ood-as wcII as Ihcir mcIaboIism, Ihc
amounI o! rcsourccs Ihcy had Io consumc Io conIinuc moving and
prcvcnI dcaIh. U our casc vision and mcIaboIism musIbc Iakcn
Io rc!cr Io Ihc pcrccpIuaI capaciIics and aIimcnIary nccds o! cnirc
communiIics.Ihcconscnsusvicwrcachcda!IcrmanypcrsonaIobscr-
vaIionsandIhc subsisIcnccacIiviIicsnccdcdIokccpIhccnIircgroup
aIivc.KuIcsaIsospccicdIhcbchavioro!rcsourccsinIhcsimuIaIion.
aI cach poinI inIimc cach squarc inIhc Iwo-dimcnsionaI spacc had
boIh a givcn quanIiIy o! sugar, a raIc aI which iI rcpIcnishcd iIscI!,
and a II onIhc maximum amounI IhaI couId cxisI Ihcrc. AgcnIs
uscd Ihcir visionIo inspccI aII squarcs wiIhinIhcir rangc, idcnIicd
Ihosc unoccupicd squarcs wiIh Ihc mosI sugar, and movcd Io Ihc
ncarcsI onc Io gaIhcr Ihc rcsourcc. Through Ihis simpIc ruIc Ihcy
acquircdIhc cmcrgcnIabiIiIyIocIimbrcsourcc gradicnIs.
AIIhoughIhc agcnIsinIhissimuIaIionhad |non-cmcrgcnI) prc!cr-
cnccs, Io movc Io Ihc ncarcsI squarc conIaining morc sugar, !or
cxampIc,IhcydidnoI ycIhavc Io makc choiccsbcIwccnaIIcrnaIivc
rcsourccs.ToaIIow!orIhisIohappcna sccondrcsourcc IogcIhcrwiIh
a sccond mcIaboIism wasinIrduccd. spicc. NowasIhcagcnIsmovcd
aroundIhcycon!ronIcdchoiccsbcIwccnsquarcsmorc orIcssrichin
sugarandIhoscwiIha grcaIcr orIcsscr abundancc o!spicc. SinccIhc
rcsourccswcrcdcncdbyruIcsiIwouIdbcrcIaIivcIysimpIcIochangc
Ihcm Io simuIaIc choiccs bcIwccn high-quaIiIy!IinI and parIiaIIy
workcd IooIs, buI Ihis wouId noImakc a di!!crcnccin a simuIaIion
dcsigncd Io cxpIorc Ihc mcchanism o! cmcrgcncc o! priccs. IcIs
Ihcrc!orc sIick Io Ihc originaI!ormuIaIion. To dcIcrminc an agcnIs
dccisions iIs prc!crcnccs !or di!!crcnI combinaIions o! quanIiIics o!
sugarandspiccadIobccharIcd,IhaIis,cachagcnIhadIobcassigncd
anindi!!crcncc map. ScvcraI !acIors cnIcrcd inIo Ihc dcIcrminaIion
o!Ihcindi!!crcncc curvcs in Sugarscapc. Ihc agcnIs mcIaboIisms!or
sugarandspicc:IhccurrcnIamounIso!cachgoodIhcycurrcnIIypos-
scsscd: and Ihc currcnIdcgrcc IohichIhcirmcIaboIicnccdswcrc
saIiscd. Thcsc !acIors dcIcrmincd Ihc rcIaIivc dcsirabiIiIy o! sugar
MUL1LN1b N b1NL L LLNMLb
andspiccaIanygivcnmomcnI!orcacho!IhcagcnIsasa!uncIiono!
IhcirrcIaIivcinIcrnaIscarciIyandIhcircurrcnIcndowmcnIs. IinaIIy,
Ihc agcnIs wcrc givcnIhc |non-cmcrgcnI) capaciIyIoIradc byadding
a ncw ruIc. Thc Iradc ruIc canbc dcscribcdindgworIh Box Icrms.
i! Ihc indi!!crcncc curvcs o! a givcnpair o! agcnIs wcrc IangcnI Io
cach oIhcrIhcrc wcrc no inccnIivcs Io Iradc so Ihc coupIc did noI
inIcracI:i!Ihc curvcs wcrc noIIangcnIIhcnadirccIiono!Iradcwas
csIabIishcd, oncagcnImayhavcnccdcdsugarandIhcoIhcrspicc, so
sugar movcd inonc dirccIion and spicc inIhc oIhcr. Thcna raIio o!
sugar Io spicc was csIabIishcd |via abargainingruIc) and Ihc quanIi-
Iics o! cach good IhaI wouIdbc Iradcd aI IhaI pricc wcrc chcckcdIo
scc H no agcnI was madc worsc o!!, IhaI is, i! Ihc IransacIion was
wiIhinIhcarcadcningIhcgains!romIradc,inwhichcascIhcIrans-
acIionwascarricdouI.
To scc whcIhcr a!Icr many such Irading inIcracIions an cnIirc
popuIaIion couId coIIccIivcIy rcach Ihc poinI o! compcIiivc cquiIib-
rium 200 agcnIswcrc un!cashcd inaspaccwiIhaccrIaindisIribuIion
o!boIh sugar and spicc. AI Ihc sIarI o!Ihc simuIaIionIhcrc was sig-
nicanI variaIion inpriccs across Ihc popuIaIionbuI as iI procccdcd
IhcrcwasaIcndcncyIoconvcrgcIowardIhccquiIibriumpricc,ncvcr
quiIc rcaching Ihc singuIariIy buI comng quiIc cIosc and sIaying
wiIhinasmaIIncighborhoodaroundiI.U oIhcrwords,priccscmcrgcd
sponIancousIy away !rom cquiIibrium wiIhouI Ihc nccd !or an
aucIionccr.'Thcmcchanismo!cmcrgcnccdcpcndcdonIhc!acIIhaI
as agcnIs rcpcaIcdIy inIcracIcd Ihcir inIcrnaI vaIuaIions !or Ihc Iwo
rcsourccs bccamc morc and morc aIikc: and Ihc morc simiIar Ihcir
vaIuaIions bccamc Ihc cIoscr Ihcy aII goI Io a singIc pricc. On Ihc
oIhcr hand, somc dcgrcc o!pricc dispcrsion survivcd unIiIIhc cndo!
Ihc simuIaIion,adcgrcco!dispcrsionIhaIwasincrcascdbyany!acIor
IhaI inIcr!crcd wiIh Ihc inIcnsiIy o! Ihc inIcracIion. rcducing Ihc
rangc o!visiono! Ihc popuIaIion |sincc Ihis madc agcnIs Icss abIc Io
IocaIcpoIcnIiaIIradcrs): aIIowing agcnIsIodic cvcni!Ihcy mcI Ihcir
mcIaboIic nccds | shorIcr Ii!cIimcs rcduccd numbcr o! inIcracIions):
andaIIowingagcnIprc!crcnccsI ochangcinrcsponscIononcconomic
!acIors| sinccIhisincrcascdIhcIimcnccdcd!orinIcrnaIvaIuaIionsIo
bccomc simiIar). In Ihc casc o! paIcoIiIhic Iradc a Iargc dcgrcc o!
pricc dispcrsion is somcIhng Io bc cxpccIcd so Ihc ouIcomc o! Ihc
simuIaIionsupporIs Ihc Ihcsis IhaI Ihc gains!romIradc couIdhavc
IJ
IJd
NlLbN N b MUL1lN
bccnrcaIzcdnIhaIconIcxIwIhouImaknganyo! IhcusuaIassump-
IonsIhaIanIhropoIogsIsndso ob|ccIonabIc.
IcIs draw somcconcIusons!romwhaI hasbccnargucdupIoIhs
ponI.NuIIagcnIsmuIaIonscangudc usnIhc scarch!orsoIuIons
IoarchcoIogcaIprobIcmsasdcncdbyIhcavaIabIccvdcncc. dcpos-
Iso!sIoncIooIsandIhcdcbrsIc!IbchDddurngIhcrproducIon.U
somc cascs Ihc probIcm s Io accounI !or a parIcuIar dsIrbuIon o!
Ihcsc dcposIs, a probIcm Iobc dscusscdbcIow, buI Ihcrc s aIso Ihc
qucsIon o!cxpIannghow rarc mporIcdmaIcraIs goI Io Ihc pIacc
whcrc Ihcy wcrc !ound. Bccausc wc do noI know Ihc mcIhod o!
IransporI IhaI was uscd muIIagcnI smuIaIons can hcIp nc-Iunc
our nIuIons abouI Ihosc unknown IransporI mcchansms. WhIc
IhccurrcnIIyprc!crrcd cxpIanaIons somc kndo! ccrcmonaIIrans-
porI, anaccounIIhaIdrawsIspIausb!Iy!romIhc!acIIhaI nsIanccs
o! I havc bccn obscrvcd n somc conIcmporary hunIcr-gaIhcrcrs,
Ihcrc s aIso Ihc smpIcrmcchansm o! Iradc bcIwccn communIcs.
Ths opIon has bccnundcrcsImaIcdbyarchcoIogsIsparIIybccausc
o!IhcprcsupposIonsbuIInIocconomcmodcIs. BuI!IhcIaIIcrarc
IakcnIo addrcss onIyIhcmcchansm-ndcpcndcnI componcnI o! an
cxpIanaIon, IhaIs, Ihc snguIarIcs sIrucIurng Ihc possbIIy sacc
o!aIradDgsIuaIon, andnoI Ihc acIuaImcchansmsnvoIvcdnIhc
makng o! dccsons or n prcc ncgoIaIons Ihcn Ihc assumpIons
bccomc Icss probIcmaIc. A!IhoughsmuIaIonsaIsoopcraIc aIa rcIa-
IvcIy mcchansm-ndcpcndcnI IcvcI Ihc capacIy o! compuIaIonaI
proccss Io Irack hsIorcaI paIhs Ioward a snguIarIy, and Ihc addcd
advanIagc o! usng cmbodcd and spaIaIIy sIuaIcd agcnIs, makcs
Ihcm dcaI Io cxpIorc poIcnIaI canddaIcs !or Ihc IransporImccha-
nsmIhaIaccounIs!orIhc avaIabIccvdcncc.
Togo bcyondIhs rcIaIvcIyhumbIc roIc and IackIc archcoIogcaI
probIcms !kc cxpIanngIhc acIuaI dsIrbuIon and composIon o!
dcposIs o! sIonc arI!acIs n a spccc gcographcaI arca, wc nccd a
morccxp!cIway o!modcIngIhc d!!crcnI cIcmcnIso!amuIIagcnI
smu!aIon. Ihcrcsourcc Iandscapc, IhcbchavoraIru!cs,andIhcIcm-
poraIscaIco!Ihc subssIcncc acIvIcs. Thss ahgh!ynIcrdscp!n-
aryIaskbuIarchcoIogyIsc!!has!ongrc!cdonoIhcrc!ds |chcmsIry,
gcoIogy, ccoIogy, cIhnoIogy) Io rcconsIrucI subssIcncc acIvIcs and
scIIIcmcnIpaIIcrns !rom sIonc IooIs, !ossIzcdpo!Icn, anma!boncs,
and hcarIh rcmans. Smu!aIcd rcconsIrucIons o! Ihc subssIcncc
MUL1LN1b N b1NL L LLNMLb
cconomy o! hunIcr-gaIhcrcrs vary n Ihc dcgrcc o! dcIaI o! boIh
bchavoraIpaIIcrnsandrcsourcc gradcnIs.U IhccarIcsIsmuIaIons
Ihc Iypc o!pIanIs andanmaIprcy assumcdIobcIhc IargcIo! !orag-
ng acIvIcs wcrc gucsscd cIhcr !romconIcmporaryhunIcr-gaIhcr-
crsnIhc samcrcgonor!romIhc !oss!zcdrcmans!oundn a cavc
oranoIhcr sIc. Thcn Ihc conIonsa!!ccIngIhc rcIaIvc abundancc
o! Ihosc rcsourccs Ioday |ran!aII !or pIanIs, popuIaIon dynamcs
!oranmaIs)wcrcmodcIcdIocrcaIcsIaIsIcaIscrcs o!good, bad, and
avcragc ycars Io bc uscd B Ihc smuIaIons. AIhough Ihc rcsourcc
dsIrbuIonsgcncraIcdIhswaywcrccIcarIymorcrcaIsIcIhanIhosc
o!IhcSugarscapcsmuIaIons|usIdscusscd, cxIrapoIaIng!rompr

s-
cnI condIons canbc probIcmaIc!Ihc Iandscapc s noI IcmporaIIy
sIabIc, ! IargcI spcccs arc now xIncI, or ! ccoIogcaI dvcrsIy has
bccn grcaIIy rcduccd. "
To avodIhcsc poIcnIaIsourccso! crror daIaandIcchnqucs !rom
muIIpIc dscpIncs musI bc uscdIo rcconsIrucI anccnI Iandscapcs.
usng Ihc n!ormaIon sIorcd n cc shccIs, Ircc rngs, scdmcnI, and
rockspaIcocImaIoIogy canhcIpIo rccrcaIc IhcIcmpcraIurc andprc-
cpIaIon condIons o! a gvcnImcpcrod: gcomorphoIogy can bc
uscdIo undcrsIand Ihc hsIory and dynamcs o!Iand!orms, andIo
rccrcaIcIhcc!!ccIo!rvcrs,voIcanocs,andgIaccrsonIhcdsIrbuIon
andquaIIyo!soIs: ccoIogycan!urnshn!ormaIonabouIIhcsucccs-
son dynamcs o! d!!crcnI pIanIs andIhs, IogcIhcrwIhIhc ouIpuI
!rompaIynoIogy |Ihc sIudy o!poIIcn, sporcs, andparIcuIaIc organc
maIIcr), cangudcIhcrcconsIrucIono!anccnIdsIrbuIonso!pIanIs
o!a gvcn spcccs." Oncc Ihc spaIaIcomponcnI o!rcsourcc dsIrbu-
Ions has bccn workcd ouI wc nccd Io add a IcmporaI dmcnson
Io Ihc smuIaIons. Ths s smpIcdby Ihc !acI IhaI Ihc subssIcncc
acIvIcso!hunIcr-gaIhcrcrsIcndIohavcacycIcaIsIrucIurc.U soc
cascs Ihc pIanIs IhaI arc Ihc IargcI o!!oragng acIvIcs rpc B d!!cr-
cnI scasons and Ihs gvcs Ihc acIvIcs a ycarIy cycIc, somcImcs
accompancdbyscIIIcmcnIpaIIcrnsIhaIdspIayasm!arpcrodcIy.
InoIhcrcascs amuch shorIcrpcrodmaybcuscdsuchasadaIycycIc
dvdcdnIo daI andnocIumaIacIvIcsThspcrodcrcpcIIon
o! acIvIcs can bc uscd Io gvc Icmpora! sIrucIurc Io a smu!aIon
and Io corrcIaIc Is ouIpuIwIh Imc scrcs !or IcmpcraIurc andprc-
cpIaIonchangcs, Ihcrscand!aIIo!avaIabIc groundwaIcr, andIhc
changcsnsoIqua!Iy crcaIcdbyIhc dcposIon o!scdmcnIs.
IJV
I+
NLbN N blMUL1N
Thcncxtstcpistomakcmorcrca!isticthcbchaviora!andctabo!ic
ru!csthatcapturc thc dccision-makng andconsumption activiticso!
agcnts. tLno!ogica!studicso!contcmporaryhuntcrgathcrcrscanbc
ava!uab!c sourcc o!in!ormationinthis rcgardbutwcmustbccarc!u!
not to rci!y particu!ar cascs into a timc!css sct o! propcrtics. In thc
I60s, !or cxamp!c, thc conscnsus vicw o! contcmporary huntcr-
gathcrcrs charactcrizcdthcmas!ivingingroups o!25 to 50 mcmbcrs,
withnomatcria!posscssionstoconstrainthcirhighmobi!ity, ascxua!
divisiono!!abor,!!cxib!cmcmbcrship, andvcrycga!itarianauthority
structurc. But a dccadc !atcr othcr tribcs wcrc !ound that did not
match this !ist o!propcrtics, bcing scdcntary part o!thc ycar, !iving
in !argcr groups, and storing somc posscssions Thcsc two cascs can
bc uscd to dcnc thc two cxtrcmcs o!a continuous possibi!ity spacc
andthcbchaviora! ru!cs!orthcsmu!ations canbc dcrivcdbymaking
cducatcd gucsscs about thc !ikc!y position o! pa!co!ithic huntcr-
gathrcrs in this continuum. Scvcra! smu!ations may bc pcr!ormcd
withru!cs dcrivcd!romdi!!crcntpartso!thccontinuumtochcckthc
scnsitivityo!thcrcsu!tstodi!!crcntassumptions.Iina!!y, tobcab!cto
uscthcscsmu!ations tomakcspccicprcdictionsaboutthcdistribu-
tiono!dcpositso!stoncarti!actsthc!attcrmustbcavai!ab!cin awc!!
prcscrvcd statc. H gcncra!, thc !urthcrback intmcwcgo thc morc
disturbcd thc archco!ogica! rccord is so most dcti!cd simu!ations
havc!ocuscdonmorcrcccntpcriods, suchasthcmcso!ithic,thctran-
sitiona!pcriodbctwccnthcuppcrpa!co!ithicandthca!rcadyagricu!-
tura! nco!ithic. This pcriod di!!crs !rom thc oncs discusscd so !ar in
onccrucia!rcspcc, thcavai!abi!ityo!!anguagc, a changc rc!!cctcdin
thcsmu!ationsbythcnccdtoincorporatcthc!!owo!scmanticin!or-
mationamongthcagcnts.
In onc simu!ation, ca!!cd Nagica!, thc goa! was to rcproducc
thc distribution ad composition o! stonc arti!acts on an is!and o!
Wcstcrn Scot!and whcrc mcso!ithic huntcr-gathcrcrs oncc !oragcd
!or hazc!nuts. Uthissmu!ationrcsourcc gradicntswcrc modc!cdin
dctai!using thcpa!cocnvironmcnta!approach|ustdiscusscd, inc!ud-
ing compctition among typica! tcmpcratc !orcst trccs |oak, birchI
withhazc!. Ycar!y, month!y, anddai!ytmcsca!cswcrcinc!udcd.Thc
activiics on a ycar!ytmc sca!c wcrc dctcrmincdby thc!i!c cyc!c o!
thc trccs thcmsc!vcs. onc scason with a hazc!nut harvcst and onc
without. This, in turn, was assmcd to a!!ccthuman activitics !ikc
MUL1LN1b N b1NL L LLNMLb
gathcringonabasccamp!o!!owcdbydispcrsing around agivcnarca
o! thc is!and. Thc month!y timc sca!c was dctcrmincd by dccisions
rcachcd by conscnsus about thc !ocation o!thc basc camp. whcthcr
or not to movc it to a ncw p!acc with bcttcr chanccs o! incrcasing
thcproductivity o!!oragngactivtics. Iina!!y, on a dai!ybasis agcnts
!oragcdor cxp!orcdduringthcdayandgot togcthcratnightto sharc
thcrcsu!ts o!thciractivitics.Unikcthcprcvious!ydicusscdsimu!a-
tions in which thc agcnts wcrc cntirc communtics hcrc thc acnts
wcrc takcnto bc individua! pcrsons so thc night!y rcports had tobc
modc!cd cxp!icit!y. Thcso!utiontothisprob!cmwasvcryingcnous.
Sinccthcactiviticso!thcagcntshadtotakcp!accinasim!atcdspacc
withthcsamc charactcristics asthc origina!is!and,a standard!ormat
!orthcstoragc o!spatia!data,ca!!cdaCcographicIn!ormationSystcm
| or CISI , was uscd. A CIS isbasica!!yamaponwhichmu!tip!c !aycrs
arc supcrimposcd cach containng n!ormation about a particu!ar
!andscapc propcrty |soi! composition, distributions o! p!ant spccics,
watcr avai!abi!ityI . InNagica! thcsc maps wcrc givcn an additiona!
usc. to rcprcscnt thc know!cdgc that thc agcnts acquircd about thc
!andscapc. In othcr words, thccognitivcmapsthatthcagcnts !ormcd
o!rcsourccdistributions,mapsthatwou!dhavc bccn crcatcdbyrca!
huntcr-gathcrcrs through a combination o! cpisodic and scmantic
mcmory, wcrc cach rcprcscntcd by a CIS map. That waywhcn thc
agcnts gathcrcd at night to rcport thcir dai!yndings thc !inguistic
cxchangc among thcm cou!d bcmodc!cdbymcrging a!! thcndivid-
ua! CIS maps to !orm a compositc onc that was thcn rcdistributcd
amonga!!thc mcmbcrs o!thc group.
Un!ikc thc agcnts in Sugarscapcthc usc o!smu!atcd vsua!in!or-
mation did not !orm thc basis on which movcmcnt dccisions wcrc
madc,thcagcntso!Nagca!usinginstcadthcirintcma!CIS maps!or
thatpurposc But as in Sugarscapc Nagica!s agcnts wcrc hctcrgc-
ncous inthis capacity. whi!c in thc !ormcr thcyposscsscd di!!rcnt
rangcs o!vision in thc !attcr thcy ana!yzcd di!!crcnt!y sizcd portions
o!thc map around thcir currcnt position bc!orc moving to a ncigh-
boring !ocation. Agcnts who madc dccisions bascd on a sma!!cr por-
tiono!thcmapwcrcbcingconscrvativcinthc scnscthatthcywcrc
tryngto rcduccriskandmprovcthcirshort-tcrmgains:thoscusng
!argcrportions took morc riskbymovingto!ocationswithunknown
hazc!nut abundancc but cou!d bcnct !rom!ongcrtcrm gains i!thcy
I+I
I+Z
NLbN N b MUL1N
dscovcrcd ncw sourccs o! !ood. Fna!!y, Ihc sIonc arI!acIs wcrc
dvdcd nIo d!!crcnI caIcgorcs |wasIc !rom producIon proccsscs,
sma!! Ioo!s, !argcr scrapcrsI and probab!Ics IhaIIhc agcnIs wou!d
dscardIhcmwcrc assgncdaccordngIoIhcrposIon rc!aIvc IoIhc
basc camp andIhcsIagco!Ihcycar!yscason. So !arIhcmanrcsu!Is
o!usngNagca!havcbccnIorcducc Ihc!kc!hoodo!somc prcvous
sccnaros proposcd Io accounI !or Ihc cxsIngarchco!ogca!daIa. In
parIcu!ar, Ihc hypoIhcss IhaIIhc on!ydcIcrmnanI o! Iand usc was
IhcgaIhcrngo!hazc!nuIs, anadmIIcd!ymporIanIrcsourcc, has!osI
somc crcdb!Iy. Nanyruns o!Ihc smu!aIon wcrc pcr!ormcd, cach
run rcprcscnIng abouIIcnycars o! !oragng acIvIy, usngd!!crcnI
!andng sIcs around Ihc s!and as wc!! as d!!crcnI dsIrbuIons o!
rsk-Iakngbchavor. YcI, nonc o! Ihc ouIcomcs managcd Io rcpro-
duccIhccxsIngdsIrbuIono!dcposIso!sIoncarI!acIs.AIIhough
Ihc smu!aIons rcsu!Is wcrc ncgaIvc Is conIrbuIon was c!car!y
posIvc. I showcd IhaI oIhcr spaIa! consIranIs musI havc bccn n
opcraIon, such asIhc ava!ab!Iyo! hghquaIIy BnIncarIhc coasI
IhaImayhavcbccn annccnIvc Io crcaIcspccaIsIbasc campsuscd
!or Ihc producIon o! Ioo!s. U oIhcr words, Ihc smu!aIons rcsu!Is
ponIcdIousc!u!a!IcrnaIvchypoIhcscs.
Sm!ar!y ncgaIvc buI hgh!y n!ormaIvc rcsu!Is havc bccn
obIancd n a Sugarscapc smu!aIon nwhch rcsourcc dsIrbuIons
wcrc rcconsIrucIcd usng pa!cocnvronmcnIa! modc!s. Thc smu!a-
Ion Iack!cd an cvcn morc rcccnI pcrod, Ihc Nco!Ihc, a pcrod n
whch agrcu!Iurc was a!rcadypracIccd and Ihc nomadc I!csIy!c o!
hunIcr-gaIhcrcrs had bccn abandoncd. Wh!c sIonc Ioo!s conInucd
Iobc uscd andIhcrncrcascdsophsIcaIonand!uncIona!d!!crcn-
IaIon !c!I bchnd a rchcr archco!ogca! rccord, Ihc Nco!Ihc nIro-
ducc a morc scnsIvc mcdum. poIIcry. Thc p!asIcIy o! c!ay gvcs
poIIcryagrcaIcrrangco!sIyIsIcvaraIonandIhsIogcIhcrwIhIhc
!acI IhaI I wasIypca!!yproduccd !oca!!y supp!cs us wIh cvdcncc
abouIrcgona!groupngs and abouI IhcnIcracIonsbcIwccn ncgh-
borngcommunIcs.AnoIhcrsourcco!n!ormaIon!romIhspcrod
sscII!cmcnIpaIIcrns.ScII!cmcnIs!cavcmncra!zcdrcmansarrangcd
nscparaIc sIraIa. somc!aycrsmayshowIhaI!or awh!c mosI o!Ihc
popu!aIonwas conccnIraIcd nonc!argcv!!agc, !or cxamp!c,wh!c
oIhcr!aycrsmayrcvca!IhaI aI oIhcr ImcsIhc scII!crs dspcrscd and
Ivcdnmany sma!!v!!agcs.For archco!ogsIs IryngIo cxp!anIhcsc
MULlLN1b N b1NL L LLNMlLb
varaIonsIhcqucsIonswhcIhcrIhcycanaI!bcaIIrbuIcdIochangcs
n rcsourcc gradcnIs |so! producIvIy, ava!abIIy o! waIcrI or
whcIhcrcu!Iura! changcs wcrc a!sonvo!vcd.Nu!IagcnIsmu!aIons
can bc uscdIo Iack!c Ihs qucsIon !or spccc hsIorca! cascs. Onc
parIcu!ar casc s Ihc Anasaz cu!Iurc IhaI nhabIcdIhc Iong Housc
VaI!cynnorIhcasIcmArzona,rsIashunIcr-gaIhcrcrs|bc!orc ! 800scI
Ihcnasagrcu!Iura!sIssurvvng ona cconomybascdonIhc cu!Iva-
Iono!mazc unI!Ihcy abandoncd Ihc rcgonnP ! 300!orrcasons
sI!I unknown.
Un!kcIhc sIaIc rcsourcc !andscapcs o!Ihc Sugarscapc smu!aIons
dscusscd abovc Ihosc o! Ihc Iong Housc Va!!cy had Io bc modc!cd
dynamca!!ybccauscIhcproducIvIyo!mazccrops,Ihcmangrad-
cnI drvng Ihc proccss, undcrgocs changc aI scvcra!Imc sca!cs. Is
n!!ucnccd by rc!aIvc!y shorIIcrm varaIons n Ihc mosIurc and
IcmpcraIurc o!IhcaImosphcrcaswc!!asby!ongcrIcrmvaraIonsn
Ihc dynamcs o!rvcrs, a!!ccIngIhcava!abIIy o! groundwaIcr and
Ihc crcaIon and dcsIrucIon o! ncw !crI!c so!. Thc Imc sca!c !or
Ihc smu!aIon was dcIcrmncdbyIhc agrcu!Iura! cyc!c, IhaI s, Ihc
smu!aIonprocccdcdnycar!ypcrods. Thc agcnIsnIhssmu!aIon
wcrc noIndvdua! pcrsons buIndvdua!houscho!ds, composcdo!
vc mcmbcrs cach, whosc mcIabo!c rcqurcmcnIs wcrc ca!cu!aIcd
usng anIhropo!ogca! sIudcs o! oIhcr subssIcncc agrcu!Iura!sIs.
cach mcmbcr o!a!am!ywas assumcd Io consumc ! 60 k!ograms o!
mazcpcrycar, sIornghaIcvcrwas!c!I!orup IoIwoycars.Fam!cs
wcrcgvcnbchavora! ru!csIhc mosImporIanIo!whchwasarcscI-
I!cmcnI ru!c. aI Ihc cnd o! cach ycar cach !am!y ca!cu!aIcd on Ihc
bass o! Ihc currcnI harvcsI and Ihcr sIorcd gran Ihc chanccs IhaI
Ihcrc wou!d bc cnough !oodncxIycar: u I dccdcd IhaIIhcrc wou!d
noIbc cnoughImadc adccsonIorc!ocaIc consIrancdon!y byIhc
ava!ab!Iyo!unoccupcd!arm!andandproxmIyIodrnkngwaIcr.
I!Ihc wrong dccson was madc and noI cnough !oodwasproduccd
Ihc!am!ydsappcarcd!romIhc smu!aIon.achruno!Ihcsmu!a-
Ion gcncraIcd, as Ihc cmcrgcnIcumu!aIvc c!!ccI o! Ihc movcmcnI
dccsons bymany!am!cs ycar a!Icr ycar, apossb!c scII!cmcnI hs-
Iory bcIwccn Ihc ycars P oUU and ! 300. To capIurc sIaIsIca!Ircnds
many such possb!c hsIorcs wcrc gcncraIcd sIarIng !rom d!!crcnI
nIa! condIons and varyng ccrIan paramcIcrs. Thc ouIpuI was
boIha gcographca!rcprcscnIaIon o!scII!cmcnI!ocaIonsIhaIcou!d
I+J
I++
NlLbN N b MUL1N
bccomparcdwIhmapscrcaIcdbyarchco!ogsIs, aswc!IassIaIsIca!
scrcs onpopu!aIon growIh and aggrcgaIon paIIcms IhaI cou!d bc
chcckcd agansIIhosc n Ihc archco!ogca!rccord.
Thc comparsons, howcvcr, wcrc noI nIcndcd Io bc cxaI and
quanIIaIvc. I wou!dbc unrca!sIc Io cxpccI agrccmcnI on ovcra!I
popu!aIonszcs,numbcro!houscho!ds, orszco!scII!cmcnIs.U !acI,
Ihc smu!aIon conssIcnI!y gcncraIcd quanIIcs IhaI wcrc Ioo !argc
packng many morc houscho!ds n a sng!c rcsdcnIa! zonc Ihan s
conssIcnI wIh Ihc cvdcncc. BuI Ihc qua!IaIvc agrccmcnI was
sIrkng. !orpopu!aIon growIh, !or nsIancc, Ihc qua!IaIvc !caIurcs
o! Ihc sIaIsIca! scrcs, Ihc pcrods o! growIh and dcc!nc, as wc!! as
Ihosc n whch numbcrs rcmancd consIanI, wcrc mosI!y rcpro-
duccd." Sm!ar!y, Ihc rcsponscs o!Ihc smu!aIcdagcnIsIo cnvron-
mcnIa!changcs-IoaggrcgaIc n a !argc v!Iagc Iopcr!ormnIcnsvc
agrcu!Iurc whcngroundwaIcrwas scarcc and Io dspcrsc nIo many
sma!! oncs whcnp!cnI!u! groundwaIcr a!!owcd cu!IvaIonn many
p!accs-gcncraIcd scII!cmcnI paIIcms qua!aIvc!y sm!ar Io Ihosc
n Ihc daIa. Thc man dscrcpancy bcIwccn Ihc smu!aIon and Ihc
archco!ogca! rccord was Isc!! hgh!y n!ormaIvc. Around Ihc ycar
I250!ong- and shorIIcrmvaraIonsn cnvronmcnIa!!acIors con-
cdcdIoproducca subssIcncc crssnIhcIongHousc Va!!cyandby
Ihc ycar I 300 Ihc Anasaz had abandoncdIhc arc. Thc smu!aIon
!a!cdIorcproducc Ihs cvcnIsuggcsIngIhaIIhc dccsonIocmgraIc
wasnoIncccssar!y!nkcdIogradcnI dcgradaIon. I!acI, Ihc smu-
!aIonndcaIcd IhaI Ihc ava!ab!c rcsourccs cou!d havc supporIcd a
sma!!crpopu!aIon dspcrscd nIo sma!IcrcommunIcs. ThaI Ihc rca!
AnasazddnoIIakc advanIagc o!Ihs opporIunIy suggcsIs IhaI Ihc
cu!Iura! and soca! !acIors IhaI wcrc noI nc!udcd n Ihc smu!aIon
musI havc p!aycd an mporIanI ro!c nIhcr dccson."IparIcu!ar,
whcn dca!ng wIh agcnIs IhaI a!rcady posscss !anguagc wc musI
assumc noIon!yIhaI Ihcy cancxchangc!acIua!n!ormaIonbuI a!so
IhaIIhcyan!ormandsharcmagca!bc!c!sIhaIcanp!ayan cqua!!y
mporIanIro!c nIhcr own cxp!anaIons abouI rcsourcc ava!ab!Iy,
and hcncc on Ihcr dccsons Io sIay or Io cmgraIc abandonng Ihc
arca a!IogcIhcr.
Wc w!! rcIurn n ChapIcr I ! Io Ihc ro!c IhaI magca! bc!c!s p!ay
n Ihc dcIcrmnaIon and cxp!anaIon o! human bchavor, and Io
Ihc waynwhchIhc c!!ccI o!Ihosc bc!c!s can bc ncorporaIcd nIo
MUL1LN1b N b1NL L LLNMlLb
mu!IagcnI smu!aIons. BuIbc!orc wc Iack!c IhaIprob!cm wc musI
con!ronI Ihc onc IhaI u!ImaIc!y gvcs rsc Io I. Ihc cmcrgcncc o!
!anguagc. A!Ihough hunIcrgaIhcrcr communIcs durng Ihc hun-
drcds o!Ihousandso! ycas IhcyspcnIsk!!!u!!y workng sIoncmay
havc a!rcadydcvc!opcdsmp!cr!ormso!communcaIon, bascdpcr-
hapsonhandand!aca!gcsIurcs,Ihc gradua!dcvc!opmcnIo!arIcu-
!aIcd !anguagc markcd a Ihrcsho!d bcyond whch an cnIrc!y ncw
wor!dbccamc possb!c. Thccruca!propcrIy o!!anguagc sIhaI!rom
anIcscIo!wordsannnIcnumbcro!scnIcnccs canbcproduccd.
Thus, whaInccds Io bc cxp!ancd s howIhs nnIc combnaIora!
producIvIy cou!d havc cmcrgcd sIarIng wIh a !cw mono!Ihc
symbo!c arI!acIs ncapab!c o! cnIcrng nIo comp!cx parI-Io-who!c
rc!aIons. lIsIoIhs IaskIhaIwc nowIum.
I+b
I+
LHPJLJLN
MJ|||dO|5 d0 |||0| ||VO LdJdO
WhcnhumansrsIbcganIoshapc!!owso!arwIhIhcrIongucsand
pa!aIcsIh acousIc maIIcr Ihcy crcaIcd nIroduccd ycIanoIhcr!aycr
o! comp!cxIynIoIhc wor!d. Thc producIon o!soundrc!cdonIhc
samc mcchansm uscd by oIhcranma!s-asourcc o! osc!!aIng ar
shapcdbyavoca! cavIyacIngasa!Icr-buInhumansIhsmccha-
nsmundcrwcnIscvcra!mporIanIcvo!uIonarychangcs.Thcsourcc,
Ihc !arynx and Is voca! chords, !owcrcd IsposIonrc!aIvc Io Ihc
boncaIIhcbasco!Ihcsku!!cxIcndngIhcrangc o!vowc!sIhaIcou!d
bc produccd, cvcn aI Ihc cosI o! !osng our ab!Iy Io swa!!ow and
brcaIhc aI Ihc samc Imc. Thc !Icrs, n Ium, consIIuIcd by a!! Ihc
d!!crcnI organs IhaI can changc Ihc !orm o! Ihc rcsonaIng cavIy
|Ionguc, !ps, IccIh) bccamc capab!c o!manpu!aIng prcssurc grad-
cnIs wIhcvcr grcaIcrprccson. rcsonanccs nIhc voca! cavIycon-
ccnIraIcdIhc cncrgyo!Ihcar!!owaroundparIcu!ar!rcqucnccsand
Ihcsc conccnIraIons | ca!!cd !ormanIs) bccamc Ihc bu!dng b!ocks
IhaIind!!crcnIcombnaIonswou!dcvcnIua!!yyc!da!!Ihc d!!crcnI
sounds o!!anguagc. Sncc cvo!uIonhasno !orcsghI ncIhcr o!Ihcsc
cvo!uIonary changcs was aimcd aI makng !anguagc possb!c. Thc
dcsccnI o!Ihc !arynx may havc occurrcd, as I has m oIhcr spcccs,
ducIoscxua!sc

ccIon.a!owcr!arynxcanproduccsoundsIhaImmc
Ihc c!!ccI o! a !argcr body szc, an c!!ccI IhaI anma!s cxp!oI durng
IcrrIora! dspuIcs. ' AndIhc grcaIcr conIro! o!Ihc organs producng
!ormanIs was mosI !kc!y a byproducI o! Ihc ncrcascd szc o! Ihc
bran. I!c n Ihc savanna had !ong ago changcd dcIary paIIcrns
addng ncw nuIrcnI sourccs |Iubcrs, mcaI) and ncw Icchnqucs !or
MULlLN1b N KlMl1lYL LNUL
!ood prcparaIon |Ihc usc o! Ioo!s Io crush Iubcrs and rc Io cook
mcaI)-ncrcasng Ihc amounI o! cncrgy consumcd nd makng
Is dgcsIon morc c!ccnI. Thc cxIra mcIabo!c cncrgy ava!ab!c, n
Iurn, was nvcsIcd n a !ongcr cmbryo!ogca! dcvc!opmcnI, wIh a
morc c!aboraIc!o!dng o!ncurona!!aycrspackngmorcbranmaIIcr
nIo agvcnvo!umc.
BuI!IhcrcwasnoIhng unquc!yhuman abouIIhcbo!ogca!cvo-
!uIon o!IhcsoundproducIonmcchansm, IhcacousIcmaIIcrIsc!!
wasunquc bccausc as soon asI cmcrgcdIbccamc parIo!an cv!u-
IonIhaI was scparaIc !rom gcncIcs. Irc!ngusIchuIcrgaIcrcrs
posscsscd !carncd habIs and ski!!s IhaI wcrc IransmIIcd across
gcncraIons and IhaI cou!d Ihcrc!orcp!ayIhc ro!c o!rcp!caIors n a
cu!Iura! cvo!uIonary proccss, a proccss no whch Ihc ncw!y born
!ngusIc sounds cou!dbcnsIanI!ygra!Icd. Thc dcIa!s o!whaI hap-
pcncdIoIhosccar!ysoundsmayncvcrbcknownbuIwcmusIassumc
IhaI!ngusIccvo!uIonbcganwIhmono!Ihcsymbo!carI!acIs.ThaI
s, wc musI assumc IhaIIhc rsI words wcrc noI madc ouIo!parIs
IhaI cou!d bc rccombncd Io !orm oIhcr words, and Iha Ihc words
Ihcmsc!vcs wcrc noIparIsIhaIcou!d cnIcrnIo !argcr scnIcnccs.Thc
on!y communcaIvc !uncIonIhaIIhosc mono!Ihc symbo!carI!acIs
cou!d havc had s IhaI o! !abc!s IhaIrc!crrcdIo drccI!y cxpcrcnccd
ob|ccIs and acIons. Simp!c concaIcnaIons o! Ihosc arI!acIs cou!d
havc!aIcronbccn uscdIocxprcssIhcobscrvcdconsIanIcon|uncIon
o!!ncar causcs andIhcrc!!ccIs, anobscrvaIonIhaIhadbccn madc
rouInc by Ihc pracIccs assocaIcd wIh sIonc Ioo! producIon. Thc
ncxIcvo!uIonary sIcp, gong!rom a scIo!mono!Ihc sounds Io onc
o! words IhaI cou!d bc combncd nIo an nnIc scI o! possb!c scn
Icnccs,smuchmorcd!cu!IIocxp!an.WcnccdnoIon!yIoaccounI
!or Ihc capacIy Io combnc a scI o!words !kc Ihc prcy and runs
away nIo a scnIcncc !kc Thc prcy runs away. Wc a!so nccd Io
accounI !orIhc ab!Iy Io !ccdIhssmp!c scnIcnccbackas annpuI,
IogcIhcrwIh anoIhcr scI o!words !kc Ihc hunIcrs andchasc, Io
producc a comp!cx scnIcncc !kc Thc prcy Ihc hunIcrs chasc runs
away. lnoIhcrwords, wc nccdIocxp!anIhc unquchumanab!Iy
Iousc rccurson.
TogcIa scnsc o!Ihcprob!cmIhaIhadIobc so!vcdbycu!Iura!cvo-
!uIonIw!!bcusc!u!IoconsdcrIhc spacco!a!!possb!c !anguagcs.
Thc sIrucIurc o! Ihs spacc canbc cxp!orcd usng auIomaIa Ihcory,
I+
I+d
NLbN N blMUL1lN
a Ihcory in which mcmory, rccursion, and Ianguagc compIcxiIy
arc cIoscIy inIcrIwncd. AuIomaIa can bc cIassicd inIo a scrics o!
ncrcascd compuIaIionaI capaciIy dcIcrmincd by Ihcir dcgrcc o!
acccss Io mcmory rcsourccs niIc sIaIc auIomaIa, pushdown auIo-
maIa,Iincar-boundcdauIomaIa, andTuringmachincs.This scrics, in
Iurn, can bc rcIaIcd Io a scrics o!Ianguagcs o! incrcascd compIcxiIy.
IiniIc sIaIc auIomaIa posscss no mcmory and can handIc onIy Ian-
guagcs wiIhouI rccursion. rcjulurlunjuujcs. ThcncxIIypchas acccss
onIy Io Ihc IasI iIcm sIorcd in mcmory |Ihc oIhcrs bcing pushcd
downasIackIbuIIhisgivcsiI Ihc abiIiIyIohandIcsimpIc rccursion
and masIcr ccnrcxr-]cc lunjuujcs. U parIicuIar, pushdown auIomaIa
havc Ihc abiIiIy Io kccp Irack noI onIy o! cmbcddcd cIauscs | Ihc
hunIcrschascIbuIIomaIchvcrbsandnounsscparaIcdinascnIcncc,
IhaI is, IomaIch Ihc prcy Io runs away. Thc ncxIIcvcI o!compu-
IaIionaI capaciIy invoIvcs a morc powcr!uI mcmory in Ihc !orm o!
an inniIc Iapc on which symboIs can bc sIorcd buI acccss onIy Io
a smaII porIion o! iI. This is cnough, howcvcr, !or Iincar-boundcd
auIomaIaIohandIcIhcsIorcdsymboIsIogcIhcrwiIhIhcirimmcdiaIc
conIcxI and masIcr ccnrcxr-scnsirivc lunjuujcs. xampIcs o! Ihc morc
compIcxrccursivc consIrucIions IhaI Ihis aIIows arc rarc B naIuraI
Ianguagcs buI do cxisI, as iIIusIraIcd by cross-dcpcndcncicsinwhich
vcrbs arc noI maIchcd Io nouns in Ihc ordcr Ihcy appcar. IinaIIy,
Ihc mosIpowcr!uIauIomaIa,Turingmachincs, havc unIimIcdacccss
Io Ihc mcmory Iapc, can Iakc as much conIcxI inIo accounI as
nccdcd,canhandIcany!ormo!rccursion,andmasIcrrhrusc-srrucrurc
lunjuujcs.' A IpicaI dcskIop compuIcr Iics somcwhcrc bcIwccn a
Iincar-boundcd auIomaIon and a Turing machinc approximaIing
Ihc IaIIcr cvcr morc cIoscIy as mcmory gcIs chcapcr and morc
abundanI.
As mcnIioncd in ChapIcr 2 auIomaIa Ihcory is imporIanI phiIo-
sophicaIIy bccausc iI providcs !or capaiIics whaI sIaIc spacc and iIs
aIIracIors do !or Icndcncics. a cIcar and rigorous way o! cxpIoring
Ihc sIrucIurc o! a possibiIiIy spacc. Thc singuIariIics in Ihis casc arc
consIiIuIcdbyIhc!ourauIomaIa, wiIhniIcsIaIcauIomaIaandTur-
ingmachincs consIiIuIing aminimumand amaximum o! compuIa-
IionaI capaciIy, rcspccIivcIy. ThcIcrm Ianguagc inauIomaIaIhcory
rc!crs noI onIy Io Ihc approximaIcIy 6,000 currcnIIycxisIing naIuraI
Ianguagcs buI aIso Io d Ihc programmng Ianguagcs uscd Io crcaIc
MUL1LN1b N KM1YL LNUL
so!Iwarc !or Ihc di!!crcnI auIomaIa, as wcII as manyIanguagcsIhaI
arcpossibIcbuIhavc ncvcrbccn uscd. Iikcmanyo!Ihc combnaIo-
riaIpossibiIiIyspaccsIhaIhavcbccndiscusscdso!arIhisspaccisin-
niIc in sizcbuI !orIhc purposc o!cxpIoringIhc cvoIuIion o! naIuraI
Ianguagcs wc can conccnIraIc on Ihc rcgion conIaimng conIcxI-
scnsiIivc Ianguagcs. This impIicsIhcassumpIionIhaI IhcparI o! Ihc
human brain IhaI handIcs Ianguagc musI havc Ihc compuIaIionaI
capaciIyo! Iincar-boundcdauIomaIon. UsingncuraI ncIsIocxpIorc
howIhiscapaciIyisimpIcmcnIcdinourbrainswouIdsccm aIrsIIo
bcanipossibIc IaskbccauscIhckindo!ncuraI ncI IhaIcanproccss
IhcIcmporaIscqucnccso!soundsIhaImakcuphumanspccch,rccur-
rcnIncuraIncIs, hasonIyIhccompuIaIionaIcapaciIy o!aniIcsIaIc
auIomaIon. WcwiII rcIum IoIhis imporIanI qucsIion aI Ihc cnd o!
Ihis chapIcr.
Having dcncd Ihc naI sIaIc o! Ihc cvoIuIionary proccss IcIs go
backIo iIs bcginnng. Ihccmcrgcncco!rc!crcncc. Having Ihc abiIiIy
Io rc!cr impIics Ihc crcaIion o! a convcnIionaI associaIion bcIwccn
monoIiIhicsymboIicarIi!acIsandrcaIcnIiIicsmcdiaIcdbymcanings.
BuI whaI couId mcanngs bc in a worId o! prc-IinguisIic hunIcr-
gaIhcrcrs? In ChapIcr 6 iI wasargucdIhaIinboIhanimaIsandcarIy
humansmcanings arc IhcproIoIpcs IhaIncuraIncIscxIracI!rom
scnsorycxpcricncc, nonIinguisIic caIcgoricsorgcncraIizaIions sIorcd
asdisIribuIcdpaIIcrnso!acIivaIion.ThcscdisIribuIcdrcprcscnIaIions
arc associaIcd Io Ihcir rc!crcnIs dirccIIy bccausc Ihcy arc cxIracIcd
!rom rcpcaIcd cncounIcrs wiIh manyparIicuIar insIanccs o! a cIass
o! rc!crcnIs. In addiIion Io giving us an accounI o! Ihc associaIion
bcIwccn mcanings and rc!crcnIs ncuraI ncIs can aIso cxpIain Ihc
associaIion bcIwccn mcanings and sounds bccausc Ihc proccss o!
Iraining IhaI gcncraIcs Ihc disIribuIcd rcprcscnIaIions is o!Icn pcr-
!ormcd wiIh Ihc goaI o! producing a parIicuIar moIor ouIpuI. I! wc
Ihinko!IhismoIorouIpuIasinvoIvingIhcIarynxasasourccandIhc
vocaIcaviIyas aIIcrIhcnIhcnaIproducI is avocaIizaIionIhaIis
auIomaIicaIIy associaIcd wiIh Ihc disIribuIcd rcprcscnIaIion. Bcing
in posscssion o! an cxpIanaIion !or Ihc Ink bcIwccn mcanings and
rc!crcnIs and mcanings and sounds aII IhaI is Ic!I Io cxpIain is Ihc
cmcrgcncc o! Ihc Iink bcIwccn scunds und rc]crcnrs. This is hardcr
Ihan iI sccms bccauscprc-IinguisIic hunIcr-gaIhcrcrs couIdnoI rcad
minds, IhaI is, Ihcy did noI havc acccss Io cach oIhcrs disIribuIcd
I+V
Ib
NlLbN N b MUL1N
rcprcscnIaIons. Ths mpIcsIhaIcarIyhumans hadnochoccbuIIo
usc pubIcIy avaIabIc n!ormaIon Io convcrgc on a sharcd scI o!
sound-rc!crcnI assocaIons.
To cxpIorc Ihc cmcrgcncc o!Ihs sharcd scI wc nccd Io dcpIoy a
popuIaIono!ncuraIncIs sIuaIcd na spaccconIanngsmpIc rc!cr-
cnIs andIogvc Ihcmamnmum o! cmbodmcnI Iocnsurc IhaIIhc
agcnIs havc no acccss Io cach oIhcrs mnds. U !acI, ncuraI ncIs
arc noI sIrcIIy ncccssary. wc can usc any oIhcr mpIcmcnIaIon
o! proIoIypccxIracIon, usng vdco camcras and sIandard magc
proccssng so!Iwarc, !or cxampIc, as s donc n a smuIaIon caIIcd
scmicricdynumics.U IhssmuIaIonparso!agcnIs IakcIumspIayng
Ihc roIcso! spcakcrandIsIcnr.Thc agcnIsnhabIasmpIcworIdo!
gcomcIrc gurcs |Ihc poIcnIaI rc!crcnIs) and posscss Ihrcc smpIc
capacIcs. Ihcy can pcrccvc and caIcgorzc Ihc gcomcIrcaI gurcs,
Ihcy can ponI aI a gurc, and Ihcy can obscrvc n whaI drccIon
oIhcr agcnIs arc ponIng. Thcy arc aIso abIc Io producc arbIrary
symbo!c arI!acIs |random syIIabIc combnaIons) Io usc as words,
arI!acIsIhaIcan cIhcrbc nvcnIcd orbcaIrcadyB usc n Ihc com-
munIy. Thus,usngscmoIc dynamcs Io modcIIhc cmcrgcncco! a
sharcd scI o! sound-rc!crcnI assocaIons mpIcs onIy Ihc assump-
IonsIhaIprc-IngusIchunIcr-gaIhcrcrscouIdusc gcsIurcsIoachcvc
sharcd aIIcnIon and had Ihc wIngncss Io parIcpaIc n rIuaIzcd
Iasks, suchasIakngIumsIo voca!zc.
AscmoIcdynamcsmuIaIonun!oIdsnIhc!oIIowngway.AgcnIs
arcparcd aI rando Io aIIcmpI Io communcaIc. Thc agcnI whosc
IumIs Iospcak scIccIs a gcomcIrc ob|ccIas Ihc Iopco! convcrsa-
Ion,chccksIsmcmoryIo sccwhaIprcvoussymboIc arI!acIshavc
bccn assoaIcdwIhI, andpcks onc sncc aI Ihc bcgnnng o!Ihc
smuIaIon no arI!acI s avaIabIc I nvcnIs a ncw onc. Thc agcnI
pIayng Ihc roIc o! !sIcncr Ihcn chccks Is own mcmory Io scc ! a
prcvous assocaIonbcIwccnIhaIarI!acIandsomcrc!crcnIhasbccn
madcanduscsI

aIasIsrsIgucss.U IhcvcryrsIcncounIcrIcan-
noIndanyIhng soInvcnIsIsown sound. Thus, IhcsIarIngponI
!or Ihc smuIaIon s a sIuaIon n whch cach agcnI has Is own
dosyncraIc vocabuIary and Ihc !sIcncrs aIways !aI Io gucss Ihc
spcakcrsrc!crcnIaInIcnIons. BuI spcakcrs canaIsohcIpIsIcncrsby
ponIng Io Ihc nIcndcdrc!crcnI and Ihs can somcImcs rcsuII n a
MUL1LN1b N KlM1YL LNUL
corrccIgucssa!IcrscvcraIIraIs.ThsrandomgucssngcannoIbyIscI!
convcrgcIo asngIc sharcdvocabuIary!oraIIIhc poIcnIaIrc!crcnIs
buI convcrgcncc can bc achcvcd ! wc gvc agcnIs Ihc capacIy Io
kccpscorco!whaI parso!symboIcarI!acIsandrc!crcnIshavcIcdIo
communcaIvc succcss n Ihc pasI.` Whcn a spcakcr scarchcs Is
mcmory !ora prcvous arI!acIassocaIcdwIhIs nIcndcd rc!crcnI,
!orcxampIc,ImaydscovcrIhaIIhcrcarc scvcraIchoccswIhd!!cr-
cnI scorcs. Thc hsIory o!Ihc d!!crcnI aIIcmaIvc assocaIons, somc
o!whchhavc Icd n Ihc pasI Io morc corrccIgucssngs, src!IccIcd
n Ihosc scorcs soaIIIhc spcakcrhas Io do spkIhc onc wIhIhc
hghcsIscorc.ThcIsIcncrIooscarchcsamongcompcIngassocaIons
!or Ihc IcasI ambguous onc |Ihc onc wIh a sngIc rc!crcnI) and !
Ihcrc arc scvcraI I pcks Ihc onc wIh Ihc hghcsI scorc as Is rsI
gucss.IIcraIngaproccssnwhchagcnIsarcaIIcmpIngIogucss cach
oIhcrs rc!crcnIaI nIcnIons and hcIpng cach oIhcr rcpar brcak
downs n communcaIon cvcnIua!Iy Icads Io a sIabIc rcpcrIorc o!
sharcdsound-rc!crcnIassocaons.
ThcouIcomco!IhssmuIaIonsvaIuabIcbuIImIcd.Ihcsymbo!c
arI!acIsrcprcscnIonIyonc cIass o!IngusIccnIIcs, IabcIs orpropcr
namcs, whIc Ihc scI o! mcanngs s rcsIrcIcd Io ob|ccIs IhaI havc
acIuaIIybccncxpcrcnccd,noIIomcanngsIhaIcanbcdcncdcom-
posIonaIIy by oIhcr words. NcvcrIhcIcss, wIhn Is ImIcd goaIs
Ihc smuIaIons a succcss bccausc I shows how Ihc proccss o! In-
gusIccvoIuIoncouIdhavcgoIsIarIcd.TosmuIaIcIhcncxIsIcp,Ihc
concaIcnaIon o! IabcIs nIo smpIc monoIIhc scnIcnccs, Ihc agcnIs
musI bc sIuaIcd n a worId IhaI dcmands !rom Ihcm cngagng n
causaI rcasonng. In Ihc prcvous chapIcr wc saw IhaI prc-IngusIc
hunIcr-gaIhcrcrs wcrc capabIc o! masIcrng compIcx scqucnccs o!
manuaIopcraIons as parI o! Ihc producIon o! sIonc IooIs. Ths Iask
prcscnIcd Ihcm wIh Ihc opporIunIy Io Icarn abouI Incar causa!
rcIaIons by Ihc drccI obscrvaIon o! Ihc consIanI con|uncIon o! a
causc andIsc!!ccI. xprcssngIhsconsIanIcon|uncIonsymbo!ca!Iy
wouId dcmandnomorc IhanconcaIcnaIngIhcIabcIs!orIhc acIons
or cvcnIs acIng as causc and c!!ccI. Thc rcsuIIng symboIc arI!acI
wouIdnoIbcarcaIscnIcncc, snccIsmcanng wouId noI rcsuII!rom
Ihc composIon o! hc mcanngs o! Ihc componcnI IabcIs, buI I
wouId ncvcrIhcIcss bc capabIc o! bcn Iruc or !aIsc. OnIhc oIhcr
IbI
IbZ
NLbHN N b MUL1N
hand, ! sharcd pcrsona! obscrvaIonswcrcsu!ccnI Io !camabouI
causa!rc!aIonswhaIwou!dhavc bccnIhcmoIvaIon!or cxprcssing
Ihcmsymbo!ca!!y?
To answcr ILs qucsIon Ihc ncxI smu!aIon poscs Ihc !o!!owng
cha!!cngc. whaI ! Ihcrc wcrc causa! rcgu!arIcs IhaI cou!d noI bc
cas!y obscrvcd by pcrsons buI on!y by cnIrc communIcs? Wou!d
Ihc producIon o! arI!acIs cxprcssng gucsscs abouI co-occurrcnccs
o! cvcnIs mprovc Ihc chanccs IhaI ovcr scvcra! gcncraIons a!!
mcmbcrs o! Ihc communIy wou!d makc Ihc corrcI assocaIon?
Thc sIuaIoncon!ronIngIhc smu!aIcdagcnIswasnsprcdbya rca!
cxamp!c !romconIcmporaryhunIcr-gaIhcrcrs. Whcn Ihc occan Idc
s !ow a gradcnI o! conccnIraIcd proIcn |shc!!sh) !orms aI Ihc
bcach. l!Ihc agcnIs !vcd ncarIhc bcach Ihcy cou!ddrccI!yobscrvc
!owIdcsandknowwhcnIoco!!ccIIhc shc!!sh.BuI!Ihcy!vcd!ar
awayIhcywou!d havc Io usc anoIhcr obscrvab!c cvcnI IhaI corrc-
!aIcdwIha !ow dc, such asIhc phasc o!Ihc moon. As I happcns
whcnIhc moonsncwor!u!!IhcIdc rcachcs boIhshghcsIandIs
!owcsI poinIs, maxmzngIhc chanccso!nding shc!!sh. So a com-
munIy cou!d bcncI !co!!ccIvc obscrvaIons o!Ihc con|uncIon o!
Idc sIaIcsandmoon phascs cou!dbc cncapsu!aIcd nIo amono!Ihc
scnIcncc.Thcmcmbcrs o!IhscommunIywou!dnoIhavcIoposscss
anyknow!cdgc o!rca!mcchansms,IhaIs,IhcynccdnoIknowIhaI
whcn Ihc moon sncw or !u!!Is gravIaIona!!orcc smphasc wIh
IhaIo!IhcsunandIsc!!ccIonIhcsIaIc o!IhcIdcs grcaIcsI. OnIhc
oIhcrhand,Ihcrcsu!Iso!ChapIcr7 a!!ows IoassumcIhaIprc-!n-
gusIchunIcr-gaIhcrcrs cou!dpcr!orm pcrccpIua! ana!yscs o! sIua-
IonsusngscrpI-!kcsIrucIurcs,assgninIcracIngob|ccIsIoIhcro!cs
o! causa! agcnI and causa!paIcnI, andrcmcmbcr Ihosc cxpcrcnccs
usngcpsodcmcmory.
Wh!c Ihc symbo!carI!acIs musI havc Ihc capacIy Io bc Iruc or
!a!sc, a capaciIy IhaI proposIons havc, Ihcy nccd noI ncccssar!y
havc aproposIona!!orm sincc Ihc smu!aIcd agcnIs havcIo so!vc a
sng!c causa! prob!cm in Ihcr !vcs a smp!c numcrca! vocabu!ary
spcca!zcdIo rc!crIoIdc sIaIcs andmoon phascs W su!cc. Ncvcr-
Ihc!css, I W bc usc!u! !or us Io maginc Ihc symbo!c arI!acIs as
mono!Ihc scnIcnccs!kcIhs
Iu!!NoonCa uscsIowTdc
MUL1LN1b N KM1YL LNUL
U Ihsprccursoro! a rca!scnIcncIhc abscncc o! spaccsbcIwccn
wordsndcaIcsIhaIIhc!abc!s!or!u!!moon and !owIdc cannoI
bc uscd composIona!!y Io gcncraIc oIhcr scnIcnccs, wh!c Ihc vcrb
causcs smp!y marks Ihc !acI IhaI Ihc co-occurrcncc o! Ihc Iwo
!abc!s sIands !or Ihc co-occurrcncc o!Ihc cvcnIs p!aying Ihc ro!cs o!
causc andc!!ccI. AIanyraIc,bccauscnIhssmu!aIonIhcagcnIsarc
cnIrc!y dscmbodcd and arc noI sIuaIcd n a spacc n whch Ihc
rc!cvanI cvcnIs cou!d Iakc p!acc Ihc Icrms !abc! and cvcnI rc!cr
Io acIvaIon paIIcrns in ncura! ncIs supp!icd Io Ihc agcnIs by Ihc
cxpcrmcnIcr. ach agcnI conssIs o! Ihrcc ncura! ncIs, Iwo uscd Io
hand!c Ihc assocaIon bcIwccn !abc!s and Ihc cvcnIs Io whchIhcy
rc!cr,andoncuscdIocrcaIcanassocaIonbcIwccnIhccvcnIsIhcm-
sc!vcs. Thc dcsgno! Ihc Iwoncura!ncIs IhaI hand!c Ihc !ngusIc
parIo!IhcIaskcanbc vcrysmp!cbcausIhccmcrgcncco!a!cxcon
Io rc!cr Io cvcnIs has a!rcady bccn cxp!ancd by smu!aIons !ikc
scmoIc dynamcs. Norc spccca!!y, Ihc ncura! ncIs IhaI hand!c
!abc!s and Ihcr rc!crcnIs do noI nccd Iohavc hddcn !aycrs capab!c
o! cxIracIng proIoIypcs. P Ihcy nccdIo do sproducc a !abc! asan
ouIpuI!gvcnancvcnIasannpuIandvccvcrsa,produccancvcnI
!gvcna!abc!.ThcIhrdncura!ncIsarcgu!armu!I!aycrpcrccpIron
IhaI Iakcs cvcnIs as Is npuI andproduccs a causa! assocaIon as Is
ouIpuI. AIany oncImcnIhs smu!aIonacommunIyo!20 agcnIs
cxsIs, cach spcndingIs !!c !carnngaouI Ihc !ink bcIwccn moon
phascs andIdc sIaIcs !rom drccI ccrcncc and!romIhc mono-
!IhcscnIcnccsproduccdbyprorgcncraIons.AIIhccndo!Is!!can
agcnI dcs !cavng bchind a mono!ILc scnIcncc cxprcssng Is bcsI
gucssabouIIhc causa!!nkandancwunIrancdagcnIIorcp!accI.
BccauscIhcagcnIsnIhssmu!aIonarcdscmbodcdIhcrcspons-
biIIy!orrccrcaIingIhc dcsrcdsIuaIon!a!!sonhc Iranngproccss
Isc!!.!Ihcmu!I!aycrpcrccpIronwasIrancdusngon!yn!ormaIon
abouI cvcnIs Ihcn cu!Iurc wou!d noI maIIcr, and vcc vcrsa, ! Ihc
on!y Irainng cxamp!cs wcrc pasI symbo!c arI!acIs Ihcn Ihc cvcnIs
and Ihcr ob|ccIvc causa! connccIons wou!d noImakc a d!!crcncc.
SoIhc scIo!Iranngcxamp!csnccdsIondagoodba!anccbcIwccn
Ihc Iwo. On onc hand, I musI makc I hard !or anyagcnI Io !carn
Ihc assocaIon bcIwccn cvcnIs !rom drccI cxpcrcncc, mmckng
Ihc c!!ccIs o! c!oudy wcaIhcr, dsIancc, and oIhcr nIcr!crcnccs IhaI
in rca! !!c wou!d prcc!udc pcrsona! obscrvaIions o! Ihc consIanI
IbJ
Ib+
NlLbN N blMUL1lN
con|uncIionbcIwccnIuII or ncw moon and cxIrcmc Iidc variaIion.
OnIhcoIhcrhand,iImusIrc!IccIIhcaccumuIaIcdknowIcdgcoIpasI
gcncraIions soIhaI abias Ioward IrucarIiIacIsmusI bc addcd IoIhc
Iraning, abiascapIuringIhcIacIIhaIinarcaIcommuniIyIhcrcIaIivc
succcss oI agcnIs Irom prcvious gcncraIions wouId in!Iucncc Ihc
choicc oI gucsscs uscd as asIarIingpoinI byancw gcncraIion. WiIh
Ihis bias acIing as a scIccIionprcssurc cuIIuraI cvoIuIion arrivcs aI a
siIuaIioninwhichIhc cnIirccommunIycan rcIiabIyIcam abouIIhc
rcIaIionbcIwccnmoonphascs and IidcsIaIcs.
Thc ouIcomcs oI Ihis and Ihc prcvious smuIaIion suggcsI IhaI
monoIiIhicsymboIicarIiIacIs couIdhavc cmcrgcdnacommuniIy oI
hunIcr-gaIhcrcrsIopcrIormsmpIc rcIcrcnIiaI IncIions and cpIurc
simpIc causaI rcIaIions. This is noI in iIscII surprisngbccausc as wc
argucdnChapIcr7chmpanzccsposscssIhcabiIiIyIoassocaIcIabcIs
|coIorcd shapcs) and Ihcir rcIcrcnIs and Io chain scvcraI IabcIs
IogcIhcr Io cxprcss simpIccausaIrcIaIions, aIIhoughIhcy nccd Iobc
Irancd by humans Io dcvcIop Ihis abiIiIy and havc ncvcr bccn
obscrvcd cxcrcisng iI in Ihcir naIuraI habiIaI. BuI as was aIrcady
poinIcd ouI Ihis consIIuIcs Ihc parI oIIhcprobIcmIhaIis rcIaIivcIy
casy Io soIvc. Thc hard parI is Io smuIaIc a proccss IhaI can brcak
down a scI oI monoIiIhic scnIcnccs inIo rccursivcIy rccombinabIc
componcnIs, a proccss rcIcrrcd Io as jrummuriculizuricn. BcIorc dis-
cussing a smuIaIion IhaI IackIcs Ihis probIcm iI wiII bc uscIuI Io
dcscribc Ihc IradiIionaI accounI. InsIcad oI cxpIaining Ihc human
capaciIyIouscandundcrsIand rccursionbyposIuIaIngancvoIuIion-
aryproccss bascd onrcpIicaIngacousIicmaIIcr,aswcarcdoinghcrc,
Ihc IradIionaI cxpIanaIion assumcs IhaI Ihis capaciIy was produccd
bybioIogicaIcvoIuIion.
This accounI sIarIs by crcaIing a IormaI modcI IhaI can producc
scnIcnccsIhrougharccursivcproccss. ThcmodcIisbascdonrcwriI-
ing ruIcs, opcraIors IhaI Iakc as Ihcir inpuI sIrngs oI inscripIions,
rcwrIc Ihcm, and producc a ncw sIring as Ihcir ouIpuI. Thc sIrngs
arc noI random chains oI sounds |or IcIIcrs) buI scqucnccs aIrcady
assigncdIogrammaIicaIcaIcgorics, Iikcnoun, vcrb, scnIcncc, and so
on. BoIh Ihc rcwriIing ruIcs and Ihc grammaIicaI caIcgorics arc
assumcdIobcinnaIcconsIiIuIingaunivcrsaIgrammarcommonIoaII
Ianguagcs. '" Thc innaIc hypoIhcsis is dcIcndcd onIhc grounds IhaI
whcnchiIdrcnIcarnaIanguagcIhcyacquircacapacIyIoproducc an
MUL1LN1b N KM1YL LNUL
inniIc numbcr oI scnIcnccs cvcn Ihough Ihcy arc onIy cxposcd
Io Ihc vcry smaII numbcr oI scnIcnccs acIuaIIy produccd by Ihcir
parcnIs.In oIhcrwords, IhcpovcrIyoIsImuIaIioncharacIcrizingIhc
IcarnngsiIuaIion isIakcnIobc cvidcnccIhaI chIdrcn donoI Icarna
grammarbymakngsIcp-by-sIcpnIcrcnccs IromaduII scnIcnccsbuI
raIhcr IhaI Ihc IaIIcr acI as mcrc Iriggcrs Ior ruIcs and caIcgorics
inhcriIcd bioIogicaIIy. To puI Ihis diIIcrcnIIy, H as auIomaIa Ihcory
shows Ihc spacc oI possibIc Ianguagcs is inniIc in sizc how can
chiIdrcn Icarn a Ianguagc givcnIhc povcrIy oI Ihc sIimuIaIion Ihcy
rcccivc Irom Ihcir parcnIs? II wouId sccm IhaI wc wouId nccd Io
posIuIaIc a way oIconsIrainngIhis spaccIomakc Ianguagc acquisi-
Iion noI |usI possibIc buI highIy probabIc. A univcrsaI grammar
isprcciscIy suchaconsIrainI. iI cIiminaIcsIromIhcspacc oIpossibiIi-
Iics IhaI Ihc chiId musI conIronI aII cxccpI Ihc humanIy IcarnabI

Ianguagcs.
I l
Thc IradiIionaI cxpIanaIion runs inIo diIcuIics whcn Irying Io
spcciIyIhccvoIuIionaryproccssIhaIIcdIoIhcdcvcIopmcnIoInnaIc
ruIcsandgrammaIicaIcaIcgorics.ThisisnoI|usIaqucsIionoIcxpIain-
ing Ihc survivaIvaIuc oI rccursion givcnIhaIIinguisIic communica-
Iioncanbc achcvcdwiIhouIiI. 1isaIsoIhcprobIcm oIshowingIhaI
gcncIic variaIion Ior Ihc ncuraI subsIraIc oI caIcgorics Iikc noun
and vcrb cxisIcd.Thc aIImaIivcaccounI, onIhcoIhcrhand, musI
noIonIycxpIainhowIanguagccvoIvcdcxIcrnaIIydrivcnbyiIscapac-
iIy Io cncodc proposiIionaInIormaIion abouI causc and cIIccI rcIa-
Iions. buI in addiIion iI musI cxpIainhowiI wasIaIcrinIcmaIizcd.
lJ
Thc sccond rcquircmcnI is ncccssarybccausc wc nccd Io accounI
IorIhccxisIcnccoIbrainorgans spcciaIizcdIorIanguagc, organsIhaI
can acI as nhcriIabIc consIrainIs on Ihc spacc oIpossibIc Ianguagcs.
Thc Iwo aspccIs oI Ihis aIIcrnaIivc accounI havc bccn succcssIuIIy
invcsIigaIcd using compuIcr smuIaIions. grammaIicaIizaIion has
bccn shown Io cmcrgc IhroughcuIIuraIcvoIuIionin groups oIinIcr-
acIing agcnIs inwhich onc gcncraIion pIays Ihc roIc oI Icachcr and
Ihc oIhcr onc oI sIudcnI: andIhc mcchanism oI inIcrnaIizaIion, Ihc
so-caIIcdBuldwinc]ccr. has bccn shownIobcviabIc insimuIaIionsoI
cvoIving ncuraI ncIs. Thc rsI simuIaIion docs noI havc Io producc
Ihc compIcx Iypc oI rccursion IhaI characIcrzcs conIcxI-scnsiIivc
Ianguagcs buI onIy Ihc smpIcr Iypc oI conIcxI-Ircc Ianguagcs. Wc
canmagnc IhaIonccsucha smpIc rccursivc Ianguagc cmcrgcd in
Ibb
Ib
NLbN N bMUL1N
ancicnI hunIcr-gaIhcrcr communiIics Ihc BaIdwin cIIccI bcgan Io
opcraIcandIhaIIhc rcmaindcr oIIhc|ourncyIhroughIhcspacc oI
possibIc IanguagcsIook pIacc by cvoIuIionary scarch bascd on boIh
gcncIicandIinguisIicrcpIica Iors.
Thc agcnIs uscd in Ihc simuIaIion oI gr aIicaIizaIion arc cvcn
morcdiscmbodicdIhanIhocoIIhcsimuIaIion|usIdiscusscd.Ipar-
IicuIar,IhcagcnIshavcacccssIoIhcinIcndcdmcaningsoImonoIiIhic
arIiIacIs. Sincc agcnIs arc supposcd Io Icarn Irom obscrving cach
oIhcrs bchavior noI by rcadDg cach oIhcrs minds Ihis is a ma|or
IimiIaIion. I addiIion, Ihc simuIaIion docs noI usc ncuraI ncIs buI
sIayswiIhinIhcworIdoIauIomaIa, pushdownauIomaIain Ihc casc
oI conIcxI-Ircc Ianguagcs, and Ihc noIaIion Ihcy undcrsIand. Thus
Ihcmcaning oIamono!iIhicscnIcnccIikc.
IuIINoonCauscsIowTidc
isaproposiIionIhaIis cxprcsscdin auIomaIon noIaIion as.
Causcs |IuIINoon,IowTdc)
This choicc is |usIicd on Ihc basis IhaI using symboIic rcprcscnIa-
IionsIaciIiIaIcsIrackingIhcprogrcssivccvoIuIionoIcomposiIiona!iIy
andrccursion. chcckingIhaIancwsynIacIicaIIcaIurchas cmcrgcdis
simpIyamaIIcroIIookingaIIhccurrcnI rcpcrIoirc oIproposiIionsin
IhcpopuIaIionoIagcnIs.'cruciaIcomponcnIoIIhcproccssIhrough
whichcomposiIiona!iIycmcrgcsisIhcabiIiIyoIIhcagcnIsIogcncraI-
izcIrom cxpcricncc. In a simuIaIionusingncuraIncIsIhiswouIdbc
impIcmcnIcd by Ihc cxIracIion oIproIoIypcs Irom scnsory sIimuIa-
Iion. InIhc prcscnI casc, onIhc oIhcrhand, iI is casicrIoimpIcmcnI
IhiscapaciIyIhroughan inducIivcaIgoriIhmspcciaIizcdIoopcraIcin
Ihc worId oIpushdown auIomaIa. IinaIIy, Ihc agcnIs invoIvcd musI
bcabIc Io inIcracIpIaying Ihc roIc oI Icachcr or sIudcnI, an inIcrac
Iion IhaI in Ih casc oI prc-IinguisIic hunIcr-gaIhcrcrs wouId havc
IakcnIaccbcIwccnparcnIsandoIIspring, orbcIwccnaduIIsbcIong-
ing Io an cxIcncd IamiIy group and kindrcd chiIdrcn. uring Ihis
inIcracIion IhcsIudcnIshavc dirccI acccss IoIhc mcaning oIdccIara-
IivcscnIcnccs,IhaIis, IhcIcachcrprcscnIsIhcmwiIhpairs oIpropo-
siIions and IhcirmonoIiIhicsymboIic cxprcssions. To |usIiIy Ihc usc
oImindrcadingwcwouIdhavcIoassumcIhaIinanoIhcrvcrsionoI
MUL1LN1b N KM1YL LNUL
Ihis simuIaIion, a vcrsion in which agcnIs wouId bc cmbodicd and
siIuaIcdinaspaccconIainingIhcrcIcrcnIsoIIhcIabcIs, Ihcproposi-
IionswouIdbcrcconsIrucIions oIinIcndcdmcaningsIhaIIhc agcnIs
wouIdarrivcaI asIhcvobscrvcd cach oIhcrsinIcracIionswiIhIhosc
rcIcrcnIs. WcwiII rcIurn Io Ihc qucsIion oI how Io cIiminaIc mind
rcadingbcIow.
ThcworIdinwhich IhcagcnIsIivcisxcdand simpIc.U a IvpicaI
run, Ior cxampIc, Ihc simu!aIion incIudcs as rcIcrcnIs Ior Ihc IabcIs
vc ob|ccIs and vc acIions. CivcD IhaI cach proposiIion has Iwo
argumcnIs, such as IuII moon and Iow Iidc, and IhaI Ihcsc musI
bc diIIcrcnI Irom cach oIhcr, Ihis vicIds I00 possibIc proposiIions.
AgcnIs spcnd haII Ihcir IiIc as sIudcnIs oI Icachcrs bcIonging Io Ihc
prcvious gcncraIion, Ihc oIhcr haII as Icachcrs Io a ncw gcncraIion.
ToincIudcIhccIIccIoIIhcpovcrIv oI sIimuIaIion cachagcnI Icachcs
onIvhaIIoIIhcpossibIcproposiIionsiniIsIiIcIimc. WhcnIhcsimu-
IaIionsIarIsaIIsvmboIicarIiIacIsarcmonoIiIhicandIcachinginvoIvcs
makingIhcsIudcnImcmorizcaIong!isIoIassociaIions.IromIhcIacI
IhaI IhcscociaIions arc IaughI as cusIomarv, howcvcr, sIudcnIs
caninIcrIhcsimpIcruIcIhaIaproposiIionIikc.
Causcs |IuIINoon, IowTdc)
bcIongsIoIhccaIcgorvscnIcnccandIhaIiI hasIobccxprcsscdas.
IuIINoonCauscsIowTdc.
ThisruIccanbccxprcsscdinanoIaIionIhaIapushdownauIomaIon
wouId undcrsIand as.
SlCauscs |IuIINoon, IowTdc) IuIINoonCauscsIowTdc
In which Sl mcans bcIongs Io Ihc scnIcncc caIcgorv and Ihc
arrowmcans musIbc cxprcsscd as. AIIhispoinIIhcsvmboIicarIi-
IacIs uscd IorIcaching Ihc prcvious scnIcncc as wcII asIhc diIIcrcnI
scnIcncc.
IuIINoonCa uscsHighdc
arcIrcaIcdasHIhcIacIIhaIIhcvboIhincIudcIhcIabcIIuIINoon docs
noImaIIcr. achmonoIiIhicscnIcncc has Iobc Icamcd indcpcndcnIIv
and cach has Io bc associaIcd as a whoIc wiIh iIs own proposiIion.
Ib
Ibd
NLbN N b MUL1N
BuI IhcnIhcgcncraIzing abiIIics oI Ihc sIudcnIs comcinIo acIion.
Ihc common parI oI Ihcsc Iwo scnIcnccs is dcIccIcd and Ihc Iwo
proposiIions arcrcpIaccdbyamorc gcncraI onc.
Cascs |IuIINoon, X)
IogcIhcrwiIhanassignmcnIoIIhccaIcgorynounIopossibIcvaIucs
oI Ihc variabIc X. Whcn Ihc sIudcnI IhaI madc Ihc gcncraIzaIion
bccomcs a Icachcr iI docs noIIcachIhc Iwo originaIassociaIionsbuI
Ihcncw onc.
SlCauscs |IuIINoon, NlX) IuIINoonCauscsX
AI Ihis poinI Ihc povcrIy oI sIimuIaIion characIcrizing Ihc Icarning
siIuaIion sIarIs Io work in Iavor oI Ihc morc gcncraI Iorms bccausc
symboIicarIiIacIsIhaIarcuscdmorcIrcqucnIIyIcndIobcovcrrcprc-
scnIcdu IhcimpovcrishcdsampIcuscdIorIcaching.InoIhcrwords,
a scnIcnccIkc IuI!oonCauscsX is uscdIwiccas oIIcnasIhcIwo
symboIicarIiIacIsiIrcpIaccssincciIcan cxprcssIhcmcaning oIboIh,
soIhcprobabiIIyIhaIiIwiIIbcscIccIcdasaIcachingcxampIcisIwicc
ashigh. IronicaIIy, IhcsamcpovcrIyoIIhcsIimuIusinvokcdas a|us-
IihcaIion Ior a univcrsaI grammar works hcrcIocIiminaIc Ihcnccd
Ior innaIc ruIcsby acIing as a boIIIcncck Iavoring Ihc Iransmission
oImorc gcncra!IormsIoIuIurcgcncraIions. ' BuIiI aIso Icads IoIhc
sIarIoI Ihc scgmcnIaIion oIIhc monoIIhic arIiIacIs sincc Ihc morc
gcncraIIormsarcaIsoIhconcsIhaIdispIaycomposiIionaIiIy. U oIhcr
words, IhcboIIIcncckacIs as a sorIingdcvicc, Iavoringsomc rcpIca-
Iors aI Ihc cxpcnsc oI oIhcrs, whiIc aIso crcaIingIhccondiIions Ior
Ihc IurIhcr scgmcnIaIion oI monoIIhic scnIcnccs. U IacI, a!I Ihc
simuIaIions discusscd in Ihis chapIcr, as wcII as oIhcr rcIaIcd oncs,
opcraIcbyusingaboIIIcncckasascIccIionmcchanism. ' InscmioIic
dynamics, IorcxampIc,Ihc originaIpIuraIIyoIidiosyncraIicvocabu-
Iarics is sIowIy narrowcd down Io a singIc sharcd Icxicon by Ihc
IcndcncyoIagnIsIochooscsymboIcarIiIacIsaccordingIohowrcI-
crcnIiaIIysucccssIuIIhcyhavcbccninIhcpasI.AndsimiIarIyIorIhc
simuIaIion oI IabcI concaIcnaIion as a mcans Io cxprcss causaIiIy.
cvcry gcncraIionhas a IcndcncyIo sIarI wiIhIhoscmonoIiIhicscn-
IcnccsIhaIwcrcmorcprcdicIivcoIIhcrcaIIinkbcIwccnmoonphasc
and Iidc sIaIc, andIhis acIs a boIIIcncck IaciIiIaIing Ihcpassagc oI
somcbuInoIoIhcrarIiIacIs IoIhcncxI gcncraIion.
MUL1LN1b N KM1YL LNUL
AsancxampIcoIhowIhcgrammaIicaIzaIionboIIIcncckcouIdIcad
Io IurIhcr scgmcnIaIion IcIs imaginc IhaI Ihc sac proccss |usI
dcscribcdwasappIcdIoIhc monoIIhicscnIcnccs.
IuI!oonCauscsIowTidc
NcwNoonCauscsIowTidc
andIhaIIhcrcsuIIingmorcgcncraIproposiIion, Causcs |Y,IowTidc) ,
wascomparcdbyasIudcnIwiIhIhcprcviousonc.
Causcs |IuIINoon, X)
Causcs |Y. IowTdc)
Thc sIudcnImayIhcnrcaIzcIhaIIhcIabcIinIhchrsIproposiIionis
apossibIcvaIucIorIhcvariabIcuIhcsccondoncandviccvcrsa, and
produccby gcncraIizaIionIhcabsIrcIproposiIion.
Causcs |Y, X)
AIIhis poinI, a monoIIhicsymboIicarIiIacIhas bccncIIccIivcIybro-
kcndowninIopicccsIhaI arcnow rcadyIocnIcrinIoparIIo-whoIc
rcIaIions Io composc oIhcr scnIcnccs. NoI onIy havc Ihc IabcIs Ior
cvcnIs bccn assigncd Io Ihc caIcgory noun buI Ihc prcviousIy
impIiciIprcdicaIc Causcs has bccn isoIaIcd and can bc assigncd Io
Ihc caIcgory vcrb. U oIhcr words, composiIionaIiIy has bccn
achicvcd and grammaIicaI caIcgoricshavccmcrgcdin apurcIycuI-
IuraI cvoIuIionary proccss. BuI whaIabouI Ihc oIhcr rcquircd com-
poncnI oI a grammar, rcwriIing ruIcs IhaI can bc uscd rccursivcIy?
AsimpIcIormoIrccursion couId cmcrgc iIIoIhcprimiIivcprcdicaIc
CauscsIhaIIakcs cvcnIsasiIsinpuIswcaddaprcdicaIcIkc Says
Iaking proposiIions IhcmscIvcs as inpuIs. ' Thc ncw prcdicaIc can
gcncraIc proposiIions IikcIhcsc.
Says|TadiIion, Causcs|IuIINoon,IowTidc) )
Says|NyTcachcr, Causcs|IuIINoon, IowTdc) )
OrinngIsh, TadiIion says IhaIIuIImooncauscs Iow Iidc or Ny
IcachcrsaysIhaIIuIImooncauscsIowIidc. ThcscarcIhckindoIprop-
osiIionsIhaIwou!darisc sponIancousIyinaIcachingsiIuaIionwhcn-
cvcrIhcconIcnIoIwhaIisbcingIaughImusIbc|usIihcd byascrbing
iI Io a IadiIion or Io Ihc now dcad Icachcr oI Ihc currcnI Icachcr.
IbV
I
NLbN N bMUL1N
ThcIypc o!rccursonnvoIvcds,o!coursc,vcrysmpIc,buIIhaIsaII
wc nccdcd n Ihc prcscnI conIcxI. Oncc an cxIcmaI Ianguagc had
acqurcd Ihs mnmaI ompIcxIy I couId sIarI cocvoIvng wIh Is
boIogcaI subsIraIumIhroughIhcBaIdwnc!!ccI.
ThcBaIdwnc!!ccIsnoIa gcncraImcchansm!orIhc Irans!orma-
Iono!Icamcd capacIcsnIonhcrIabIc oncs. Such a gcncraImcch-
ansmdocsnoIcxsI.KaIhcr,IsamcansIogvcgcncIcadapIaIons
IhaI havc IIIIc survvaI vaIuc n IhcmscIvcs a chancc Io ncrcasc n
vaIuc by gcIIng parIaIcrcdI!or Ihcr conIrbuIonIo Ihc Incss o!
a gvcn gcncraIon. A Icamcd capacIycanbccomc nhcrIcdonIyn
vcry spccaI crcumsIanccs. I musI aIrcady havc a gcncIc bass buI
onc IhaI docs noI!uIIy dcIcrmnc I I musIdrccIIyncrcasc rcpro-
ducIvc succcss IohcIpIsassocaIcdgcncspropagaIc andIhspropa-
gaIonmusIbc suchIhaIwIhouIIhc acIuaIcxcrcsc o! Ihc Icamcd
capacIy Ihc gcncs wouId bc cImnaIcd by naIuraI scIccIon. Onc
waynwhchIhc cxcrcsc o!acapacIycana!!ccIboIogcaIcvoIuIon
s!IscuIIuraIIyIransmIIcd!rom onc gcncraIonIo anoIhcrgvng
I Ihc ncccssary IcmporaI sIabIIy Io havc rcIcvanI c!!ccIs. ' Thc
capacIy n qucsIon musI aIso prccxsI Ihc cmcrgcncc o! IngusIc
communcaIon andbc co-opIcd!orIhspurposc. Thc pcr!ccI cand-
daIc wouId bc Ihc abIIy Io cxpIoI scqucnIaI dcpcndcnccs n Ihc
opcraIonsnvoIvcdnsIoncIooImanu!acIurc. ThsabIIyddhavc a
parIaIgcncIcbass, IwasIransmIIcdacross gcncraIons, andI had
IobcacIuaIIycxcrcscdIompngconrcproducIvcsucccss.IIs cxIcn-
sonIoscqucnccs o! sounds, onIhc oIhcrhand, msIhavc bccn aI
rsIaIcamcdcapacIy. owIhscxIcnsonIookpIaccssuggcsIcdby
Ihc opcraIon o!rccurrcnIncuraI ncIs. Thcsc canbc uscd Io cxIracI
IcmporaI dcpcndcnccsn scqucnccs o! manuaI opcraIons aswcIIas
n scrcs o!vocaIzaIons. In !acI, a smpIc rccurrcnI ncuraI ncI, onc
wIhIhc samc compuIaIonaIcapacIyasIhcsmpIcsIauIomaIon,can
IcarnhowIoprcdcIIhcncxIwordnascnIcncc cvcn!IhcscnIcncc
has cmbcddcd Iauscs. Ths docs noI conIradcI Ihc dca IhaI nIc
sIaIc auIomaIa cannoI handIc rccursonbccausc unIkc apushdown
auIomaIonIhcncuraIncIdocs noIhavc IobcabIcIo IcIIwhcIhcror
noI a scqucnccbcIongsIoa conIcxI-!rcc Ianguagc. AIIIhasIo do s
dcIccI sIaIsIcaI paIIcms n a sampIc o! scqucnccs and cxIracI Ihc
probabIIyIhaIa gvcnwords a IkcIy conInuaIon o! a parIcuIar
scqucncc. '"
MUL1LN1b N KM1YL LNUL
ThcnIcracIono!IcarnngandnhcrIanccnvoIvcdnIhcBaIdwn
c!!ccIcanbcundcrsIoodIhroughIhc|onIusc o! gcncIc aIgorIhms
andncuraIncIs. Thcrc arcIwo cxIrcmc cascs o!Ihs combncd usc. a
gcncIc aIgorIhm can bc uscd Io nd Ihc naI wcghIs o! a ncuraI
ncIsconnccIons,nwhchcascIhcrcsnoroom!orIcarnng, oruscd
Iospcc!y onIy ccrIanpropcrIcs o!IhcncuraIncI |Ikc Is connccIv-
Iy or Ihc nIaI vaIucs o! Ihc wcghIs) aIIowng Ihc naI scIIng o!
Ihc wcghIs Io bc donc by Iranng." A smuIaIon o! Ihc BaIdwn
c!!ccI musI sIarI wIh Ihc sccond casc, nwhch a Icamcd abIIy s
onIyparIIygcncIcaIIydcIcrmncd, andcndwIhIhc rsIcasc, a!uIIy
gcncIcaIIy spcccdncuraI ncI. H onc smuIaIon cxIrcmcIy smpIc
ncuraI ncIs wcrc uscd, much Ioo smpIc Io bc usc!uI n Ianguagc
proccssng, buI Ihc probIcm IhaIhadIo bc soIvcd Ihrough Ihc com-
bncdusc o!IcarnngandgcncIcshadonIyoncsoIuIon.Thsmcans
IhaIIhc sIrucIurc o!Ihc scarchspacc hadIhc !orm o! a nccdIc n a
haysIack, wIh Ihc onIy good soIuIonrcprcscnIcdby a hgh Incss
pcak surroundcd by Iow Incss ponIs. WIhouI Ihc avaIabIIy Io
Icarn!rom cxpcrcncc Ihc IncsspcakcorcspondngIo Ihc soIuIon
was so sIccp IhaI I was bascaIIy mpossbIc Io cImb. InIroducng
Icarnng, on Ihc oIhcr hand, changcd Ihc shapc o! Ihc scarch spacc
surroundngIhc pcakwIhagcnIIcsIopcIhaI!acIIaIcdcvoIuIonary
cImbng
In our casc, Ihc nccdIc n a haysIack wouId bc Ihc probIcm o!
gcncraInga!uIIIanguagc, IogcIhcrwIhspccaIzcdbranorgans!or
producngandparsngIsscnIcnccs,usngasasIarIngponIancxIcr-
naIIycvoIvcdIanguagcposscssngonIyamnmumo!composIonaI-
Iy and rccurson. Bc!orc dscussng how Ihs proccss couId havc
un!oIdcdwc nccd Io addrcss aImIaIono! Ihc prcvous smuIaIon.
Ihc assumpIon o! drccI acccss Io proposIonaI conIcnI. ThaI s, wc
nccdIorcpIaccIhccapacIyIorcadmndswIhIhcn!crcncco!com-
muncaIvcnIcnIons!romIhcobscrvaIono!bchavor.Wcargucdn
prcvous chpIcrs IhaI prc-IngusIc humans sharcd wIh brds and
oIhcr mammaIs Ihc abIIyIo anaIyzc scnsory cxpcrcnccnIcrms o!
compIcx sccncs n whch ob|ccIs wIhcndurng dcnIIcs pIay roIcs
o! agcnIs andpaIcnIs, andIhaIIhcysharcdwIhoIhcrprmaIcsIhc
abIIyIodcIccIIcndcnccsIocoopcraIcorchcaInsocaInIcracIons.
UndcrsIandng bchavoraI roIcs and dcIccIng bchavoraI Icndcnccs
snoI,o!coursc, IhcsamcIhngasgucssngnIcnIonsbuIIhc abIIcs
II
IZ
NLbN N b MUL1N
invo!vcd cou!d havc bccn !urIhcr dcvc!opcd in IhaI drccIion oncc
smp!c rc!crcncc andcausa!prcdcaIon cmcrgcd.
Thc ab!Iy Io acLcvc sharcd aIIcnIon by Ihc usc o! a common
scIo!soundrc!crcnIassocaIonscou!dhavc!cdIoIhcab!IyIon!cr
nIcnIons ! Ihc ob|ccIs rc!crrcd Io had a capacIy Io a!!ccI and bc
a!!ccIcd byIhc agcnIs, ! Ihc rc!crcnI was a prcdaIor a!!ordng Ihcm
dangcr or a prcy a!!ordng Ihcm nuIrIon, !or cxamp!c. ln oIhcr
words, Ihc inIcnIon Io warn oIhcrs o! poIcnIa! opporIunIcs and
rskswou!dbccasyIogucss!romIhcproducIono!asound!Ihc!aI-
IcrIookp!accnasIuaIonIowhcha!!agcnIsaIIrbuIcdsgncancc.
Andsmi!ar!y !or Ihc symbo!c arIi!acIs dcnoIingIhc con|uncIon o!
causcs andc!!ccIs. l! Ihc producIon o! Ihc sound !or Iu!!Noon-
CauscsIowdc had acqurcd Ihc sIaIus o! an urgcnI ca!! Io go Io
IhcbcachIo!oragc !orshc!!sh, agcnIswou!dcomcIoin!crIhccus-
IomaryinIcnIonbchindIhcproducIiono!IhaIsound.Thcscab!cs,
primIvc as Ihcy arc, cou!dhavc su!ccd Io rcp!acc Ihc c!cIprc
scnIaIono!proposIona!conIcnIina!camngsIuaIon!Ihc Icach-
ngo!!ingusIcassocaIonshadIakcnp!accsparIo!Ihcpcr!ormancc
o!non!ngusIcIasks, such ashunIing, gaIhcring, or working sIonc.
On Ihc oIhcr hand, oncc mono!ILc arI!acIs bcgan Io bc brokcn
down a ncw Iypc o! obscrvab!c bchavor bccamc avai!ab!c. wcrd-
chcicc bchuvicr. m parIcu!ar, words !kc !u!! moon and !ow Idc
IhaI Icndcd Io occur IogcIhcr morc o!IcnIhan in Ihc company o!
oIhcr words wou!d havc crcaIcd paIIcrns in Ihc !rcqucncy o! co-
occurrcncc IhaI wcrc dcIccIab!c n Ihc choccs o! words madc by
agcnIs.Thcsc arc cxacI!yIhckndo!sIaIsIca!paIIcmsIhaIcanbc
cxIracIcd!romapopu!aIon o!scnIcnccsby cvcnIhc smp!csIrccur-
rcnIncura!ncIs.
IromIhcnonIhchsIoryo!!anguagccanbcimagncdIohavcpro-
cccdcdonIwo!cvc!saIIhcsamcImc.bo!ogca!cvo!uIonconInucd
IoopcraIcIhroughIhcBa!dwnc!

ccIhc!pngIopropagaIc Ihc gcncs

undcr!yng Ihc capacIy Io dcIccI paIIcms o! word co-occurrcncc,
wL!ccu!Iura!cvo!uIonbcganIoIrans!orm cusIomarypaIIcrns inIo
ob!gaIoryconsranIsonword-chocc bchavor.Wh!ccar!yoninIhc
cu!Iura! cvo!uIonary proccss Ihc Icachcr-sIudcnI inIcracIon may
havc invo!vcd on!y Ihc Iransmsson o! paIIcms IhaI had bccomc
abIua!, !aIcronIhcpaIIcmscou!dhavcbccnIransmIIcdas!Ihcy
MUL1LN1b N KM1lYL LNUL
consIiIuIcdan cn!orccab!csoca!ob!gaIon.ThaIs, awordIhaIuscd
Io rcqurc anoIhcr onc smp!y bccausc Ihc Iwo Icndcd Io co-occur
vcry o!Icn now ob!gcd Is uscr Io provdc Ihc sccond word. Ths
way cu!Iura! cvo!uIon cou!d havc provdcd a mcchansm !or Ihc
cmcrgcncc o! !ingusIc ru!cs, IhaI s, amcchansmIonsIIuIona!zc
habIua! usagc inIo an ob!gaIory convcnIon. Thc bass !or such a
mcchansm wou!d havc a!rcady bccn ava!ab!c !, as was argucd in
ChapIcr 8, Ihc habI o!punsLngIhosc IhaI donoIpunshdcparIurcs
!rom coopcraIvc bchavor |mcIanorms) had prcvous!y cmcrgcd.
Oncc mcIanorms had bccomc parI o! Ihc cu!Iura! conIcnI IhaI was
IransmIIcd !rom onc gcncraIon Io Ihc ncxI Ihcr usc cou!d havc
bccn cxIcndcd I paIIcrns o!bchavorIhaI dsIingushcd onc com-
munIy!romanoIhcr.moIhcrwords,mcIanormscou!dhavcbccomc
a mcchansm noI on!y!orIhccmcrgcncc o!so!darIyand a scnsc o!
communa!idcnIIybuIa!so!orIhccn!orccmcnIo!bchavora!paIIcms
IhaIwcrc unquc IoacommunIyandconso!daIcdIhaIdcnIIy.
A mcchansm Io Irans!orm habI inIo convcnIon s an mporIanI
componcnI o! Ihcorcs o! non-bo!ogca! ngusIc cvo!uIon aI hc
!cvc! o!boIh synIax and scmanIcs. Thc way synIax cou!d havcbccn
a!!ccIcd by nsIIuIona!zaIon can bc shownby gongback Io Ihc
prcvous cxamp!c o! Ihc proposIon Causcs |Y, X) . Oncc such an
absIracI proposIon had cmcrgcd and Ihc cusIomay co-occurrcncc
o! a prcdcaIc and Is argumcnIs had bccomc a convcnIona! con-
sIranI, Ihc paIIcrn Isc!! cou!d havc bccn uscd as a Icmp!aIc |Ihc
opcraIor-argumcnI Icmp!aIc) !or many oIhcr crcssons. m oIhcr
words, Ihc paIIcm cou!dhavc bccn uscdbyana!ogyIoconsIranIhc
crcaIon o!ncwpaIIcmsIhcrcsu!IbcngndsIingushab!c !rom scn-
IcnccsproduccdbyIhcapp!caIon o!grammaIca!ru!cs. Thc c!!ccI
o!nsIIuIona!zaIonon scmanIcs, nIu, L rc!aIcd IoIhc !acIIhaI
whcnIwowordsIcndIoco-occurwIhvcryLgh!rcqucncy, soIhaIa
!sIcncr can a!rcady Ic!! whaI Ihc sccond word w!!bc oncc Ihc rsI
onc s hcard, Ihc sccond word canbc omIIcd!roma scnIcncc. ThaI
s,vcryLgh!rcqucncyo!cooccurrcncccou!dhavcmadcsomcwords
rcdundanI !or a spcakcr sncc Ihcy cou!dbc supp!cd by Ihc !sIcncr.
m somc cascs Ihs cou!d havc a!!owcd cnIrc phrascs or cvcn scn-
IcnccsIobc compacIcdnIoasng!cwordwh!cprcscrvngIhcmcan-
ng o! Ihc phrasc or scnIcncc. Whcn Ihc producI o! packng Ihc
IJ
I+
NLbN N b MUL1N
composIcmcanngo!scrcso!wordswasmadcob!gaIoryapowcr!u!
mcchansm!orIhccxIcnson o!vocabu!ary, andcvcn!orIhcproduc-
Iono!ncwcaIcgorcso!words, wou!dhavc comcnIocxsIcncc.
U Ihssccnaro scorrccIwc canconc!udcIhaIIhcpossbi!Iyspacc
IhaI was scarchcd Ihrough cvo!uIon, boIh bo!ogca! and cu!Iura!,
was somorphcwIhbuInoI dcnIca! IoIhc spacc sIudcdbyauIom-
aIa Ihcory. AuIomaIa Ihory charIs Ihc spacc o!possb!c !anguagcs
and o! Ihc compuIaIona! capacIcs nccdcd Io masIcr Ihcm n an
cnIrc!y mcchansm-ndcpcndcnI way nvo!ving no commImcnI Io
anyparIcu!arimp!cmcnIaIono!IhcauIomaIaIhcmsc!vcs.ThsponI
s o!Icn ovcr!ookcd by Ihosc who Ihnk our brans musI cmbody a
gcncIca!!y codcd !incar-boundcd auIomaIon using an cxp!cI con-
IcxI-scnsIvc !anguagc o!IhoughI. Whcn I comcsIoIhc dcsgn o!
!uIurc smu!aIons o!

ingusIc cvo!uIon kccpng Ihc dsIncIon

bcIwccn mcchansms and mcchansm-ndcpcndcnI sIrucIurc rm!y
nmndcanhavcimporIanI conscqucnccs.mparIcu!ar, usc!u! asIhc
|usI dscusscd smu!aIon o!grammaIca!zaIonsIosIudyIhc dccom-
posIon o! mono!Ihc symbo!c arI!acIs, pcr!ormng I n a !ormaI
IhaI auIomaIa can undcrsIand rcduccs Ihc dcgrcc Iowhch Ihc paI-
Icrns IhaI rcsu!I can bc consdcrcd Io bc Iru!y cmcrgcnI. Norc prc
csc!, wh!c comosIona!Iy dd ndccd cmcrgc nIhaI simu!aIon
Ihc grammaIca! ru!cs and caIcgorcs dd noI cmcrgc. Ihcy wcrc
parIa!!y bu!I in by Ihc chocc o! symbo! scqucnccs and in!crcncc
a!gorIhm.` WhaI wc nccd arc simu!aIons IhaIcanyc!d ru!cs and
caIcgorcs as an cmcrgcnIproducI by usng rccurrcnI ncura! ncIs Io
cxIracI paIIcms o! co-occurrcncc among words and by sIuaIng
Ihc agcnIs n a soca! spacc n whch Ihc Irans!ormaIon o! habIua!
pracIccsnIo ob!gaIory convcnIonscanIakc p!acc.
1 sImc nowIo rcIumIo a sub| ccIIhaIwas!c!IunnshcdaI Ihc
cndo!IhcprcviouschapIcr.Thcrcwc sawIhaIdcIai!cdarchco!ogca!
simu!aIons o! mcso!Ihc and nco!Ihc communiIcs !a!cd aI Iimcs
Io maIch Ihc avai!ab!c cvdcncc ad IhaI a possb!c cxp!anaIon!or
IhssIhaIoncc omp!cxmcanngscou!dbc!ormcdbyIhccombna-
Ion o! words, and oncc Ihc rc!crcnIs o! Ihosc words ccascd Io bc
drccI!yobscrvb!c ob|ccIs andacIons, magca!bc!c!shaving apow-
cr!u!c!!ccIonbchavorcou!dhavccmcrgcd. mIhcncxIchapIcrwc
wi!!Iack!cIhs knd o!bc!c!s buIwIhouI scparaIing Ihc magnary
wor!dsIhcygvc rscIo!romIhcmaIcra!wor!do!rcsourcc gradcnIs.
MUL1LN1b N KM1YL LNUL
To kccp Ihcsc Iwo wor!dsIogcIhcrmagca! bc!c!s W bc dscusscd
as a mcans Io gcncraIc a ncw knd o! soca! gradcnI, a gradcnI o!
!cgImacy, IhaI govcrnmcnI organzaIos can usc Io manpu!aIc
physca! gradcnIs Io an cxIcnI IhaI car!yhumans wou!dhavc !ound
mysI!ying. m parIcu!ar, Ihc consIrucIon o! hugc !uncrary monu-
mcnIs, !kc Ihc pyramds o! gypI, W bc shownIo havc dcpcndcd
onIhc coup!ingo!magca!and pracIca! bc!c!s, Ihcpyramds Ihcm-
sc!vcs consIIuIng a powcr!u! mcchanism !or Ihc producIon and
mainIcnancc o!!cgIimaIc auIhorIy.
Ib
I
LHPJL LLLVLN
MJ|||dO|5 d0 |C0d|C b|d|O5
Nco!ImccommunIs, !kcIhcrhunIcr-aIhcrcrprcdcccssors, wcrc
rc!aIvc!y ca!Iaran CcrIan pcrsona! qua!Ics such as charsma,
oraIory prowcss, orIhc ab!Iy Io !cad may havc crcaIcd d!!crcnccs
amon Ihcr mcmbcrs buI Ihcsc Icndcd Iobc IranscnI and had!II!c
mpacInIhc !on run. IorIhsrcasonIhcsc communIcshadacccss
Ioon!yonc soca!radcnI, a radcnIo! so!darIy. BuIwhcnd!!cr-
cnccs n prcsIc wcrc noI dsspaIcd quck!y and wcrc a!!owcd Io
accumu!aIc Ihcy bcan Io!orm anoIhcr radcnI, ajrudicnrc]srurus.
marknIhc bcnnn o!Ihc proccss o! soca! sIraIcaIon. Covcrn-
mcnI oranzaIons wIh a wc!!-dcncd auIhorIy sIrucIurc, on Ihc
oIhcrhand,nccdcdmorcIhaIsIaIusd!!crcnccsIocmcrc. AuIhorIy
can a!ways bc corccd n a maIcra! way Ihrouh physca! punsh-
mcnI or conncmcnI buI I s much !css cosI!y andmorc sIab!c ! I
sbc!cvcdIobc!cImaIc.AndImsmp!csIhaIajrudicnrc] lcjirimun
has bccn crcaIcd, a conccnIraIon o!Ihc capacIy Io command|usI-
cd by a rc!ous IradIon c!Ic mcmbcrs Io supcrnaIura!
!orccsor, D somc cascs,|usIcdbyIhc succcss!u!pracIca!rcasonn
o! spcca!zcd burcaucraccs Thc msIorca! proccss IhaI avc rsc Io
Ihcsc Iwo ncw xp!oIab!c rcsourccs uscd Io bc conccpIua!zcd as a
smp!c !ncar c

o!uIon. ca!Iaran arcu!Iura! Irbcs wcrc supcr-

scdcdbychc!doms oranzcdaroundsIaIus d!!crcnccswhchwcrc,
nIum,rcp!accdbyarchacsIaIcsandIhcrccnIra!zcdauIhorIy. BuI
archco!osIshavcmodcdImssmp!cmodc!nscvcra!ways.
IrsI, I has bccomc c!carIhaI Ihcrc s morc Ihan onc IransIona!
cnIIy bcIwccn rura! and urban !orms o! soca! oranzaIon. AI Ihc
MUL1LN1b N KLNL b1Lb
vcry!casI wc musI makc a dsIncIonbcIwccnsmp!c and comp!cx
cmc!doms, Ihc !aIIcr a!rcady dsp!ayDrd soca! sIraIcaIon buI
noI ccnIra!zcd dccson-makn. Sccond, Ihc !ocus o! rcscarch has
sh!Icd !rom !ookn !or sm!arIcs Io cxp!orn Ihc wdc ranc o!
varaIono!cacho!Ihcsc soca!!orms.Andna!!y, IhcdcaIhaIIhcsc
soca!!orms !o!!owcd cach oIhcr n Imc has bccn rcp!accd by Ihc
sIudyo!IhcrcocxsIcnccnspacc. anarchacsIaIcmayhavc!ormcd
Ihc corc o!a!arc rcondomnaIna!cwcomp!cx cmc!doms, wIh
smp!c cmc!doms and arcu!Iura! v!!acs !ormn an cxp!oIcd
pcrphcry.Thc conccpI o!a!ncarcvo!uIonusua!!y !cadsIomodc!s
n wmch a sn!c cnIIy, soccIy as a who!c, dcvc!ops !rom onc
sIac o! dcvc!opmcnI IoanoIhcr. BuI oncc wc rcp!acc IhaI conccp-
Ion wIh onc nvo!vnIhc cocxsIcncc and comp!cx nIcracIon o!
arcu!Iura!communIcs,cmcy!ncacs, andnsIIuIona!oraDza-
Ions, wc nccd scvcra!modc!s.Insomc cascswc canIakc !orranIcd
Ihc cmcrcnI propcrIcs o!Ihcsc !arcrndvdua!cnIIcs-so!darIy,
sIaIus, or !cImacy-and modc! Ihc cnIIcs Ihcmsc!vcs as acnIs.
Indvdua! oraDzaIons, !or cxamp!c, can bc modc!cd as acnIs
bccausc Ihcy havc Ihcr own oa!s |o! wmch mosI o! Ihcr sIa!!
maybc unawarc) : arc capab!co!a!!ccInandbcn a!!ccIcdbyIhcr
cnvronmcnI |as whcn an cnvronmcnIa! crss a!!ccIs an cnIrc
oraDzaIon) : posscssnsIIuIona!know!cdc and mcmory |wrIIcn
rccords, rouIncs): havc Ihcrown!ca! sIandn, rhIs, and ob!a-
Ions: and arc capab!c o!coordnaInIhc acIvIy o! many workcrs
andmanacrsu Ihcpcr!ormancc o!comp!cxIasks.
Nu!IacnI smu!aIons n wmch Ihcbchavoro!acnIss cnIrc!y
dcIcrmncdby!orma!ru!csarc adcquaIc Iodca!wIhIms casc, aswc
saw n ChapIcr whcrc ndvdua! communIcs wcrc modc!cd as
sn!c acnIs. BuI !wc wanIIocxp!an howpropcrIcs!kchc sIaIus
o! a !ncac or Ihc !cImacy o! an oranzaIon cmcrc wc musI
smu!aIc Ihc nIcracIons bcIwccn pcrsons cxp!cI!y and modc! Ihcr
bchavorasdcIcrmncdbybc!c!sand dcsrcs. Ihc bc!c!IhaIasacrcd
IcxI|usIcs Ihc auIhorIyo! a suprcmc !cadcr, !or cxamp!c, or Ihc
dcsrc Io bc parI o! Ihc msson o! oraDzaIon. Thc acnIs wc
havc dscusscd so !ar do noI mccI Ims rcqurcmcn Ihc ru!cs IhaI
udc IhcrbchavormaymmcIhc c!!ccI o!havn bc!c!s ordcsrcs
buI Ihc !aIIcr arc noI cxp!cI!y modc!cd and cannoI nIcracI Io
producc cmcrcnI c!!ccIs. Thc smp!csI way o! modc!n bc!c!s and
I
Id
NLbN N b MUL1N
dcsircs arc as aIIiIudcs orsIanccsIakcn bv pcrsonsIowardproosi-
Iions. IhaI is, as proposiIionaI aIIiIudcs. Using proposiIions as Ihc
ob|ccIs o! mcnIaI aIIiIudcs disrcgards Ihc !acI IhaI prc-ImguisIic
humansaIrcadvposscsscdbcIic!sanddcsircsrcIaIivcIoIhcconIcnIo!
Ihcircpisodicmcmorics. Ihc bcIic!IhaI somcIhinghappcncdbccausc
onc wiIncsscdiI happcnng, !or insIancc, or Ihc dcsirc !or a Bavor,
aroma, sound, orsighI IhaI onc has prcviousIv cxpcricnccd. BuI Ihc
bcIic! on supcrnaIuraI !orccs or Ihc dcsirc noI Io o!!cnd Ihosc !orccs
cannoI cxisIwiIhouIIanguagc. ToIackIcmorccompIcx cascs o!pcr-
sonaIagcncvrcccnIapproachcsIoIhcdcsigno!simuIaIcdagcnIshavc
rcpIaccdrigidruIcswiIhmorc!IcxibIcproposiIionaIaIIiIudcs.Thisis
Ihc sIraIcgv !oIIowcd in Ihc so-caIIcd BcIic!-csirc-InIcnIion |BI)
approach.
BI agcnIs arc noI, in !acI, dcsigncd Io dcaI wiIh rcIigious or
idcoIogicaIbcIc!sbuIIorcasonabouIpracIicaImaIIcrs.AsinIhccasc
o! oIhcr agcnI dcsigns Ihis invoIvcs mapping scnsorv inpuIs mIo
appropriaIc bchavioraI ouIpuIs buI insIcado!usingruIcsIo mcdiaIc
bcIwccnpcrccpIionandacIionamorccIaboraIcproccssiscmpIovcd.
m!ormaIionabouIIhcworId,abouIaccrIaindisIribuIiono!rcsourccs,
!or cxampIc, isrsI uscd IoupdaIc an agcnIsbcIic!s: Ihcsc arc Ihcn
uscdIogcncraIca scIo!opIions conccrnngIhcavaiabIc opporIuni-
Iics and risks: Ihc opIions arc Ihcn chcckcd !or how dcsirabIc and
achcvabIc Ihcv arc: opIions IhaI arc dcsircd and can bc achicvcd
bccomcgoaIsandIhcsc, combmcdwiIhacommiImcnIIoa coursco!
acIion, bccomc Ihc inIcnIions driving Ihc bchavior o! Ihc agcnI.
Usmg BI agcnIs !or Ihc siuIaIion o! supcmaIuraI bcIic!s wouId
mvoIvcIwosimpIcmodicaIions.IirsIwcwouIdhavcIogivcagcnIs
IhccapaciIvIoascribcproposiIionaIaIIiIudcsIooIhcragcnIsinordcr
Io makc scnsc o! Ihcir acIions, a capaciIv rc!crrcd Io as Iaking Ihc
mIcnIionaI sIancc Ioward Ihc cxpIanaIion o! bchavior. Sccond,
agcnIs shouId bc abIc rc rrcurrcscurccsusujcnrs.'TogcIhcr Ihcsc Iwo
modicaIionswouIdgivcBI agcnIsIhccapaciIvIoadopIIhcinIcn-
IionaIsIanccIowardmaIcriaIgradicnIs,aIIribuIingbcIic!sand dcsircs
Io cnIiIicsIkcIc sun, Ihc rain, Ihc soiI, and so on. ToIrans!orm a
!cw magicaI bcIic!s inIo an idcoIogv onc morc mgrcdicnI wouId bc
ncccssarv. dcbaIcsamongagcnIsaimcdaIpcrsuasion. InIhcaccounI
o! Ihc cmcrgcncc o! Ianguagc givcn in Ihc prcvious chapIcr iI was
imporIanIIo assumc IhaI agcnIs did noIhavc acccss Io Ihc conIcnIs
MUL1LN1b N KLNL b11Lb
o!cachoIhcrsmindsandIhaIIhcvcouIdonIv!orminIcnIionsabouI
changing cach oIhcrs bchavor. BuI oncc Ianguagc cmcrgcd and
couIdcxrcss IhcconIcnIo!proposiIionaI aIIiIudcs humansbccamc
abIcIo!ormmIcnIionsinwhichIhcgoaIwasIomodi!vIhcvcrvcon-
IcnIo! Ihosc bcIic!s anddcsircs. U Ihc sub| ccImIIcro! suchpcrsua-
sionaIIcmpIsmvoIvcdcxpIanaIionso!rcsourccscarciIvininIcnIionaI
Icrms, Iikc Thc soi has bccomcbarrcnbccausc Ihc!crIiIiIv goddcss
is angrv, Ihcn!romIhccnsumgdiscussions acohcrcnI scI o!supcr-
naIuraIbcIic!scouIdcmcrgc.IroIocoIs!orncgoIiaIionIhroughwhich
BI agcnIs can csIabIish Ihcir posiIions on an issuc, makc conccs-
sions, andrcachagrccmcnIs arc currcnIIvbcingdcvcIopcd.
ThcBI approach Io agcnI dcsignisiniIsin!ancv andIhckind o!
appIicaIion Io archcoIogicaI simuIaIions |usI suggcsIcddocs noI vcI
cxisI. BuI simpIcr approachcs using ruIcs Io gcncraIc bchavior IhaI
mimics Ihc c!!ccI o! proposiIionaI aIIiIudcs arc aIrcadv vicIding
inIcrcsIingrcsuIIs. Onco! Ihcm,caIIcdmcrgcncco!SocicIv | OS) ,
cxpIorcs Ihc rsI sIcp i nIhc cvoIuIion o! organizaIionaI auIhoriIv.
Ihc risc among cgaIiIarian communiIics o! cndurinj lcudcrshir. In a
IvpicaI OS simuIaIion agcnIs proposc pIans abouI spccic wavs o!
harvcsIingrcsourccs.whaIrcsourccIoIargcI,howmanvoIhcragcnIs
arc nccdcd, how Io aIIocaIc Ihcir Iabor. I! oIhcr agcnIs bcIicvc IhaI
IhispIan is bcIIcr Ihan Ihcir own Ihcv adopI iI. Whcn an agcnI has
parIicipaIcdinanoIhcr agcnIs pIanonscvcraI occasionsiI bccomcs a
!oIIowcrwhiIc IhoscwhoscpIansarc!oIIowcdbccomcIcadcrs.Whcn
agrouphas!ormcdaroundaIcadcrccrIainIcndcncicsbccomcmani-
!csI IhaI sIrcngIhcn Ihc bonds hoIding Ihc group IogcIhcr. iIs mcm-
bcrs Icnd Io communicaIc wiIh cach oIhcr morc o!Icn abouI Ihc
IocaIion and conccnIraIion o! rcsourccs and Icnd Io Iakc cach oIhcr
inIoaccounIinIhc!ormaIiono!!uIurcpIans. OnccIhis dvnamic o!
group !ormaIion has bccn csIabIishcd Ihc agcnIs can bc con!ronIcd
wiIh cnvironmcnIaI criscs Io chcck whaI c!!ccIs Ihcsc havc on Ihc
sIrucIurco!Ihcgroups.InoIhcrwords,whaIIhissimuIaIioncxpIorcs
is Ihc rcIaIion bcIwccn cndurmg Icadcrship and a spacc o! possibIc
rcsourcc disIribuIions. This spacc mav bc picIurcd as consisIing o!
Iwo dmcnsions, onc rcIaImg Io rcIaIivc abundanccs and scarciIics,
Ic oIhcr Io rcIaIivc conccnIraIions and dispcrsions. Abundancc
wcakcnsIcadcrship smccmosIagcnIscanmccIIhcirmcIaboIicnccds
bv working aIoncwhiIc scarciIv acIs an inccnIivc IoworkIogcIhcr.
IV
I
N LbN N b MUL1N
spcrson o! rcsourccs, n Iurn, makc crcaIng succcss!u! p!ans
hardcr snccIncrcascsIhc d!cu!Iy o! coordnaIng Ihc acIons o!a
group wh!c conccnIraIon!ac!IaIcs p!annng. Thc ouIcomc o! OS
smu!aIons hasncrcascdIhcp!ausb!Iyo!ancxsIng archco!ogca!
hypoIhcss. IhaI Ihc hsIorca! cmcrgcncc o! cndurng !cadcrshp
wasIrggcrcdby cnvronmcnIa! changcs !cadng Io boIh scarcIy and
conccnIraIon o!rcsourccs. '
Ths smu!aIon capIurcsIhcmaIcra! aspccI o!Ihc IransIon !rom
a sIrcI cga!Iaran communIy Io onc n whch an accumu!aIon o!
prcsIgc around a !cadcr s noI rapd!y dsspaIcd. Thc manIcnancc
o! such a gradcnI ovcr !ongcr Imc sca!cs, howcvcr, s prob!cmaIc
bccausc nhunIcr-gaIhcrcr andagrcu!Iura!communIcs dcparIurcs
!rom an cga!Iaran organzaIon arc acIvc!y prcvcnIcd by nIcrna!
dcvccs. Icadcrs arc dcmandcd, !or cxamp!c, Io bc gcncrous and
rcdsIrbuIcnccrcmona! !casIs whaIcvcr !ood surp!uscs havc accu-
mu!aIcd n Ihcr hands, or c!sc Io rIua!!yburn Ihcm. CusIoms !kc
IhsmayacI asanunconscous mcchansmIoguardagansIIhc crys-
Ia!!zaIon o! !orma! auIhorIy. Norcovcr, war hcrocs, prophcIs, or
promoIcrso!agrcu!Iura!producIvIymayndIhcr!cadcrshpcom-
promscd ! anyms!orIunc |a dc!caI nwar, Ihc !a!urc o! a p!an, a
naIura! dsasIcr) s bc!cvcd by Ihcr !o!!owcrs Io bc a supcrnaIura!
sgn IhaI Ihc !cadcrs do noI n !acI posscss Ihc admrcd charsmaIc
vrIuc. Nagca!bc!c!s, onIhcoIhcrhand, cannspcca!crcumsIanccs
sIab!zcraIhcrIhandsspaIc asIaIusgradcnI.ThccaIa!ysI!orsucha
changc can bc a succcsson crss cnsung a!Icr Ihc dcaIh o! a chcr-
shcd !cadcr. Thcsc crscs can !cadIoIhc Irans!ormaIon o! charsma
!rom a pcrsona! characIcrsIcnIo oncIhaI canbc IransmIIca!ong
a b!ood !inc, a Irans!ormaIon Iypca!!y |usIcd by Ihc nvocaIon
o! supcmaIura!bc!c!sAndoncc!cadcrshp canbcnhcrIcdI can
p!ay a cruca! ro!c n Ihc gcncraIon o! cndurng conccnIraIons
o! prcsIgc around parIcu!ar knshp groups. Smu!ang Ihs crIca!
IransIon wou!d nvo!vc augmcnIng smu!aIons !kc Ihc onc |usI
dcscrbcd wIh BI agcnIs capab!c o! havng magca! bc!c!s and o!
cngagng ndcbaIcsabouI IhcsourccsandpoIcnIa!Iransmssb!Iy
o!charsmaIcpowcrs.
Oncc sIaIusbccomcs apropcrIyIhaIcanaccumu!aIc I opcns Ihc
way!orIhc d!!crcnIaIon o!communcs, or o!groupswIhn com-
munIcs, nIo soca! sIraIa. Archco!ogca! cvdncc !or sIraIcaIon
MUL1LN1b N KLNL b11Lb
comcs !rom avarcIy o! sourccs. Sncc comp!cx chc!doms organzc
cnIrc gcographca! rcgons Ihc rcmans o! scII!cmcnIs n a rcgon
maybc rankcdby szc, Ihc numbcr o!ranks counIcd, and Ihc dsIr-
buIon o! scII!cmcnI szcs pcr rank sIudcd. Ths rcgona! hcrarchy
canIhcnbc uscdasanndcaIono!Ihc numbcro!hcrarchca!!cvc!s
o! dccson-makng IhaI cxsIcd n Ihosc communIcs. AI a sma!!cr
sca!cbura! sIcs o!!cr anoIhcrwndownIo soca! sIraIa. SysIcmaIc
bura!wasa!rcadypracIccdbyhumansBIhcmdd!cpa!co!Ihcand
by Ihc uppcr pa!co!Ihc I had bccomc ccrcmona!. Ihc dcad wcrc
burcdc!oIhcdandwcarngpcrsona!adornmcnIs. ' Thcbura!sIcso!
chc!s conIan n addIon Io pcrsona! n!ormaIon va!uab!c ndca-
Ions abouI Ihc srccrncss o! sIaIus gradcnIs. Ihc szc o! a Iomb, Is
dcmarcaIon !rom oIhcr gravcs, and morc mporIanI!y, Ihc rchncss
o! Is conIcnIs. O!parIcu!arsgncancc arc Ihc prcscncc o! adorn-
mcnIsmadc !rom cxoIcmaIcra!srangng !romobsdan, !mcsIonc,
andbasa!I, Iocoppcr,!aps!azu!, Iurquosc, anda!abasIcr. Thcscrarc
ob|ccIs arc sgncanI bccausc Ihcy consIIuIcd a d!!crcnI !orm o!
wca!Ih Ihan agrcu!Iura! producIs. Ihcy cou!d bc uscd noI on!y Io
cxprcss sIaIus d!!crcnccs buI a!so as a ndo! po!Ica!currcncy Io
con!cr sIaIus Io oIhcrs, as whcn Ihc !cadcr o! a comp!cx chc!dom
dsIrbuIcdIhcmIoIhosc o!smp!c chc!domsIoccmcnIa!!anccsand
dcnc !oca! rankngs.' Ths movcmcnI o! ob|ccIs o! sIaIus !rom a
p!acc o!hghconccnIraIonIoonco!!owconccnIraIonconrmsIhaI
prcsIgc can!ormagradcnI. ccpngI!romdsspaIngnvo!vcdIhc
sIrcIconIro! o! Ihc !ong-dsIancc Iradc o! rarc raw maIcra!s and o!
Ihc spcca!zcd cra!Is nccdcd Io producc c!Ic adornmcnIs. Indccd,
buryng Ihcsc rarc ob|ccIs IogcIhcr wIh Ihcr !ormcr owncrs may
havcbccn awayIoprcvcnI n!!aIonaryprcssurcs!romdcvc!opng
bypcrmancnI!yIakngIhcmouI o!crcu!aIon.
Thc cxsIcncc o! an cndurng gradcnI o!sIaIuscan!cadIoIhccrc-
aIon o!rank d!!crcnccsIhaIarc !!ud and conInuous as n smp!c
chc!doms or xcd and sharp!y dsconInuous as n comp!cx chc!-
doms. To cxp!orc how Ihcsc d!!crcnI ouIcomcs may bc produccd
wc do noI nccdsmu!aIonsn whchpcrsona!agcncys modc!cdn
dcIa! buI raIhcr oncs IhaI capIurc Ihc nIcracIon bcIwccn !argcr
sca!c soca!cnIIcsIakngp!acc aI!ongcrIcmpora! sca!cs. Onc cxam-
p!csasmu!aIon| ca!!cdTongaSm)nsprcdbyaconIcmporarycasc.
Ihc Tonga chc!dom n Io!yncsa. Ths chc!dom s organzcd nIo
II
IZ
N LbN N b MUL1N
knshp groups ca!!cd !ncagcs, cach wIh Is own chc! or chc!s,
IhaIarcrankcdnIcrmso!sIaIus. Thcscrankngscanchangcdcpcnd
ng onIhc rc!aIons bcIwccn!ncagcs. Spccca!!y, marragcs among
Ihc o!!sprng o! II!cd chc!s can ncrcasc or dccrcasc Ihc rank o!
a !ncagc. ! Ihc II!cdhcrmarrcs somconc !roma hghcr sIaIusn-
cagc hs or hcr own !ncagc w!! gan n sIaIus and vcc vcrsa, hcrs
marryng down n sIaIus brng down Ihcr own !ncag. SgncanI
changcsnIhcrankngo!!ncagcs canIakc up Io!ourIovc gcncra
Ions so on!y a scrcs o! bad marragc dccsons canhavc Ihs c!!ccI.
ach!ncagc s nvo!vcdn cconomc acIvIy, Is agrcu!Iura! wca!Ih
dcIcrmncdby numbcr o!kn, Ihc !andava!ab!cIo I, and Ihc pcr
sona!ab!Iyo!Is chc!IomoIvaIc boIhknandcommoncrsIowork
hardcr. Thsagrcu!Iura!wca!Ih, Ium, !!owswIhn!ncagcs n Ihc
!orm o! rcdsIrbuIon Io comoncrs, and bcIwccn !ncagcs !rom
w!c-rcccvng Io w!c-gvng oncs. Agrcu!Iura! wca!Ih a!so !!ows n
Ihc !orm o!IrbuIc pad Io !ncagcs o!hghcr rank.
Thus n chc!doms maIcra! and sIaIus gradcnIs arc nIcrIwncd,
Ihcr nIcracIon pvoIng on marragc dccsons as wc!! as on dcc
sonsa!!ccIngIhcsc!ccIono!II!cdhcrs.Thc qucsIonIhaIIhc sm
u!aIonscIouIIoanswcrshowIhc rgdsoca!sIraIcaIonobscrvcd
ncomp!cxchc!domscou!dcmcrgc!romIhcdynamcso!IhcscnIcr
acInggradcnIs. Oncpossb!c answcrsIhcrc!axaIono!prohbIons
Io marry c!osc rc!aIvcs. InccsI Iaboos comc n d!!crcnI sIrcngIhs.
IhcymayprohbIIhcmarryngo!broIhcrsorssIcrs:Ihcmarryngo!
rsI cousns: or Ihc marryng o! any rc!aIvc howcvcr dsIanI. Thc
sIrongcrIhc prohbIons Ihc morc !kc!yIs IhaI an hcrbc!ongng
Ioa hghsIaIus!ncagcw!!bc !orccd Iomarrysomconc wIh!owcr
sIaIus. II s Ihcrc!orc n Ihc nIcrcsI o!Ihosc wIhIhc hghcsI sIaIus
Io!owcrIhc sIrcngIh o!Ihc Iaboo. HsIorca!!yIhs cou!d havc bccn
achcvcdIhrough po!Ica! sIraIcgcsbychc!s buIwIh smp!c smu
!aIcd agcnIs Ihs !cvc! o!sca!c cannoIbc capIurcd. Thc rc!axaIon o!
Iaboos, onIhc oIhcr hand, canbc pcr!ormcd cxogcnous!y Io chcck
whcIhcrIhasIhcosIu!aIcdc!!ccIondcgrcc o!sIraIcaIon.mIhc
smu!aIon Icnchc!!y !ncagcs sIarIcd Ihc proccss. As popu!aIon
ncrcascd and omc !ncagcs ran ouI o! agrcu!Iura! !and Ihcy sp!I
nIoIwoncrcasngIcovcra!!numbcr. |Thc smu!aIoncou!hand!c
upIo 50 !ncagcs. ) Thc smu!aIon wcnIIhrougha scrcs o!cyc!cs as
cach!ncagcrankcdpoIcnIa!spouscsandsc!ccIcdhcrs,hundrcdso!
MUL1LN1b N KLNL b1Lb
smu!aIcd maragcs Iook p!acc, agrcu!Iura! wca!Ih !!owc among
!ncagcs, and sIaIus was IransmIIcd. Oncc Ihs basc dynamc was
csIab!shcd cxpcrmcnIs on Ihc c!!ccI o! Iaboos o!d!!crcnIsIrcngIhs
cou!dbc conducIcd. Ncasurcs o!sIraIcaIonnc!udcdIhc sIccpncss
o! Ihc sIaIus gradcnI |how !argc Ihc d!!crcnccs n sIaIus bcIwccn
!ncagcswcrc) andIhcmob!Iyo!Ihc!ncagcs |howo!Icnchangcsn
Ihc ranngoccurrcd) . Thc ouIcomc o!Ihc smu!aIon conrmcdIhc
basc!ogcbchndmarragcsIraIcgcs.Ihc!owcrIhc sIrcngIho!nccsI
prohbIons Ihc hghcr Ihc dcgrcc o!sIraIcaIon '
I! Ihc bura! sIcs o! II!cd chc!s can bc sad Io bc our wndow
nIo soca! sIraIcaIon IhcnarchIccIura!monumcnIs !kcpyramds,
Icmp!cs, and pa!accs arc our man sourcc o! cvdcncc !or Ihc cxs-
Icncc o! ccnIra!zcd dccson-makng n archac sIaIcs. Thc ccnIra!
dccsons !cadng Io Ihc crccIon o! monumcnIa! sIrucIurcs wcrc
bascdonboIh Icchnca! and magca!bc!c!s andn!ormcdboIhaIc-
ra! andccrcmona!pracIccs. IcIs Iakc Ihc cxamp!c o! Ihc !amous
pyramd o! Cza n gypI. On onc hand, Ihs !uncrary monumcnI
wasc!car!ynIcndcdIohavcmagca!!!ccIs sncc Iwas supposcdIo
!uncIon!kc a rcsurrccIon machnc. Nccd!css Io say, Ihc pyramds
nIcrna!mcchansm ddnoIa!!owIIoacIua!!yIransmuIcakngnIo
a godbuIIncvcrIhc!css!uncIoncd!kcamachnc!orIhcproducIon
o! !cgImacy, boIh by Is powcr!u! c!!ccI on Ihc rc!gous bc!c!s o!
Ihosc who conIcmp!aIcd I and by scrvng as a rIua! ccnIcr!or Ihc
conInuous pcr!ormancc o! IradIona! ccrcmoncs assocaIcd wIh
Ihc cu!I o! a dcad kng. On Ihc oIhcr hand, Ihc dcsgn o!Ihc monu
mcnInvo!vcdhgh!yIcchnca! know!cdgc such as Ihc asIronomca!
know!cdgc nccdc!ors propcr orcnIaIon Ioward Ihc hcavcnsI
s vcry prccsc!y orcnId Iruc norIh-and Ihc app!cd maIhcmaIca!
know!cdgcnvo!vcdnca!cu!aIngIs shapc. IIs consIrucIon, nIum,
dcmandcd wc!!-organzcd maIcra! pracIccs. Thc pyramd s madc
o! morc Ihan Iwo m!!on b!ocks o! sIonc cach wcghng ovcr Iwo
Ions on avcragc. Thcsc b!ocks wcrc orgna!!y covcrcd by po!shcd
whIc !mcsIonc IhaI, un!kc Ihc corc sIonc, was an cxoIc maIcra!
IransporIcd !rom !ar away. Bu!dng such a sIrucIurc n |usI ovcr
Iwo dccadcs s cvdcncc o!Ihc rc!aIvc c!ccncy o!Ihc burcaucraIc
organzaIons nvo!vcd, Ihc !cgImacy o! whch dcrvcd !rom Ihcr
pragmaIcab!IyIo ovcrcomc Ihc cognIvc, physca!, Icmpora!, and
nsIIuIona! !mIaIons o! pcrsona! agcncy. In shorI, Ihc pyramds
IJ
I+
NLbN N b MUL1N
wcrcboIhccrcmona!andmaIcra!cnIIcs, IhcrsI aspccIrc!aIcdIo
Ihcr capacIy Io cxprcss n Ihcr so!d pcrmancncc Ihc va!dIy and
conInuIy o! auIhorIy andIhc sccond Io Ihc !acIIhaI Ihcy consI-
IuIcd a !rozcn ndcaIon o! Ihc IoIa! amounI o! cncrgy IhaI a gvcn
c!Ic wasab!c Iohamcss.'
Bc!orcIhcgrcaIpyramd o!CzaIhcrcwason!y a ccnIury o!con-
sIrucIon cxpcrcncc sIarIngwIhIhc vcryrsIgypIan monumcnI
bu!I cnIrc!y ouI o! sIonc, Ihc SIcp pyramd aI Saqqara. mbcIwccn
Ihcsc Iwo Ihcrc wcrc a !cw cxpcrmcnIs nIhc bu!dng o! smooIh-
sdcdpyramds onc o!wLchsccms Io showIhaIIhc archIccIs wcrc
sI!!!cangsnccIhcychangcdIhcs!opco!IhcmonumcnIasIgrcw
Ia!!cr |Ihc BcnIpyramdaI ashur) . OnIhc oIhcrhand,prorIoILs
rc!aIvc!y shorIbursI o! crcaIvc acIvIy Ihcrc wasncar!y a m!!cn-
num o!cxpcrcncc nIhc consIrucIon o!Iombs and oIhcr!uncrary
monumcnIs gong backIo Ihc !aIc prc-dynasIc cra whcn cLc!Ians
Iombs bu!I ouI o! brck wIh somc sIonc c!cmcnIs sccmcd a!rcady
nIndcdIo!asI!or an cIcmIy. 'Thcsc Iwo d!!crcnIImc sca!cs ca!!
!or d!!crcnI cxp!anaIons. Nodc!ng a !cang proccss !asIng many
ccnIurcs shou!d usc as agcnIs Ihc organzaIons Ihcmsc!vcs. roya!
workshops and Ihc parIs o!Ihc rsng ccnIra! govccnI on wLch
Ihc workshops dcpcndcd. Nodc!ngaproccssnvo!vngIwo orIhrcc
gcncraIons, on Ihc oIhcr hand, a pcrod o! Imc n wLch pro|ccIs
grcwso!argcso!asIIhaIburcaucraIshadIo!camIoso!vcnovc!!ogs-
Ica!prob!cms durng Ihcr!!cImcs, shou!d usc smu!aIcdpcrsona!
agcnIs. Onc wayIoapproachIhc smu!aIon o!organzaIona! !cam-
ngIakngp!acc aI m!!cnna Imc sca!css Io modc!I as an cvo!u-
Ionary proccss. OrganzaIons canbc modc!cd as cnIIcs capab!c o!
cvo!uIonH Ihcrda!yrouIncs andproccdurcs, andIhcrdvsono!
!aborandcoordnaIonmcchansms, arc passcd!romoncgcncraIon
Io Ihc ncxI, or cvcn !rom onc organzaIon Io anoIhcr wILn Ihc
samc gcncraIon. '"
Wc canmagncIhcacIvIyo!Iomb bu!dng car!y on in gypIan
LsIory as dvdcd nIo scparaIc Iasks cach pcr!ormcd by a scparaIc
arIsan nwhcorganzaIona!!carnngnvo!vcd Ihc c!mnaIon o!
nc!ccnccsIhrough!urIhcrsubdvsonnIosubIasks.Norc spcc-
ca!!y, H Ihcrc wcrcsubIasksIhaIhadIobc cxccuIcd asparI o! scvcra!
Iasks Ihcn a ccrIan dcgrcc o! ovcr!ap cxsIcd among Ihc acIvIcs.
l! asng!c arIsan wcrc assgncd noIoncbuI Iwo ovcr!appng Iasks
MUL1LN1b N KLNL b11Lb
IhcnIhc sharcdsubIaskswou!dhavc hadIobcpcr!ormcdonIyoncc
rcducng Ihc ovcra!! amounI o! c!!orI nvo!vcd. On Ihc oIhcr hand,
Ihc c!!orI o! coordnaIng Ihc d!!crcnI acIvIcs by ovcrsccrs wou!d
havc ncrcascd. Thus, Ihcrc was a Iradc-obcIwccnIhc advanIagcs
o! c!usIcrng ovcr!appng Iasks and Ihc coordnaIon cosIs ncurrcd
bysuchc!usIcrng. Tosmu!aIc organzaIona!cvo!uIonusnggcncIc
a!gorIhmsIhc rouInc IaskassgnmcnIs musI bc codcd nIo a chro-
mosomc nwLhIhc oncsand zcrocssIand!oragcnIs andsubIasks,
whIcIhcIncss!uncIonmcasurcsIhcovcra!!amounIo!c!!orIIakcn
Io comp!cIc Ihc Iask. Smu!aIons !kc Ihcsc havc bccn carrcd ouI
andhavc shownIhaIc!usIcrngo!Iasks docs occurbuInoIn a way
IhaIIakcs!u!!advanIagc o!!uncI

ona!comp!cmcnIarIcsamongsub-
Iasks." Onc way o! mprovng organzaIona! !carnng wou!d bc Io
modc! InoI asab!ndcvo!uIonary scarchbuIraIhcras a hybrd o!
gcncIc a!gorIhms and rcn!orccmcnI !carnng. Thcsc hybrds arc
ca!!cd clussmcrsysrcms.' m a c!asscr sysIcm whaI cvo!vcs arc ru!cs
IhaIgudcIhc pcr!ormancc o!Iasks, ru!csIhaI spcc!yIhaI !a cond-
Ion arscs Ihcn a ccrIan acIon musI bc Iakcn. IIncss s cva!uaIcd
noIbyIhcamounI o!c!!orIspcnIpcr!ormngaIaskbuIbyIhc ab!Iy
Io gcI a rcward !rom a pcr!ormanc. Norcovcr, Incss s assgncd
noIIo ndvdua! ru!cs buIIo groups o! rc!aIcd ru!cs, a sharcd crcdI
assgnmcnI IhaI Icnds Io producc coadapIcd scIs o! ru!cs raIhcrIhan
a sng!c IIcsI ru!c. m Ihc prcscnI casc Ihc condIon-acIon ru!cs
wou!d rcprcscnI cIhcr cusIomary acIvIcs n wLch bchavora!
rcsponscs Io spccc condIons havc bccomc ru!c-!kc or wrIIcn
rcgu!aIons spcc!yng Ihc connccIon bcIwccn a condIon and an
acIon. Koya! workshops wou!d bc Ihc cvo!vng cnIIcs wh!c Ihc
sourcc o! rcwards wou!d bc oIhcr organzaIons on wLch Ihc work-
shops dcpcndcd !or!cgImacy andnancng.
m addIon IoIhc s!ow organzaIona! !arnngbchndIhc pracIcc
o! Iomb bu!dng Ihcrc s Ihc much !asIcr onc !cadng Io pyramd
bu!dng. TogcI an dca o!Ihc cha!!cngc ILsIaskposcdIo arcLIccIs,
o!ca!s, and burcaucraIs wcnccdIo modc! Ihc consIrucIon proccss
Isc!!, a proccss nvo!vng a o! Ihc Icchnca!andIhc ccrcmona!.
On onc hand, Ihc work!orcc was moIvaIcdbyIhc nIcnsc rc!gous
!cc!ngs gcncraIcdbyIsnvo!vcmcnInadvncpro|ccI.A!Ihoughn
Ihc pasI Ihs work!orcc was assumcd Io bc composcd o! unsk!!cd
s!avcs b!nd!y !o!!owng commands !rom abovc I s now c!car IhaI
Ib
I
NlLbN N bMUL1lN
mosI o! Ihc opcraIons nvo!vcd n pyramd bu!dng dcmandcd
hghsk!!andmoIvaIon KcccnI!y cxcavaIcdccmcIcrcsnIhcIown
IhaI uscd Iohousc o!hca!s, ovcrsccrs, arIsIs, andarIsans showIhaI
cvcnIhcworkcrswcrc burcdrcspccI!u!!y, sIrongcvdcnccIhaIIhcy
wcrc noI s!avcs. OnIhc oIhcrhand, boIhIhc rc!gous!y moIvaIcd
work!orcc and Ihc rc!gous!y !cgImaIcd organzaIona! sIrucIurc
abovc Ihcm had Io Iakc nIo accounI Ihc capacIy o! Ihc maIcra!
wor!d Io bc a!!ccIcd by human acIvIy as wc!! as Ihc capacIy o!
humans Io a!!ccI Ihc wor!d aI IhaI sca!c. Norc spcchca!!y, Io undcr-
sIandIhcmaIcra!cha!!cngcposcdbyIhcacIvIyo!pyramdbu!dng
wc nccdIo dchnc boIh Ihc ob|ccIvc naIurc o!Ihc Iask cnvronmcnI
as wc!! as Ihc knds o! coordnaIng mcchansms nccdcdIo organzc
a !argc work!orcc Io mccI IhaI cha!!cngc. In whaI !o!!ows wc w!!
conccnIraIc onIhc Icchnca! componcnI o!Ihc consIrucIon proccss
buIIshou!dbckcpInmndIhaIwIhouIIhc ccrcmona!componcnI
Ihc cxp!anaIonwou!drcmanncomp!cIc.
Thc wayIhc gypIans dca!I wIh Ihc consIrucIon prob!cm was
Ihrough a rccursvc dccomposIon approach, groupng IogcIhcr
rc!aIcd Iasks and asscmb!ng Ihcm as parIs o! a !argcr who!c. Thc
ovcra!! Iask o! pyramd bu!dng was dvdcd nIo aI !casI Ihrcc man
Iask groups. quarryngIh sIonc: IransporIngI Io Ihc consIrucIon
sIc: and crccIng I. uarryng, n Iurn, was subdvdcd nIo scvcra!
Iasks. carvng dccp channc!s nIo a !mcsIonc bcd ncarby: carvng
narrowchannc!sIosubdvdc InIob!ocks:andusngwoodcn!cvcrs
Io dcIachIhcm!romIhc bcd. ansporIngIhcb!ockswas subdvdcd
nIo Ihc bu!dngo!Iracks and ramps onIop o!whchIhcy cou!d bc
movcd:!oadngIhcmnIowoodcns!cdgcs:andpu!!ng orro!!ng Ihc
s!cdgcs ovcr cy!ndrca! bcams. Ina!!y, crccIng Ihc sIoncs nvo!vcd
scIIng Ihcm usng woodcn !cvcrs: carvng marks on Ihc sIoncs Io
conIro!Ihcs!opc:andIrmmngIhc sIoncsIoIhcrhna!shapc.Thc
c!!orIs o! cxpcrmcnIa! archco!ogsIs n Ihc pasI !cw dccadcs havc
gvcn us a scnsc o!Ihc szc o!Ihcwork!orcc andIhc amounI o!Imc
IakcnIo cxccuIc cach Iask. m onc cxpcrmcnI a Icam o! I 2quarry-
manwasab!coproducc cghIsIoncb!ocksaday: 20mcncou!dhau!
Ihc b!ocks !rom Ihc quarryIoIhcbu!dngsIc aI araIc o! Icnb!ocks
aday:andaIcamo!!ourmcn,IwoIopushandad|usIandIwomasons
Iopcr!ormIhc Ig, was nccdcd Io crccI cachb!ock. ToIhswc
musIaddavarcIyo!supporIworkcrs,!rom carpcnIcrsandb!acksmIhs
MULlLN1b N KLNL b11Lb
IocrcaIcwoodcns!cdgcsandmcIa!Ioo!s,IopoIIcrs,bakcrs, andbrcw-
crs Io producc conIancrs, !ood, and drnk. Ovcra!! Ihs cxpcrmcnI
showcdIhaI a!orcc o!25, 000workcrscou!dhavchnshcdIhc Iaskn
Ihc 23 ycarsIhaIIhcbu!dno!Ihcpyramdorgna!!yIook.
In addIon IoIhcscphysca! cxpcrmcnIs Io dup!caIc somc o!Ihc
orgna!Iasksmu!IagcnI smu!aIonscanbc uscdIo cxp!orcIhcparI-
Io-who!crc!aIonsbcIwccnIasksandsubIasks.Wh!csmp!cpro|ccIs
canbcsubdvdcdnsuch awayIhaIIhcyarcnomorc Ihanasumo!
IhcrparIs, ! Ihcrc arc nIcrdcpcndcnccsbcIwccncomponcnI Iasks
IhcnIhc componcnIs cannoI smp!y bc addcd buI musIbc carc!u!!y
arIcu!aIcd. Thc ob|ccIvc spacc o!possb!Ics o! d!!crcnI Iask cnv-
ronmcnIs can bc cxp!orcd usng a !orma! !anguagc ca!!cd Task
Ana!yss, nvronmcnI Nodc!ng, and Smu!aIon |orTacms) . Tacms
can bc uscd Io modc! Ihc rc!aIon bcIwccn Ihc man Iask and Is
subIasks and Io capIurc Ihc nIcrdcpcndcnccs bcIwccn ndvdua!
Iasks. pcr!ormancc o! onc Iask, !or cxamp!c, may cnab!c Ihc pcr!or-
mancc o!anoIhcronc soImusIIakc Icmpora!prcccdcncc: orI may
mcrc!y !ac!IaIc Is pcr!ormancc so Ihc hrsI Iask nccd noI bc !u!!y
comp!cIcd bc!orc Ihc sccond onc can sIarI: onc Iask may rcp!cnsh
Ihc rcsourcc nccdcdbyanoIhcrIasksoIhcIwomayprocccd smu!Ia-
ncous!y: orIhc Iwo Iasksmay consumc Ihc samc rcsourcc n whch
casc Ihcrc canbc conBcIsbcIwccnIhcm. Thcsc rc!aIons-cnab!cs,
!ac!IaIcs, rp!cnshcs, and consumcs-a!!ow Tacms Io capIurc Ihc
sIrucIurc o! a Iask cnvronmcnI n Ihc !orm o! a hcrarchca! graph
IhcIcrmna!cndso!whcharcdrccI!ycxccuIab!c opcraIons.
Whcn uscd as a Ioo! Io cxp!orc Ihc ob|ccIvc possb!Iy spacc o!
Iask cnvronmcnIs Ihcsc drccI!y cxccuIab!c opcraIons musI bc !c!I
unspcchcd and Ihc dchnIon o! whaI Ihcy producc musI bc madc
vcry absIracIcapIurngon!yIhc conIrbuIon o! an opcraIon Io Ihc
so!uIon o! aprob!cm. Usng TacmsIhsways!ac!IaIcdby Ihc !acI
ILaI Is!orma!smwas dcsgncd Io bc cnIrc!y doman-ndcpcndcnI.
a graph rcprcscnIng a gvcn Iask dccomposIon may capIurc Ihc
sIrucIurc o! a Iask cnvronmcnI IhaI s common Io many domans
|a !acIory, a hospIa!, an arporI, a burcaucraIc agcncy) cxp!anng
why human agcnIs n Ihosc domans !acc sm!ar organzaIona!
cha!!cngcs. Thc dmcnsons o!Ihcpossb!Iyspacc arcIypca!!yprob-
ab!sIc bccausc Ihc propcrIcs o! !argc graphs can on!y bc gvcn
sIaIsIca!!y. Ihc avcragc numbcr o! subIasks nIo whch Iasks arc
I
Id
NLbN N b MUL1N
dccomposcd:Ihc avcragc duraIon o!opcraIonsandIasks: IhcsIaIs-
Ica! dsIrbuIon o!hard nIcrdcpcndcnccs |!kc cnab!cs) and so!I
nIcrdcpcndcnccs |!ikc !ac!IaIcs) . OnIhc oIhcr hand, Ihc graph
produccdbyTacms canbc coup!cdIo amu!IagcnIsysIcmIonvcsI-
gaIc a spccc Iask cnvronmcnI, n whch casc Ihc opcraIons as
wc!I as Ihcr producIs and duraIonsmusI bc !uI!y spcccd. In Ihc
casc o!pyramdbu!dngIhc drccI!ycxccuIab!copcraIonswou!dbc
acIvIcs!ikcmcasurng, a!gnng, chsc!ing,!cvcrng, cuIIng, po!sh-
ng.ThcscopcraIonscanbcassgncdIhc avcragcImc IhcyIakcIobc
pcr!ormcd and Ihc poIcnIa!va!uc o!Ihcr producI usngIhc rcsu!Is
!romcxpcrmcnIa!archco!ogy.Thsdscp!nc cana!soprovdcnsghI
nIo Ihc subdvson o! Iasks and subIasks Io bc incororaIcd nIo
aTacms graph. BuIinaddIonwcwou!dnccdanoIhcr graphspcc!y-
ng Ihc auIhorIysIucIurc o!Ihc rc!cvanIparIs o!Ihc gypIan gov-
crnmcnI. This can bc rcconsIrucIcd using cxsIng IcxIu! cvidcncc
rc!aIcdIoIhcbui!dngo!Ihc acIua!pyramds.
ConIro! o! Ihc ovcra!I Iask was in Ihc hands o! Ihc Vzcr n hs
ro!c o! Ovcrsccr o!Koya!Works. vdcncc !orIhs comcs !rom aIcxI
inscrbcdinhcrog!yphsonIhcwa!Iso!scvcra!Iombs!ownasThc
uIcso!IhcVzcr.FromIhsIcxIIsc!carIhaIauIhorIyro!cshad
noI ycI d!!crcnIaIcdnIo Ihcp!ura!Iyo! !uncIons wIhwhchwc
arc !am!arIoday. m parIcu!ar, Ihc Vzcrp!aycdscvcra!ro!cs. hcad
o!Ihc cv! admnsIraIonwIh|udca!andadmisIraIvc rcspons-
b!Ics: managing drccIor o! Ihc roya! pa!acc gcIIng da!y rcporIs
abouIsccurIyand!ogsIcmaIIcrs: anddcpuIyIoIhckng,pcrsona!Iy
hcarng hs commands and cnsurng Ihcr cnacImcnI Thc Vzcr
conIro!!cd Ihc cong!omcraIc o! ccnIra! dcparImcnIs consIIuIng Ihc
gypIan burcaucracyas wc!I asIhc!oca!auIhorIcs o!Ihcd!!crcnI
urban ccnIcrs and Ihcr rura! arcas. Thc IcxI a!so rcvca!s IhaI Ihc
d!!crcnIdcparImcnIswcrc rc!aIvc!yauIonomous. a!IhoughdcIa!cd
wrIIcn proccdurcs cxsIcd Io rcgu!aIc IhcracIvIcsIhc Vzcrdocs
noI sccmIohavc nIaIcdanyo!Ihc dcparImcnIs acIons bydrccI
commandbuIon!yIohavc supcrvscdIhc rcsu!Isa!IcrIhc !acI.Ths
suggcsIs a hcrrchca! coordnaIing mcchansm buI onc in whch
on!y sIraIcgc g

a!s wcrc gvcn as cxp!cI ordcrs wh!c Ihcr IacIca!

mp!cmcnIaIon was !c!I Io Ihc pcrso

c! wIhn Ihc dcparImcnIs.

OIhcrIcxIsgvcusIhcparo-who!crc!aIonsaIIhcboIIomcndo! Ihc
hcrarchy. Ihc work!orcc conssIcd o! scvcra! crcws, cach composcd
MUL1LN1b N KLNL b11Lb
o!Iwogangs o! I , 000mcn dvdcdnIovc groupso! 200, cach!ur-
Ihcr subdvdcd nIo Icn Icams o! 20 mcn ca!!cd dvsons. " Thc
ovcra!! hcrarchca! sIrucIurc may bc modc!cd by a graph n whch
nodcs rcprcscnI nsIIuIona! ro!cs, cdgcs rcprcscnI rc!aIonshps
bcIwccn Ihosc ro!cs, and arrows rcprcscnI asymmcIrca! rc!aIons,
suchasIhosc IhaIcxsIbcIwccnsupcrorsandsubordnaIcs. ThcunIy
o!commandundcrIhcVzcrmp!csagraphwIhasng!cnodcaIIhc
Iop, wh!c an unambguous chan o! command mp!cs a graph n
whcha!!nodcsbuIIhcIoponchavc asng!carrowpoinIngIoIhcm,
rcprcscnIngIhc !acIIhaI cvcrysubordnaIc hadasng!c drccIboss.
ThcscIwographs,oncdcscrbingIhccommandhcrarchyIhcoIhcr
dcnngIhc spcccaIonand dccomposIon o!Ihc ovcra!IIask, gvc
us an ob|ccIvc vcw o! Ihc organzaIon and Ihc prob!cm I musI
so!vc. BuI Io anmaIc Ihcm wIh agcnIs Ihc graphs musI bc supp!c-
mcnIcd by a p!ura!Iy o! parIa! vcws. Thc sub|ccIvc vcw o! Ihc
prob!cmvarcs!romagcnIIoagcnIandIs dcIcrmncdn!argcparI
byIhcwaynwhchIhcwork!orccsorganzcd.U IhsorganzaIons
achcvcdinargdhcrarchca!wayin whch commandsBowdown-
wards wh!c rcporIs Bowupwards, Ihc agcnIoccupyng Ihc Ioppos-
IonmusIbcmodc!cdashavngacccssIoIhc ob|ccIvc IasksIrucIurc
wh!c subordnaIc agcnIs shou!d havc on! a !ragmcnIary vcw, Ihc
morc!imIcdIhc !owcIhcrrank U Ihc organzaIonnc!udcdIcams
and!Ihcrmcmbcrswcrc!IowcdIocommuncaIcwIhoncanoIhcr,
IhcnagcnIsmay!ormnaddIonIoIhcr!oca!vicwsano-!oca!vcw
IhroughIhcin!ormaIonprovdcdbyIhcrpccrs.ThsdsIncIona!so
dcIcrmncs how nIcrdcpcndcnccs bcIwccn Iasks arc hand!cd by
agcnIs. n argdhcrarchyschcdu!cs anddcad!ncs!or cvcrysubIask
arc ccnIra!!y dccdcd and mp!cmcnIcd by command, wh!c wIhn
Icams agcnIs musI bc ab!c Io ncgoIaIc dcad!incs and makc commI-
mcnIs Io onc anoIhcr. m Ihc casc o! pyramd bu!dng Icams arc
nccdcdIo modc!whaIhappcns nsdc cach o!Ihchcrarchca!!cvc!s.
Thc zcrs o!cc, !or cxamp!c, conssIcdo! scvcra!pcop!c working
IogcIhcr. Ihc Vzcr hmsc!!, hs counc!, an o!ca! o!inIca! a!!ars,
and scvcra!scrbcs Tcams a!so cxsIcd nsdc cach rc!aIvc!y auIono-
mous govccnI dcparImcnI and aI Ihc !cvc! o! workcr dvsons
whcrc mosI physca! opcraIons wcrc pcr!ormcd. Ths comp!cxIy
ca!Is!oragcnIsIhaIarcnoI|usI boundcd!y raIona! | asnmosImu!I-
agcnI smu!aIons) buIa!soIask orcnIcdandsoca!Iy sIuaIcd.
IV
Id
NLbN N blMUL1N
Ina!!y, popu!aIng Ihc graphs wIh agnIs and scIIng Ihcm nIo
moIon dcmands smu!aIngarangc o!coordnaIngmcchansms.Thc
smp!csI and bcIIcr cxp!orcd o! Ihcsc arc prohbIons, IhaI s, simu-
lurcdscciulluws IhaIgcncraIc commImcnIs Io avod ccrIan acIons
wIhouIncurrngIhc cosIs o! communcaIon. Acxap!c o!proh-
bIonsarc Ira!c!aws. In a smu!aIon o! Ihc bu!dng o! a pyramd
Ihc nccd Io rcgu!aIc Ihc Ira!c o!agcnIs wou!d mmcdaIc!y bccomc
cvdcnI!IhcagcnIswcrcsIuaIcdnaIwo-dmcnsona!spacc. d!!cr-
cnIgangs wou!dhavcIobc assgncdIo d!!crcnI sdcs o!Ihc pyramd
andwIhn cach sdc agcnIsbrngng sIoncs, morIar, Ioo!s, !ood and
drnwou!dhavcIoavodco!!sonsandcrcaIc rc!aIvc!yc!ccnIcr
cu!aIon paIhs. Norcovcr, !wc addcdaIhrd dmcnson IoIhs spacc
IhcnasIhcpyramdgrcwIa!!crandIhc roomIomancuvcrdccrcascd
spaIa! assgnmcnIs a!!owng Ihc smu!Iancous pcr!ormancc o! Iasks
wou!d bccomc cvcn morc mporIanI. Crcu!aIon paIIcms cou!d bc
rgdIy spcccd by a ccnIra! conIro!!cr buI Ira!c!aws can bc a !css
cxpcnsvc coordnaIonmcchansm!Ihcyarccra!IcdIoIakcadvan
Iagc o! Ihc !acI IhaI agcnIs can pcrccvc onc anoIhcrwIhnccrIan
dsIancc and can Ihcrc!orc conIrbuIc Io Ihc so!uIon o! crcu!aIon
prob!cmsbymakng somc !oca! dccsonsIhcmsc!vcs.Idca!!y, a soca!
!aw shou!d gvc agcnIs cnough !rccdomo!acIonIo consIrucI ap!an
o!acIonndanopIma!paIh!romoncponIIoanoIhcr, !orcxam
p!c-buI noI so much IhaI Ihcy can nIcr!crc wIh Ihc acIons o!
oIhcrs.
Thc dynamcs bcIwccn d!!crcnI !cvc!s o! Ihc hcrarchy can bc
capIurcdbyaugmcnIngIhcnIcracIonsbcIwccnagcnIswIhspccch
acIs. Thcsc arc acIons n whch Ihc vcry uIIcrancc o! a scnIcncc
carrcsbndng soca!mp!caIons wIh d!!crcnI dcgrccs o! !orcc. Io
promsc,IorcqucsI,Iodcmand,andmosImporIanI!ynIhcprcscnI
casc, Io command. Usng Ihc graph IhaI capIurcs Ihc auIhorIy
sIrucIurc Ihc bndng !orcc o! Ihcsc commands can bc madc Io !!ow
!rom Ihcr ponI o! uIIcrancc Io Ihosc IhaI musI u!ImaIc!y cxccuIc
Ihcm. Thc dynam wIhnIcams, nIurn, canbc cxp!orcdusngan
approach ca!!cd Ccncra!IarIa!C!oba!I!annng |or CICI) . In Ihs
approach agcnIs usc Ihcr parIa! bc!c!s abouI a Iask Io !orm nIcn-
IonsIoacIandIo scqucncc Ihosc acIonsIocrcaIc a!oca! schcdu!c.
BccauscIhcconIcnIo!anagcnIsbc!c!sdocsnoInc!udcn!ormaIon
abouI oIhcr agcnIs Iasks Is !oca! schcdu!c may cnIcr nIo con!!cI
MUL1LN1b N KLNL b11Lb
wIhIhcrschcdu!cs. Thus, Ihc rsIcoordnaIngmcchansmnCICI
s uscd Io crcaIc non-!oca! vcwponIs. Ths mcchansm a!!ows an
agcnI Io gcI n!ormaIon abouI oIhcr agcnIs bc!c!s and Io dcIccI
coordnaIonprob!cms bcIwccn Is Iasks and Ihosc o! oIhcrs. A scc-
ond mcchansm a!!ows agcnIs Io communcaIc Ihc rcsu!Is o! Ihcr
opcraIonsIooIhcrsbuI on!yIhosc IhaI arc rc!cvanIsncc commun-
caIonIsc!!scosI!y. OIhcrmcchansmsarc IhcnuscdIoadagcnIsn
Ihc so!uIon o! prob!cms !kc avodng rcdundanccs and handIng
nIcrdcpcndcnccs. Thc cnd rcsu!I s IhaI agcnIs can modu!aIc Ihcr
!oca!schcdu!ng Ionc!udc commImcnIsIo oIhcrs. acommImcnIIo
mccI a dcadInc: Io sIop an acIon n ordcr Io !rcc a rcsourcc: orIo
sIarIanacIonaIIhccar!csIpossb!cImca!IcranoIhcragcnIsacIon
snshcd.
ArmcdwIhIhcsc d!!crcnI coordnaIngmcchansms |Ira!c!aws,
ccnIra! commands, ncgoIaIcd commImcnIs) Ishou!dbcpossb!c Io
usc a Tacms graph and a CICI mu!IagcnI sysIcm Io smu!aIc Ihc
comp!cx proccss o! bu!dng ganI !uncrary monumcnIs. cach scI o!
coordnaIng mcchansms IogcIhcr wIh a proposcd dccomposIon
o! Ihc ovcra!! Iask nIo subIasks wou!d consIIuIc an archco!ogca!
hypoIhcss abouIwhaIrca!!ywcnIondurngIhc consIrucon o!Ihc
grcaI Cza pyramd. Such a smu!aIon wou!d capIurc on!y a brc!
cpsodc n Ihc hsIory o! anccnI gypI and wou!d nc!udc on!y a
scgmcnI o!gypIan soccIy. Ihc Vzcr as Ovcrsccr o! Koya! Works,
Ihc rc!cvanI dcparImcnI undcr D, andIhc crcws o! workcrs undcr
Ihc command o! IhaI dcparImcnI. Ths mcans IhaI much o! whaI
madcIhcworkngso!Ihs soca!scgmcnIpossb!c wou!dbcnc!udcd
on!y mp!cI!y. Thc provnccs IhaI supp!cd !ood and !abor Io Ihc
ccnIra!govcrnmcnI,!orcxamp!c,wou!dbcnc!udcdon!yasrcsourcc
npuIs. BuI as wc havc argucdrcpcaIcdIynprcvouschapIcrsIhcrc
s noIhng wrong wIh uncxp!ancd prcsupposIons as !ong as a
smu!aIon cxp!ans somcIhng c!sc, suchasIhc cmcrgcnIcapacIyo!
anorganzaIonIop!anandbu!d!argcarchIccIura!sIrucIurcs,andas
!ong as Ihcrc arc oIhcr modc!s IhaI accounI !or Ihc cmcrgcncc o!
whaIsbcngprcsupposcd.
Ina!!y,IhcrcsIhc qucsIono!howIo smu!aIcIhc organzaIona!
!carnngproccssIhaI!cdIoa spccc scIo!coordnaIngmcchansms.
Thc ccnIuryo!pyramdbu!dng cxpcrcnccbc!orc Cza s Ioo shorI
!or an cvo!uIonary proccss Io havc yc!dcd any sgncanI rcsu!Is
IdI
IdZ
NLbN N bMUL1N
cvcn ! augmcnIcd by rcn!orccmcnI !canng. Ths mcans IhaI wc
havc nochoccbuIIo usc pcrsona!agcnIs capab!c o!monIorng Ihc
cxccuIon o!Ihcr ownIasks andIo dcIccI prob!cms !kcIhc pcr!or-
mancc o! rcdundanI subIasks, msIakcs n Ihc scqucncng o! opcra-
Ions, msa!!ocaIcd rcsourccs, or brokcn commImcnIs. Norc
mporIanI!y, oncc aprob!cms dcIccIcdagcnIsmusIbcab!cIo dag-
nosc Is causc and proposc a so!uIon. To havc Ihs capacIy agcnIs
musIuscIhcporIono!IhcTacmsgraphIowhchIhcyhavc acccss Io
!ocaIc Ihc dcIccIcdprob!cm n IspropcrconIcxI-Io dcIcrminc, !or
cxamp!c, ! nIcrdcpcndcnccs among subIasks wcrc onc o! Ihc
causcs-as wc!! as usc Iraccs o! pasI acIvIy rccordng noI on!y Ihc
scrcs o! dccsons Ihcy Iook abouI whaI opcraIon Io cxccuIc and
whcn Io cxccuIc I, buI a!so Ihc rcasons why Ihosc dccsons wcrc
madc. m Ihs schcmc an agcnI has drccI acccss on!y Io Is own
mcmory Iraccs buI n Ihc prcscncc o! Iask nIcrdcpcndcnccs I may
havc Io rcqucsI n!ormaIon abouI Ihc Iraccs o! oIhcr agcnIs Io bc
ab!c Io rcconsIrucI Ihc proccss IhaI !cd Io Ihc prob!cm, and Ihcsc
rcqucIscanbc cosI!y. To kccp communcaIon cosIs!ow rc!aIvc Io
Ihcgansnc!ccncyagcnIsshou!daIIcmpIIo dagnosc on!ysIua-
Ion-spccc prob!cms. m oIhcr words, agcnIs shou!d Iry Io gaIhcr
non-!oca!n!ormaIonbuIon!yIoIhc cxIcnIIhaII canbc cxp!oIcd
!oca!!y.
Through a proccss !kc Ihs an organzaIon can !carn Io mproc
coordnaIng mcchansms buI on!y aI Ihc !cvc! o! Is Icams. ThaI s,
oncc a corrccI dagnoss s madc modcaIons Io coordnaIon ru!cs
wi!Ibc app!cdon!y!oca!!y.Ths approach, on Ihc oIhcrhand, canbc
cxIcndcd Io Ihc cnIrc hcrarchy by app!yng IIo a!!Ihc d!!crcnI
ranks. HsIorca!!y, onco!IhcrsI organzaIona! ro!csIod!!crcnIaIc
bcIwccn a chc! and hs !o!!owcrs musI havc bccn IhaI o! ovcrsccr.
ar!y on nIo gypIan dynasIc hsIory scvcra! ranks o! ovcrsccrs
a!rcady cxsIcd. BuIwhaIs an ovcrsccr ! noI anagcnIIhaIspcca!
zcs n monIorng and dcIccIngprob!cms occurrng aI Ihc rank|usI
bc!owandcrcaIngcporIscxp!anngprob!cmsaddrcsscdIoIhoscn
Ihcrank|usIabvcm oIhcrwords, Ihc samc dagnosngcapab!Ics
IhaIIcammcmbcrs havcnIhs schcmc cou!dyc!danagcnIp!ayng
Ihcro!c o!ovcrsccr. Thc d!!crcncc bcIwccnIhcIwo cascs sIhaI Ihc
hghcrIhcranko!anovcrsccrIhc!argcrIhc porIono!aTacmsgraph
I wou!d havc acccss Io. Ths mcans IhaI an ovcrsccr o! su!ccnI!y
MUL1LN1b N KLNL b11Lb
hgh rank wou!d noI !carn Iomprovc Ihc scqucncng o! opcraIons
!kcsIonccuIIngorpo!shng,buIIhcscdu!ngo!cnIrcIaskgroups
!kc quarryng, hau!ng, andcrccIng. Thc modcaIons Io coordna-
Ionmcchansms wou!dbc sI!!bc!oca!and orcnIcd Io spcccIasks
buI Ihc hghcr Ihc rank Ihc morc gcncra! n scopc Ihcc !oca! so!u-
Ions wou!dbc.
Ikc gcncIca!gorIhms andncura!ncIs, mu!IagcnI sysIcmshavca
varcIyo!ndusIra!uscsIhaIhavcnoIhngIodowIhmodc!ngsoca!
rca!Iy. Thcsc rangc !rom crcaIng uscr inIcr!accs !or ncw so!Iwarc
app!caIonsIhaIcan!carnabouIIhchabIso!human uscrs, Ioncrcas-
ng Ihc c!ccncy o! Ihc a!!ocaIon o! rcsourccs m Ihc nIcmcI by
a!!owngauIonomous agcnIsIo cngagc mdcccnIra!zcdIradc. Thcsc
and oIhcr uscs o! mu!IagcnI sysIcms W conInuc Io drvc Ihcr
dcvc!opmcnIrcgard!csso!IhcraIcaIwhchsoca!sccnIsIsadopIIhs
Icchno!ogy !or modc!ng purposcs. BuI I has bccn Ihc undcr!yng
mcssagc o! Ihs chapIcr IhaI soca! smu!aIons as cnacIcd IhoughI
cxpcrmcnIscan grcaI!yconIrbuIc Io dcvc!opnsghInIoIhc work-
ingso!IhcmosIcomp!cxcmcrgcnIwho!csonIhsp!ancI.WhcIhcrIs
raIco!adopIonaccc!craIcsornoIwi!!dcpcndonIhcmagnaIvc usc
IhaI soca!sccnIsIs makc o! Ihcsc ncwrcsourccs aswc!! as onIhcr
w!!ngncss Io abandon Ihc dca o! soccIy as a who!c andrcp!acc
I wIh a scI o! morc concrcIc cnIIcs |communIcs, organzaIons,
cIcs) IhaI !cnd Ihcmsc!vcs IoparIa! modc!ing m a way IhaI vaguc
IoIa!Ics do noI. m shorI, Ihc !uIurc o! mu!IagcnI smu!aIons as
modc!so!soca!rca!Iywi!!dcpcndonhowsoca!sccnIsIscana!!ccI
Ihs Icchno!ogy by dcp!oyng I crcaIvc!y and on how Ihcy can bc
a!!ccIcdby I IhroughIhcposscssono!IhcrghI soca!onIo!ogy.
IdJ
Id9
PPLN
L|K5 |O 55O00|dO 10OO|V
ThcvarcIyo!mcchansmso!cmcrgcncc dscusscdnIhsbookshow
IhaIIhc !cars o!Ihc car!y cmcrgcnIsIs, IhaI c!ockwork mcchansms
cou!dnoIpossb!y accounI !or Ihc rrcducb!c propcrIcs o! a who!c,
wcrc msp!accd. Thcrc snoIhngn Ihc noIon o!aho!sIcpropcrIy
IhaI cmcrgcs !rom Ihc nIcracIons bcIwccn componcnIs IhaI !mIs
IhcpoIcnIa!comp!cxIyo!IhoscnIcracIons. OnIhc oIhcrhand,Ihc
conccpIo!cmcrgcnIpropcrIydocsp!acc!mIsonIhckndso!who!cs
IhaIcanbc !cgImaIc!yacccpIcd. mparIcu!ar, Im!IaIcsagansIIhc
dca o! a IoIa!Iy Ihc parIs o! whch arc so ncxIrcab!y rc!aIcd IhaI
IhcrvcrydcnIIys consIIuIcd by Ihcrrc!aIonswIhn Ihc who!c.
m a scam!css IoIa!Iy Ihc propcrIcs o! componcnIs dcrvc!romIhc
ro!c Ihcyp!ay n Ihc who!c so dcIachng Ihcm!romI wou!d auIo-
maIca!!y dcprvc Ihcm!romIhcr dcnng characIcrsIcs. BuI nonc
o!Ihcwho!cswc havc dscusscd hcrc, !romIhundcrsIormsIonsIIu-
Iona!organzaIons, arc scam!css IoIa!Ics.Thc !acIIhaIIhc propcr-
Ics o! a who!c dcpcnd on Ihc acIua!cxcrcsc o!Ihc capacIcs o!Is
parIs mp!cs IhaI rcmovng onc o! Ihcm may ndccd dcsIroy Ihc
who!csdcnIIy,buIIhcparIIsc!!nccdnoI!oscIsowndcnIIy.pu!!
nga!vc anma!s hcarI ouIw!!surc!yK IbuI Ihc hcarIIsc!!can
bc mp!anIcd nIo anoIhcr anma! and rcsumc Is rcgu!ar !uncIon.
Thus, whaIwc nccdsaconccpIIhaIa!!owsusIorcIanboIhrrcduc
b!Iy and dccomposab!Iy, a conccpI IhaI makcs Ihc cxp!anaIon
o! synIhcss and Ihc possb!Iy o! ana!yss nIc!!gb!c. Wc cou!d, o!
coursc, smp!ybu!d IhsrcqurcmcnInIo Ihc dcnIon o!Ihc Icrm
cmcrgcnI who!c buI sncc Ihs conccpI has a hsIory, and sncc n
llLNA
Ihs hsIory Ihc rcsIrcIon agansI IoIa!Ics has noI a!ways bccn
cn!orccd, I W bcusc!u!Io nIroducc a ncwword!orwho!csIhaI
arc rrcducb!c and dccomposab!c. Wc w!! rc!cr Io Ihcsc who!cs as
usscmblujcs. '
Thc advanIagc o! a ncwIcrm s noI !mIcd Io Ihc !acI IhaI I s
unb!cmshcd by pasI ph!osophca! msIakcs or IhaI I s dcvod o!
undcsrab!c connoIaIons. II a!so sgna!s a !rcsh sIarI and acIs as an
nccnIvc IogcIIhngsrghIIhsImcaround. So!Iscarc!u!!ydcnc
whaIc!scshou!dbcnc!udcdnIhcconIcnIo!IhsncwconccpI.IrsI
o!a!!,Ihc dcnIIyo!anasscmb!agc shou!da!waysbc conccvcdasIc
producI o! a hsIorca! proccss, Ihc proccss IhaI broughI Is compo
ncnIs IogcIhcr!orIhc rsIImc as wc!!asIhc proccssIhaImanIans
IsnIcgrIyIhrougharcgu!arnIcracIonamongIsparIs. Thsmp!cs
IhaI Ihc dcnIIy o! an asscmb!agc s a!ways conIngcnI andI s noI
guaranIccdbyIhccxsIcncco!ancccssaryscIo!propcrIcsconsIIuI-
ng unchangngcsscncc. OrIopuIIhsd!!crcnI!y, asscmb!agcsarc
noIparIcu!armcmbcrs o!agcncra! caIcgorybuIunquc andsngu!ar
ndvdua!s. vcn ! Iwo asscmb!agcs rcscmb!c cach oIhcr so much
IhaI no onc can Ic!! Ihcm aparI, cach w!!sI!!bc unquc duc Io Ihc
d!!crcnIdcIa!so!Isndvdua!hsIory. mIhc!asI!cwchapIcrsIwas
argucd IhaI dcspIc Ihc !acI IhaI Ihc word ndvdua! has bccomc
synonymouswIhpcrsonIcan!cgImaIc!ybcapp!cdIondvdua!
communIcs, ndvdua! organzaIons, or ndvdua! cIcs. And Ihs
argumcnIcanbc cxIcndcd IoIhc cmcrgcnI cnIIcs o!a!!oIhcrchap
Icrs. ndvdua! aIoms and mo!ccu!cs, ndvdua! cc!!sand organsms,
ndvdua! spcccs, and ccosysIcms. A!! hcsc d!!crcnI asscmb!agcs
arc born aI aparIcu!arImc, !vc a !!c, and Ihcn dc. II !o!!owsIhaI
know!cdgc abouI an asscmb!agc docs noI dcrvc !rom a boIanca!
c!asscaIonhaIIakcspropcrIcs !orgranIcdbuI !rom an accounI o!
Ihc orgn and cnduancc o! Ihosc propcrIcs. Cvcn Ihc mporIancc
o!Ihc onIo!ogca! sIaIus o!asscmb!agcswc w!!nccdaIcchnca!Icrm
Io rc!cr IoI. cvcry acIua! asscmb!agc s an individuulsinjulurin.
A sccondonIo!ogca!commImcnI musIbcbu!InIoIhc dcnIon
o!Ihc Icrm asscmb!agc bccausc Ihcsc cmcrgcnI who!cs arc dcncd
noI on!yby Ihcrpropcrcs buI a!so byIhcrIcndcnccs and capac
Ics. Tcndcnccs canmakc Ihc propcrIcs o!a wh!c vary, somcImcs
cvcn changng Is dcnIIy, aswhcnan cc scu!pIurccharacIcrzcdby
Is so!dIy and Is shapc man!csIs IsIcndcncy Io mc!I aI a ccrIan
Idb
Id
LNA
IcmpcraIurc, !osngIhoscpropcrIcsandmuch o!IsdcnIIy. Capac-
Ics, iDIu, makc who!cs cxhbI aspccIs o! IhcrdcnIIyIhaIwcrc
prcvous!yhddcn, aswhcnannnoccnI!ookngp!anIIums ouIIobc
posonous or, onIhc conIrary, Ioposscss uncxpccIcdhca!ngpowcrs.
Whi!c Icndcnccsmakc a !sI o! csscnIa! propcrIcs !ook!a!sc!y pcr-
mancnI capacIcs cxp!odc I siDcc !or a gvcnwho!c I wou!d bc
mpossb!c Io!sIa!!Ihc d!!crcnIwaysnwhchIcana!!ccI andbc
a!!ccIcdbynnumcrab!c oIhcr who!cs. Andcvcn!wc cou!d!sI a!!
acIua!!y man!csIcdIcndcnccs and acIua!!y cxcrcscd capacIcs IhaI
wou!dsIi!!noIbc cnough siDccncIhcronchasIobcacIua!.ToIack!c
Ihsprob!cmwc uscdIhcnoIono!IhcsIrucIurc o!apossb!Iyspacc,
a sIrucIurc IhaI cxp!ans how Icndcnccs and capacIcs can bc rca!
cvcn whcn Ihcy arc noI acIua!. A!Ihough wc saw how varcd Ihs
sIrucIurc can bc and acknow!cdgcd many gaps n our conccpIon
o!I, I s sa!c Io say IhaI I s characIcrzcd by spcca! or rcmarkab!c
IraIs Ihc onIo!ogca! sIaIus o! whch s IhaI o! univcrsulsinjulurirics.
Thc Icrm unvcrsa! s uscd!or Iwo rc!aIcdrcasons. rsI, Ihc cvcnIs
inwhchIcndcnccs arc man!csIcd and capacIcs cxcrcscdmay bc
cnIrc!yd!!crc

IindcIa!andycIbcshapcdbyIhc samcsingu!arIcs:
and sccond, Iwo scrcs o!cvcnIs cachconsIIuIingad!!crcnImccha-
nsmmayncvcrIhc!cssposscss ovcr!appngpossb!Iyspaccsandds-
p!aycommon!caIurcsIhaIarcmcchansmndcpcndcnI.
IaddIonIoIhcsconIo!ogca!commImcnIsanoIhcrrcqurcmcnI
musI bcbu!InIo Ihc dcnIono!Ihc Icrm asscmb!agc. A!Ihough
cach asscmb!agc s a unquc hsIorca! cnIIy I a!ways bc!ongs Io a
popu!aIono!morc or!csssim!arasscmb!agcs.InoIhcrwords, dcspIc
Ihc ndvdua! singu!arIy o!cachasscmb!agcIhc proccss o!asscmb!y
bchiDdIIcnds Iobc rccurrcnIso whaIs synIhcszcds ncvcr asng!c
ndvdua!buImany o!Ihcm.Thcrccurrcncc Isc!!scxp!ancdbyIhc
!acI IhaI Ihc asscmb!y proccss s govcmcd by unvcrsa! sngu!arIcs
buI Ihc acIua!zaIon o! Ihc !aIIcr s a!ways sub|ccI Io conIingcnI
cvcnIs so whaI s gcncraIcd s a popu!aIon in whch varanIs arc
dsIrbuIcdina ccIanway. Thc !acIIhaI sharcdpropcrIcsvary, and
IhaIIhcvaraIonnapopu!aIondsp!ays accrIaiDsIaIsIca!dsIrbu-
Ion, s anoIhcr rcason why Ihc dcnIIy o! an asscmb!agc shou!d
noI bc conccvcd n Icrms o! a xcd !sI o! propcrIcs. To !acIIaIc
popu!aIonIhnkng wc nccda mcans Io spcc!y noI on!y Ihc pos-
sb!cwaysnwhchIhcmcmbcrso!apopu!aIoncanchangcbuIa!so
LNA
IhcsIaIc o!IhrdcnIIyaIanyparIcu!arpoiDI nIhcr hsIory. This
can bc achcvcd by rurumcrrizinj Ihc conccpI o! asscmb!agc, IhaI s,
by provdng I wIh knobs wIh modab!c scIIngs Ihc va!ucs o!
whchdcIcrmnc Ihc condIono!IhcdcnIIy o! an cmcrgcnIwho!c
aI anygvcnIimc. Ths wi!I makc I cascrIopcIurc nourmnds an
cnIrc popu!aIon o! asscmb!agcs cachposscssiDg a sIghI!y d!!crcnI
dcnIIy dcpcndng onIhc scIIiDgs o!IhcparamcIcrs. AndI wi!I aIso
!aci!iIaIc Ihnkng o!Ihc popu!aIonas ancnIIyIhaIcanundcrgoan
iDIcnscshock!romIhc ouIsdcIhaIdrasIcaI!ychangcsIhosc scIIiDgs
producingvaranIsIhaI go bcyond Ihc norma! rangc o! varaIon. I
shorI, a paramcIrzcd conccpI canpopu!aIc ourmnds iD away IhaI
brngs varaIon andIhc hsIory o!IhaIvaraIonwIhI.
OncparamcIcrmusI spcc!yIhcrc!aIvc homogcncIyorhcIcrogc-
ncIyo!IhccomponcnIs o!anasscmb!agc!nagvcnpopu!aIona!!
asscmb!agcs usc Ihc cxacI samc componcnIs Ihcy wi!I sharc abcIIcr
dcncddcnIIyIhan!Ihcrcs!argcdcgrcco!varaIoniDIhosccom-
poncnIs. ThsparamcIcrmusI a!sospcc!yIhc sIaIc o!Ihcboundarcs
o!an asscmb!agc sharpandxcdor, onIhc conIrary, !uzzyanduc-
IuaIng. Thc Icrm boundary rc!crs Io vcry d!!crcnIIhngs dcpcnd-
ng on Ihc asscmb!agc Ihc ouIcr shc!! o! c!ccIrons n an aIom Ihc
mcmbranc o! a bacIcrum: Ihc ouIcr skiD o! a p!anI or anima!: Ihc
gcographcaI !ronIcrs o! an ccosysIcm: Ihc rcproducIvc barrcrs o!
a spcccs. ThcparamcIcrnccds anamc and sncc I parIIy dcIcrmncs
dcnng boundarcs wc can ca!!I rcrrircriulizuricn. 'Thc morc homo-
gcncous Ihc inIcrna! composIon o! an asscmb!agc and Ihc bcIIcr
dcncdIsouIcrboundarcs Ihc morc IcrrIoraIzcdIs dcnIIy may
bc sad Io bc. I addIon, gvcnIhaI Icndcnccs and capacIcs a!so
dcIcrmncdcnIIywcmusInc!udcbchavora! !acIorsnIhcscparam-
cIcr. Forcxamp!c, onc!acIora!!ccIingIhcrangco!capacIcsacIua!!y
cxcrcscdby anasscmb!agc s mobi!iIy, sincc a mobi!c asscmb!agc s
morc !ikc!y Io cncounIcr novc! sIuaIons Ihan animmobi!c onc. I
Ihs scnsc wcmaysayIhaIananima!IhaI canmovc aroundandIhaI
can a!!ccI and bc a!!ccIcd by a wdcr rangc o! oIhcr cnIIcs s morc
dcIcrrIoraIzcdIhanap!anIIhaIsIcdIoIhc soi!.
Forsomcasscmb!agcswcmaynccdmorcIhanoncparamcIcrsncc
ncw ways o! dcIcrmnng dcnII havc cmcrgcd hsIorca!!y. nc
novc!mcans o!xngcnIIysIhcgcncIccodc, ormorcprccsc!y, Ihc
comp!cx machncry Ihrough whch gcncIc in!ormaIon s cxprcsscd.
Id
Idd
LNA
Ccncscancxcrcsc IghI conIro! ovcrIhcmcchansms o! cmcrgcncc
bchnd organc who!cs, as whcn Ihcy coordnaIc Ihc gradcnIs and
cmcrgcnI !orms o! a dcvc!opng cmbryo. An obvous namc !o Ihs
ncw paramcIcr wou!dbc ccdinj. An organsm, !or cxamp!c, may bc
sadIo bc hgh!y codcd!cvcrydcIa!o!IsanaIomys rgd!y dcIcr-
mncd by Is gcncs and rc!aIvc!y dccodcd H Ihc cnvronmcnI a!so
conIrbuIcs Io Is anaIomca! dcmIon. And sm!ar!y !or bchavor.
HIhcbchavoro!anorgansmscnIrc!ynhcrIcdIhccodngparam-
cIcr W havc a hgh va!uc and vcc vcrsa, ! gcncs dcnc onIy Ihc
connccIvIyo!Isncurons a!!owng mucho!IsbchavorIobc dcIcr-
mncd by!carnngdurngIs!!cImc Ihc paramcIcrw!!havc a !ow
va!uc. nIIcs oIhcr Ihan gcncs can a!!ccI Ihs paramcIcr. Nuch as
IhcIcrrIora!zaIonparamcIcrcanapp!yIoa!!sorIso!boundarcs so
IhccodngparamcIcrcanbcapp!cdIooIhcrsourccso!n!ormaIon-
bascd consIranIs, such as Ihosc provdcd by!anguagc. Wc canmag-
nc a communIy o!pa!co!IhchunIcrgaIhcrcrs, !or cxamp!c, usng
Isncw!yacqurcdpowcro!spccchIorgd!ycaIcgorzca!IIhc cnIIcs
IhaI arc mporIanI !or Is conInucd cxsIcncc. !ood cou!d bc codcd
asraworcookcd: sacrcd, Iaboo, orordnary. And sm!ar!y !ordrcss,
bchavor, andoIhcrcu!Iura!cxprcssonso!IsdcnIIyAnoIhcrgood
cxamp!c wou!d bc Ihc nsIIuIona! organzaIons bc!ongng Io
archac sIaIc IhaI codc n wrIIcn !om d Ihc maIcra! !Iows Ihcy
conIro!. BoIh o! Ihcsc asscmb!agcs may bc sad Io bc hgh!y codcd.
Convcrsc!y, Ihcrc may cxsI a !cw commcrca! organzaIons in Ihc
pcrphcry o! an archac sIaIc IhaI coordnaIc maIcra! !Iows usng
prccsnsIcado!o!ca!caIcgorcsandccnIra!commands,soasasscm-
b!agcsIhcsc orgamzaIonsmaybcsadIobc rc!aIvc!y dccodcd.
IcIs summarzc whaIhas bccn sadabouI Ihc Icrm asscmb!agc
and IhcnrcvcwIhc conIcnIo!prcvouschapIcrs n Ihc !ghI o!Ihs
ncw conccpI. IrsI o! a!!, IhcIcrmmusI rc!cr Io concrcIc who!cs IhaI
arc rrcducb!c ad dccomposab!c, IhaI s, I musI accommodaIc Ihc
cpsIcmo!ogca!dcmandso!boIhsynIhcssandana!yss.Thcscwho!cs
musI bc assgncd a c!car onIo!ogca! sIaIus. vcry asscmb!agc musI
bc IrcaIcd as unquc hsIorca! cnIIy characIcrzcd boIh by a scI
o!acIua!cmcrgcnIpropcrIcs|makngIanndvdua!sngu!arIy) as
wc!! as by Ihc sIrucIurc o! possb!Iy spaccs dcnng Is Icndcnccs
and capacIcs |a sIrucIurc dcncd by unvcrsa! sngu!arIcs) . Ths
sIrucIurcmaybcIcmcdIhcdiujrumo!Ihcasscm!agc.ThchsIorca!Iy
LNA
conIngcnIdcnIIyo! asscmb!agcsmusIbc a!!owcdIochangc soIhc
conccpImusIbcsupp!cdwIhoncormorc paramcIcrs dcnngIhosc
changcs.Andna!!y, asscmb!agcsmusIaIwaysbcIhoughIasparIso!
popu!aIonsmwhchIhcrdcnIIcscanchangcwIhn!mIsproduc-
ng a ccrIan sIaIsIca! dsIrbuIon o! varaIon. Thc poIcnIa! rangc
andpossb!c !ncs o!varaIon arc dcIcrmncd byIhc dagram sharcd
by a!! popu!aIon mcmbcrs buI Ihcr acIua! dcgrcc o! sm!arIy s
dcIcrmncd by Ihc va!ucs o! Ihc paramcIcrs aI anygvcn Imc Ihc
morc IcrrIora!zcd or codcd Ihc mcmbcrs o! Ihc popu!aIon arcIhc
morcIhcyW IcndIorcscmb!concanoIhcr.ThswayIhcsm!arIcs
IhaI IcmpI us Io c!ass!y Ihc popu!aIons mcmbcrs undcr a gcncra!
caIcgory, and Ihcn Io rc!y IhaI caIcgory, can bc gvcn a concrcIc
hsIorca!cxp!anaIon.
m ChapIcr I wc cxamncdIhc smp!csI way o! crcaIng an asscm-
b!agc. p!acng Iwo scparaIc mo!ccu!arpopu!aIons |Iwobodcso! ar
or waIcr) aI d!!crcnI IcmpcraIurc or prcssurc n conIacI wIh cach
oIhcr.Ths, o! coursc, crcaIcs a gradcnIIhIhc IcndcncyIo dss-
paIc and Ihc capacIyIo acI as !uc! !or oIhcrproccsscs. Bccausc Ihc
Iwo popu!aIons arc dcnIca!B composIon Ihc onIy hcIcrogcncIy
nvo!vcd s Ihc nIcnsvc dcrcncc Isc!!. Thc IcrrIora!zaIon para-
mcIcr rcduccs m Ihs smp!csI o! a!Iasscmb!agcs Io Ihc disruncc]cm
rhcrmcdynumic cuuilibrium. IhaI s, Io Ihc dcgrcc o! nIcnsIy o! Ihc
gradcnI.Thcdagramo!Ihsasscmb!agccanbcsIudcdwIhIhchc!p
o! maIhcmaIca! Ioo!s !kc sIaIc spacc, Is Ira|ccIorcs, and sngu!ar-
Ics. Thc dmcnsons o!Ihc possb!Iy spacc arc Ihc rc!cvanI ways o!
changng!oran asscmb!agc, IhaIs,Is dcgrccso!!rccdom.U Ihcrcarc
nIcracIonsbcIwccnIhcscdcgrccso!!rccdomIhcdagramsnonIn-
car, posscssng mu!Ip!c sngu!arIcs o! d!!crcnI Iypcs, whcrcas !
Ihcrc arc nonIcracIonsIhc dagrams!ncar, sIrucIurcdby a sng!c
sngu!arIy o! Ihc sIcady-sIaIc Iypc. Ina!Iy, somc o! Ihc mcmbcrs o!
Ihcscmo!ccu!arpopu!aIonsmaybcrccruIcdIo!ormIhccomponcnI
parIs o! !argcr asscmb!agcs Ihc dcnIIy o!whchw!!bc dcIcrmncd
mparIbyIhc!ncarornonIncarnaIurc o!IhcdagramandmparIby
IhcrownIcrrIora!zaIonparamcIcr.mcrgcnIwho!csIhaI!ormb
dsspaIng a!oca!gradcnI andrcachng a !ow-nIcnsIy cqu!brum,
such as a mnmum o!bondng cncrgynIhc casc o!cc crysIa!s ora
mnmum o!sur!acc IcnsonmIhc casc o!arbubb!cs, maybc sadIo
bc hghIy IcrrIora!zcd. Thosc IhaI, on Ihc conIrary, cmcrgc whcn
IdV
IV
LNA
a IocaI gradicnI is inIcnsc cnough andiI isnoI aIIowcd Io dissipaIc,
Iikc Ihc paIIcm oI Iowdcninga convccIion ccII, maybc saidIo bc
morc dcIcrriIoriaIizcd.
I ChapIcr 3 wc gcncraIcdIhcncxIIaycr oI compIcxiIy byinIro
ducing chcmicaI inIcracIions and Ihc synIhcsis oI novcI moIccuIcs,
grcaIIyincrcasingIhchcIcrogcnciIy oIIhc moIccuIarpopuIaIions, as
wcII as caIaIysIs Io manipuIaIc Ihc rapidiIy or sIowncss wiIh which
such synIhcscs Iakc pIacc. I somc cascs Ihc ouIcomc oI chcmicaI
inIcracIionsisancquiIbriumconccnIraIionoIIhcproducIs, aIcrriIo-
riaIizcddisIribuIion oIsubsIanccswiIhasIabIcidcnIiIy. IoIhcrsIhc
caIaIyIic conIroI oI rcacIion raIcs is pcrIormcd U a Iooponc sub
sIancc acccIcraIingIhcproducIion oIanoIhcrsubsIanccIhaI, inIurn,
acccIcraIcs Ihc raIc oI producIion oIIhc rsIdrvingIhc moIccuIar
popuIaIions awayIrom cquiIbrium andIcadingIo Ihc cmcrgcncc oI
morc or Icss dcIcrrIoriaIizcd whoIcs. chcmicaI cIocks or morc com-
pIcx auIocaIaIyIic Ioops. AIIhough Ihc idcnIiIy oI an auIocaIaIyIic
IoopissIabiIzcdbyiIs comoncnIsiIis aIsosub|ccIIo changc as ncw
componcnIs arc addcd IhaI caIaIyzc and arc caIaIyzcd by cxisIing
oncs. ThcdiagramoIIhcsc asscmbIagcscansIiIIbc sIudicdusingsIaIc
spaccbuInowIhc numbcr oIdimcnsions oIIhc spacc ndIhc diIIcr-
cnIiaI rcIaIions IhaI dcnc Ihc disIribuIion oI singuIariIics musI bc
aIIowcdIo changcIo rcBccI Ihc IacI IhaI cnIircIyncwsubsIanccs can
bcproduccdandncw chcmicaIrcacIionsmadcpossibIc. TokccpIrack
oIIhc changing dimcnsions oIIhc possibiIIy spacc wc nccd a graph
inwhichIhcnodcsrcprcscnIsubsIanccsandIhc cdgcs chcmicaIrcac-
Iions. A imporIanI singuIariIy in Ihc graph iIscII is a pcrcoIaIion
IhrcshoId, a criIicaI dcgrcc oI connccIiviIy aI which Ihc numbcr oI
possibIc rcacIions changcsIromniIcIo inniIc. AIIcrIhis IhrcshoId
is rcachcd Ihc idcnIiIy oI Ihc producIs oI chcmicaI inIcracIions
canvaryin aninniIcnumbcr oIpossibIc ways, orwhaI amounIs Io
Ihc samc Ihing, Ihc diagram bccomcs IIcd wiIh poIcnIiaI Incs oI
dcIcrri IoriaIizaIin.
ChapIcr 4 addcd onc morc Iaycr oI compIcxiIyby inIroducing Ihc
capaciIyIorscIrcpIicaIioninIomoIccuIarpopuIaIions. ApopuIaIion
oIrcpIicaIors. such as moIccuIcs oInacdA, can IoIIow a paIh oI
conIinuouschangcandadapIaIionasIongascopyingcrrorsandoIhcr
muIaIionsmodiIyIhcidcnIiIyoIiIsmcmbcrs.ThcdiscmbodicdrcpIi-
caIorsIcndIoIormcmcrgcnIcnIiIicscaIIcdquasi-spccics,cIouds oI
LNA
muIanIsIhaI cvoIvc as awhoIc.ToomuchaccuracyinIhcrcpIicaIion
proccss Icads Io a high vaIuc IorIhc IcrriIoriaIizaIionparamcIcrand
a Iowcr capaciIyIo Iormquasi-spccics. BuI Ihcrc is aIso a dcIcrriIori-
aIizaIion IhrcshoId consIiIuIcd by Ioo high a muIaIion raIc IhaI
dissoIvcs Ihc idcnIiIy oI Ihc muIanI cIoud. Thc diagram oI Ihcsc
asscmbIagcsmusI capIurcIhc combinaIoriaI possibiIIics oIIhc rcpIi-
caIors as wcII as Ihc possibIc diIIcrcnccs in rcpIicaIivc succcss IhaI
wouId rcsuII H Ihc popuIaIion conIronIcdIimIcdrcsourccs. P pos-
sibIc Ascqucnccs canbc arrangcdsoIhaI Ihcyhavcas ncighbors
oIhcr scqucnccs diIIcring onIy by onc muIaIion. I such a spacc a
scrics oI onc-muIanI ncighbors dcncs a possibIc paIhway Ior idcn-
IiIy Io changc, a possibIc Inc oI dcIcrriIoriaIizaIion IhaI can Iakc a
moIccuIc oIA andIransIormiI sIcpby sIcpinIoan cnIircIy diIIcr-
cnIonc.BuIwhcIhcrsuchpaIhwayscanacIuaIIybc IoIIowcddcpcnds
onIhc disIribuIionoIIncss vaIucs.Thcsc Iorm aIopography oIvaI-
IcysandhiIIssupcrimposcdonIhccombinaIoriaIspacc, aIopography
IhaI dcIcrmincs Ihc IikcIihood IhaI a givcn paIh W bc pursucd. I
parIicuIar, scarciIyoIrcsourccsWIavorIhccIimbingoIhiIIspinning
down a popuIaIion oncc iIs Iop has bccn rcachcd, homogcnizing iIs
composiIion, and IcrriIoriaIizing iIs idcnIiIy. scaping Irom a Incss
pcak can bc doncby gcncIic driII, IhaIis, by a scrics oIchangcsIhaI
arc ncuIraI in Incss and invisibIc Io scIccIion prcssurcs . Such an
cscapcwouIdconsIiIuIc arcIaIivc dcIcrriIoriaIizaIion.
IpopuIaIions oIdiscmbodicdrcpIicaIors Ihcrc is onIy onc param-
cIcr aIIccIing Ihc idcnIiIy oI asscmbIagcs. BuI wiIh Ihc cmcrgcncc
oIIhc gcncIic codc andIhc acquisiIionoIaminimum oIcmbodimcnI
Ihrough cncapsuIaIion wiIhin a mcmbranc, idcnIiIy couId now bc
dcncd inIwo diIIcrcnI ways. As wc sawin ChapIcr 5 Ihc bchavior
oIaIIancicnIorganismswasrigidIydcIcrmincdbyIhcirgcncs,IhaIis,
Ihcy couId onIy Icarn as a spccics ovcr many gcncraIions, so Ihc
codingparamcIcrhadaxcdhighvaIuc Ior scvcraIbiIIionycars. BuI
Ihc IcrriIoriaIizaIion paramcIcr couId changc and Icad Io diIIcrcnI
Iypcs oIbchavior. arIybacIcria IackcdIhcabiIIyIomovcandIcndcd
Io IormcoIonicsIhaIaccumuIaIcdas Iaycrs aI IhcinIcrIaccbcIwccn
waIcr and boIIom scdimcnIs. To a casuaI obscrvcr Ihcsc bacIcria
wouId hav Iookcd |usI Iikc anoIhcr scdimcnIary Iaycr, moIionIcss
andunchanginginiIs anaIomy, IhaIis, vcryIcrriIoriaIizcd.BuIcnor-
mous changcs wcrc Iaking pIacc wiIhin Ihcir mcmbrancs as Ihc
IVI
IVZ
LNA
machncry Io Iap nIo cxIcrna! gradcnIs s!ow!y cvo!vcd. SIarIng
wIh Ihc chcmca! machncry bcLnd !crmcnIaIon Ihcsc crcaIurcs
!carncd how Io Iap nIo Ihc so!ar gradcnI usng phoIosynIhcss
and howIogrcaI!y ncrcasc Ihc cncrgyIhaI cou!dbcproduccd!rom
car!icrmcIhodsusngoxygcn. Thcsccvo!uIonaryachcvcmcnIswcrc
vcrIab!c mcIabo!c dcIcrrIora!zaIons. Thc rsI mob!c prcdaIors
cmcrgcd andwcrc ab!cIobrcakaway!romabchavora!!yIcrrIora!-
zcd!!cIhanksIoIhc!acIIhaIIhcydidnoIhavcIodcvc!opIhcrown
capacIy Io drvc mcIabo!c proccsscs away !rom cqu!ibrum. Ihcy
smp!y absorbcdbacIcraasawho!cdcvc!opngasymboIcrc!aIon-
shpwIhIhcm.
Bccauscwcarcdca!ngnowwIhpopu!aIonso!organsmsIhaIarc
coup!cdIo cxIcrna!gradcnIsIhc dagrams o!Ihcsc bo!ogca! asscm-
b!agcsmusIIakcILscoup!ngnIoaccounI. AnmporIanIsngu!arIy
nIhcscpossb!Iy spaccs s Ihc carryng capacIy, Ihc crIca! dcnsIy
o!apopu!aIon IhaI can bc aI !ong-Icrm cqu!brumwIhIhc ava!-
ab!c rcsourccs. !!crcnI rcproducIvc sIraIcgcs arcava!ab!cIo uni-
cc!!u!ar organsms Io coup!c Ihcrpopu!aIon dcnsIy Io gradcnIs. a
popu!aIoncanpcrodca!!y ovcrshooIIhccarryngcapacIy!cadngIo
acrashnIsdcnsIy!o!!owcdby as!owrccovcry, oronIhcconIrary,
I caad|usIIs dcnsIyIogcnI!y convcrgc on Ihcsngu!arIy.Uncc!-
!u!ar prcdaIors Icnd Io !o!!ow Ihc !aIIcr sIraIcgy wL!c Ihcr prcy
!o!!owIhc!ormcr.maddIon,Ihcdagramo!Ihcsc asscmb!agcsmusI
nc!udc Ihc combnaIora! spaccs !orIhc gcnoIypcs and Ihc assoc-
aIcd spaccs !or Ihcr phcnoIypcs) and Ihcsc a!so cxhbI coup!ng
buI o! a d!!crcnI Iypc. Whcn Ihc gcnoIypc o! a prcy popu!aIon
changcsnawayIhaIIhcrcsu!IngphcnoIypcsarcbcIIcrab!cIoprc-
vcnI prcdaIonILscauscs changcs nIhcIncssdsIrbuIons o!Ihc
prcdaIors,makngsomco!d gcnoIypcs!cssI andacIngasancvo!u-
IonarynccnIvc Io changc. And vicc vcrsa, any gcnoIypc changc
IhaI ncrcascs prcdaIory capacIy changcs Ihc Incss dsIrbuIons n
Ihc combnaIora! spacc o! Ihc prcy acIng as a sc!ccIon prcssurc on
Ihcr popu!aIons. Thcsc arms raccs bcIwccn prcdaIors and prcy
consIIuIc a !c o! dcIcrrIora!zaIon IhaI scIs Ihcr dcnIIy as
spcccsnconInuousvaraIon.
WIh Ihc advcnI o! mu!Icc!!u!ar organsms and d!!crcnIaIcd cc!!
Iypcs Ihc codng paramcIcr bcgan Io changc as ncurons s!ow!y
wrcsIcdconIro!o!bchavor!romIhcgcncs.Asdiscusscd nChapIcr 6,
LNA
smp!c ncurona! asscmb!agcs acqurcd Ihc capacIy Io modu!aIc Ihc
nIcnsIy o! nhcrIcdbchavor IhroughhabIuaIon or scnsIzaIon.
Thcsc smp!c !carnng mcchansms arc mporIanI noI on!y bccausc
IhcycocxsIwIhmorccomp!cxoncscvcnnhumanbcngs,buIa!so
bccausc IhcyconsIIuIcIhcrsI!ormIakcnby sub|ccIvc gradcnIs. a
smp!c organsm may bc surprscd or sIarI!cd by a novc! sImu!us,
cncrgcIca!!y rcspondng Io I unI! I gcIs uscd Io I and ccascs Io
rcacI. BuI IhcrsI Iruc dcparIurc !rom a rgd gcncIc dcIcrmnaIon
o!bchavora!dcnIIywasc!assca!condIoning, a!ormo!assocaIvc
!carnng n whch nhcrIcd rc!!cx s uscd Io anchor a !carncd
rcsponsc. C!assca!condiIonngmp!csIhccxsIcncc o!novc!cnIIcs
IhaI cmcrgc !rom Ihc nIcracIons bcIwccn ncurons, Ihc sIab!c paI-
Icrns o!ncurona!acIvIy wcrc!crrcdIo as dsIrbuIcd rcprcscnIa-
Ions. Thcsc sIab!c acIvIy paIIcrns arc rcprcscnIaIo

a! bccausc
IhcrconIcnI s aproIoIypc cxIracIcd!romasamp!co! a popu!aIon
o!sm!arcnIIcsIhaIs scnscdby organsm.ThaIs, IhcrconIcnI
sbascd onIhc ob|ccIvc sIaIsIca!propcrIcs o!apopu!aIon, propcr-
IcsIhaI asscmb!agco!ncuronscanIrans!ormnIoasub|ccIvcbuI
non!ngusIccaIcgory. Bccausc Ihcsc caIcgorcs havc IhccapacIyIo
a!!ccIhowIhcorganismbchavcsIoward cnIIcsrcscmb!ng Ihosc n
Ihc!carnngsamp!c, Ihcy c!!ccIvc!y dccodc Isbchavor.
Thc cxIracIcdproIoIypcs Ihcmsc!vcs, mIum, archgh!y dcIcrrIo-
ra!izcdcnIIcs.ThcycannoIbcrcduccdIo ncurons orIhcrphysca!
connccIons bccausc IhcyarcnoI sIorcd as such. a!!IhaIs sIorcdsa
conguraIon o! connccIonsIrcngIhs na ncurona! asscmb!agc IhaI,
gvcnIhcrghIsImu!aIon, can rccrcaIcIhcorgna!paIIcrn o!acIv-
Iy. m oIhcr words, whaI s sIorcd s a scI o! propcrIcs IhaI gvc an
asscmb!agc o!ncurons Ihc capacIy Io producc Ihc proIoIypc. Whcn
Ihc ncurona!asscmb!agcsnoI acIua!!y sImu!aIcda dsIrbuIcdrcp-
rcscnIaIon cxsIs on!y as Ihc poIcnIa! producI o! an uncxcrcscd
capacIy. Thc dagram o! Ihcsc asscmb!agcs s consIIuIcd by Ihc
sIrucIurc o!Ihcpossb!Iyspacc!orncurona!paIIcms o! acIvIy. As
cxp!ancdnChapIcr7.asancurona!asscmb!agc!cams!romcxposurc
Ioasamp!co!apopu!IonImapsrc!aIonso!sm!arIynIorc!aIons
o! proxmiIy n ILs spacc o!possb!Ics. sIimu! IhaI rcscmb!c cach
oIhcr cnd up n Ihc samc rcgon o! Ihc possb!Iy spacc. Cvcn IhaI
Ihcrc!aIionso!sm!arIy nwhaIspcrccvcdarcdcIcrmncdnparI
by Ihc diagram assocaIcd wIhIhcpopu!aIion o! cnIIcs producng
IVJ
IV+
LNA
Ihc scnsory sIimuIaIon. mapping cxIcrnaI simiIarIy rcIaIions inIo
nIcaI proximiIy rcIaIons may bc sccn as an in|cmuliza|icn c]|hc
cx|crnul diajrum. I addiIion. bccausc Ihc cxIracIon oI proIoIypcs
Irom scnsory cxpcrcncc oIIcn Iakcs pIacc n Ihc conIcxI oI moIor
bchavor IhaI usI bc adcquaIc Io IhaI cxpcrcncc. Ihc diIIcrcnI
rcgons oI Ihc possibiIiIy spacc arc assocaIcd wiIh asscssmcnIs oI
sgnLcancc. Ascrbng bchavioraI sgniLcancc Io nonIingusIic caIc-
gorcs IransIorms Ihc capacIcs Io aIIccI and bc aIIccIcd oI whaI is
pcrccivcdnIoascIoIpoIcnIaIopporIunIcsandrsks. sIrcngIhcnng
Ihc dca oIIhc anmaImndasannIoIdng oIcxIcrnaIdiagrams.
I Ihc casc oI insccIs asscssmcnIs oI sgniLcancc arc madc byIhc
spcccs as I Icarns ovcr manygcncraIonsbuIn animaIsIikc birds or
mammaIs Ihc asscssmcnIs arc pcrIormcdin parI during Ihc IIcIimc
oI ndivduaI organisms. nIkc insccIs. brds and mammaIs aIrcady
posscss a morc or Icss rich sub|ccIvc IIc. a sub|ccIvIy IhaI may bc
IhoughI oIas an asscmbIagc IhaI cmcrgcswiIhn popuIaIons oI ds-
IrbuIcd rcprcscnIaIons and Ihcr assocaIcd sub|ccIivc gradcnIs
|pain. angcr. hungcr. IhirsI ) . Thc sIabiIiIy oIIhcr sub| ccIvc idcnIIy
may bc cxpIaincd as an cIIccI oIhabIuaI bchavor. habiIs IcrrIoraI-
izc bchavior. maIng ccrIain paIIcrns rcIaIivcIy uniIorm and rcpcI-
Ivc. and undcr Ihcr inucncc a rcIaIvcIy dcIcrrIoriaIzcd LcId oI
cxIracIcdproIoIypcs couId bccomc consoIidaIcd nIo a morc or Icss
cohcrcnIanimaI sub|ccI. Thc acquisIon oIhabIs. inIurn. dcpcnds
on a grcaIcr dcgrcc oI dccodng. IhaI s. on Ihc IacI IhaI brds and
mammaIs can changc Ihcr bchavior Ihrough a Iypc oI assocaIivc
Icarnng IhaI docs noI dcpcnd on inhcrIcd rccxcs. nsIrumcnIaI
condiIionng. This Ind oI Icarning can cIciI novcI bchavoraI paI-
IcrnsbypccingIogcIhcrbchaviorsIhaIoccur sponIancousIy aIIow
IrcqucnccsncrcasingIhcirraIc oIoccurrcnccIhroughsomcIormoI
rcinIorccmcnI. Thc diagram oI avan and mammaIan mnds can bc
sIudicd wiIh Ihc hcIp oI sIrucIurcd rcprcscnIaIions Iikc scrpIs IhaI
capIurc Ihc acIon possbiIiIcs oIrouInc siIuaIons. ScrpIs canhcIp
us undcrsIand how an animaIs mnd canhavc Ihc capaciIy Io makc
scnsc oI sccnc in which Ihc ob|ccIs pcrccvcd havc an cnduring
dcnIiIyandpIayroIcsIikc IhaI oIagcnI orpaIicnI.
IargcanimaIscan cxcrcisc IhcrcapacIyIorsccncanaIysisnoIonIy
Io undcrsIandhowIhcir cnvirocnI can aIIccI and bc aIIccIcd by
IhcmbuI aIso Iomakc scnsc oIIhc opporIuniIcsandrskswiIhwhich
LNA
Ihcr conspccLcs suppIy Ihcm. IparIicuIar. monkcys andapcs can
noIonIyrccognizc onc anoIhcr as ndivduaIorganisms and rcmcm-
bcr Ihc ouIcomc oI pasI nIcracIions buI aIso rcIaIc Io oIhcrs u a
sIraIcgic way IhaI dcpcnds on Ihcr posIon u a socaI hcrarchy.
Thc qucsIon wc cxpIorcd in ChapIcr 8waswcIhcr scIs oImuIuaIIy
sIabiIizng nIcracIion sIraIcgcs couId cmcrgc u primaIc commun-
Iics. sraIcgics Iikc Ihc rccprocaIion oI Iood sharing or groomng.
arIy sIudics sccmcdIo indicaIc IhaI rcciprocaI aIIrusmcouIdindccd
bccomc domnanI in an cnIirc communiIy. IhaIs. IhaIa IcrrIoraI-
zcdasscmbIagc oIsIraIcgicswiIhawcII-dLncdidcnIIycouIdcsIab-
IshIscIIaIIcrscvcraIgcncraIons.BuIbccauscIhcIaIcoIanysIraIcgy
cnds Io bc rcIaIcdIo Ihc proporIion oIanimaIs usingiI Ihc ouIcomc
maybc morc dcIcrrIoraIzcd. as soon as onc sIraIcgysIarIs Io dom
naIc Is vcryhigh Ircqucncy oI occurrcncc crcaIcs Ihc condiIions Ior
anoIhcr sIraIcgy Io sprcad which. u Iurn. may Iavor Ihc cvcnIuaI
propagaIon oI ycI anoIhcr sIracgy. Ihc cnIrc asscmbIagc changing
dcnIiIy as Is nIcaI composIion varcs conIinuousIy. This rcsuII
was obIaincdnsimuIaIonsnwhichIhc sIraIcgics wcrcIuIIy inhcr-
iIcd. IhaI s. Ihc casc wc cxpIorcd was Ihc onc wiIh a high vaIuc Ior
Ihc codng paramcIcr. BuI wc saw no rcason Io cxpccI IhaI habIs
Icarncd Ihrough rcinIorccmcnI wouId Icad Io a morc IcrrIoraIzcd
ouIcomc. Thc dagramoIIhcsc asscmbIagcs was sIudicd usngapay-
oII maIrx Io capIurc Ihc opporIuniIcs Io coopcraIc and Ihc rsks oI
bcing chcaIcd. as wcII as snguIarIics Iikc Nash cquIbra dcLnng
sIabIc scIs oIsIraIcgcs.
AsmIarapproachwas uscdIoIacIcIoIransIonIromIhcprimaIc
communiIics IhaI wcrc Ihc IasI common anccsIors oI humans and
chimpanzccs Io Ihc communiIics oIhunIcr-gaIhcrcrs IhaI characIcr-
zcdhumanIyIormuchoIiIshisIory.ThcIransIonwas modcIcdas
nvoIvinga changc Irom scIs oI sIraIcgcs uscd u dyadcnIcracIions
Io sIraIcgcs nvoIvng a rcIaIon bcIwccn a pcrson and an cnIrc
communIy. A sIabIc scI oI sIraIcgcsinIhis casc s onc IhaI rcsuIIs n
a soIidary communiIy in which coopcraIivc bchavor Ioward oIhcr
mcmbcrs dcLncs Ihc boundarics oI Ihc group. Such a IcrriIoriaIzcd
asscmbIagc s aIways IhrcaIcncdbyIrcc rdcrs. communiIymcmbcrs
whorcIuscIoparIicipaIcincoIIccIvchunIingandgaIhcrngbuIcn| oy
Ihc spoiIs whcn Ihcsc arc rcdisIribuIcd. nIikc Ihc casc oI dyadic
sIraIcgics u whichpunshingIhosc who chcaI s simpIya maIIcr oI
IVb
IV
LNA
rcIa!aIng agansI Ihcm, n Ihc mu!I-pcrson casc punshmcnI s
hardIo!ocus andcanhavc coI!ccIvc unnIcndcdconscqucnccs, Ikc
wavcs o! rcIa!aIon IhaI cndangcr so!darIy and sharcd rcsourccs.
IunshmcnI sa!socosI!ysoIhcrc arcnccnIvcs Io!cIoIhcrsdo Ihc
punshng. Tms mcansIhaI an cnIrc!y ncw habI had Io bc addcd
Io Ihc asscmb!agc Io sIab!zc Is dcnIIy. mcIanorms, Ihc habI o!
punshngIhoscIhaIdonoIpunsh chcaIcrs. A!IhoughmcIanormsn
prc-!ngusIc hunIcr-gaIhcrcr communIcs do noI ycI consIIuIc a
ncw Iypc o! codng, bcng smp!ya ncw IcrrIora!zng dcvcc, Ihcy
do prcparc Ihc ground !or Ihc vcnIua! rsc o! !ngusIc caIcgorcs
IransmIIcd !rom onc gcncraIon Io Ihc ncxI by cn!orccd soca!
ob!gaIon.Thc dagramo!Ihcsc morc comp!cxasscmb!agcso!sIraIc
gcs can a!so bc sIudcd usng payo!! maIrccs and Nash cqu!bra
buI aswc saw app!yng Ihcsc dcas IoIhc mu!I-pcrson casc s morc
comp!caIcd and!css wc!!undcrsIood.
Oncc sIab!c human communIcs cmcrgcdIhcy bccamc capab!c o!
nIcracIngwIhoIhcrcommunIcsand!orm!argcrsca!casscmb!agcs.
Somc o! Ihcsc wcrc rc!aIvc!y sIab!c and cndurng, !kc Ihc !argcr
who!cs consIIuIcdby many communIcs asscmb!cd nIo anurban
ccnIcr. OIhcrs wcrc rc!aIvc!y cphcmcra! buI ! Ihcywcrc rccurrcnI
Ihcy cou!d havc nIcrcsIng cmcrgcnI propcrIcs o! Ihcr own. m
ChapIcrwccxp!orcdaparIcu!arrccurrcnInIcr-communIynIcrac
Ion, IradngnIcracIons, Ihc cmcrgcnIpropcrIy o!wmchwasprccs.
Thcsc wcrc noI, o! coursc, prccs dcncd n moncIary Icrms buI n
cusIomaryamounIso!onc ob|ccInIcrmso!anoIhcrob|ccI.Thc rca-
son wc nccdcd Io ana!yzc Iradc was Io cxp!an Ihc !ong dsIanccs
Iravc!cdbymgh-qua!IyrawmaIcra!s!orIhcproducIonsIoncIoo!s.
Onccsma!!urbanccnIcrsspcca!zngn!ong-dsIanccIradchadcomc
nIo cxsIcncc Ihcy can cxp!an Ims phcnomcnon buI cvdcncc !or
Ihc mgh mob!Iy o! maIcra!s !kc !!nI prcdaIcs Ihc cmcrgcncc o!
Iowns. Ihcrway,nIhcabscncco!apowcr!u!IradcparIncrIhaIcan
scI prccs oro!a powcr!u! govcrnmcnIorganzaIonIhaIcanrcp!acc
IhcmwIhccnIra! ommands, an asscmb!agc o!IradngcommunIcs
and Ihc ow ! goods bcIwccn Ihcm s rc!aIvc!y dccodcd. Thc
dagram !or Iwo Iradng communIcs was sIudcd wIh Ihc hc!p o!
nd!!crcncc maps capIurngIhc co!!ccIvcprc!crcnccso!Ihc commu-
nIcs. Thcsc maps wcrc combncdnIoandgcworIhboxIo dsp!ay
Ihc rcgon o! Ihc spacc o! possb!c combnaIons o! Iradcd ob|ccIs
LNA
conIanngIhcpoIcnIa!gans !romIradc.mIhs rcgona sngu!arIy
dcncsIhc opIma! ouIcomc !orIhc Iradng nIcracIon, a sngu!arIy
rc!crrcd Io as compcIIvc cquIbrum. I! acIua! Iradc a!ways Iook
p!acc aI Ihc sngu!arIy, as cconomsIssomcImcs assumc, anasscm-
b!agc o!Iradng communIcs wou!dbcrc!aIvc!yIrIora!zcd, buI
aswcsawIhs assumpIonsnoIncccssary.acIua!IradngnIcracIons
can Iakc p!acc awa !rom cqu!brum and sI!! rca!zc somc o! Ihc
gans!romIradc.
m IhaI chapIcrI was suggcsIcd IhaI Iradc may havc prcccdcdIhc
cmcrgcncc o! !anguagc buI IhaI Icmpora! prcccdcncc s noI sIrcI!y
ncccssary. !anguagc and Iradc may havc cocvo!vcd noI on!y wIh
cach oIhcr buI wIh oIhcrpracIccs!kc Ihc ccrcmona!bura! o!Ihc
dcad. As noIcd abovc, Ihc c!!ccI o!!anguagc onhuman bchavor |as
wc!!asonIhccmcrgcnIbchavoro!communIcs andorganzaIons)
s sounqucIhaIImusIbcassgncdIs ownparamcIcrn anasscm
b!agc.wordsandscnIcnccs|usI!kc gcncshavcIhc capacIy Iocodc
dcnIIy. BuI !anguagcs and gcnomcs arc Ihcmsc!vcs asscmb!agcs
IhaI can bc sIudcdndcpcndcnI!y o! Ihc ro!c Ihcy p!ay as a codng
paramcIcr. ConIcmporary !anguagcs, !or cxamp!c, can !orma dcIcr-
rIora!zcd conInuum o! da!ccIs ora IcrrIora!zcd scI o! sIandard
!anguagcs Ihc boundarcs o! wmch arc carc!u!!y xcd Ihrough Ihc
usc o!o!ca!dcIonarcs, grammars, andru!cso!pronuncaIon.`m
ChapIcr I 0wc approachcd Ihc mcrgcncc o! !anguagc as a proccss
IhaIbcganwIhmono!Imcsymbo!c arI!acIsIhaIwcrc s!ow!yIrans
!ormcd nIo Ihc knds o! cnIIcs IhaI cou!d !orm Ihc parIs o! an
asscmb!agc. anIcscIo!wordsIhaIcou!dbc synIacIca!!ycombncd
nIo an nnIc numbcr o! possb!c scnIcnccs. Cong !rom a scI o!
mono!Imc arI!acIs wIh a rgd!y dcncd dcnIIy Io a scI o! rccom
bnab!c componcnIs Ihc dcnIIy and mcanng o!wmchdcpcnds on
whaIoIhcrwordsIhcyco-occurwIh,s Isc!!aproccss o!dcIcrrIor
a!zaIon.Thcdagramo!Ihcsc asscmb!agcswassIudcdIakngadvan-
Iagc o!Ihc !acI IhaI Ihc spacc o!a!!possb!c !anguagcs, as dcncdby
Ihcr snIax, s c!osc!y!nkcdIo Ihc spacc o!possb!c auIomaIa IhaI
canmasIcrIhaIsynIax.Thc auIomaIacanbc consdcrcdsngu!arIcs
nIhcspacc o!compuIaIona! capacIcs, somc cxcmp!!yng a mn-
mumoIhcrsamaxmumo!compuIaIona!capacIy.
Our na! Ihcmc was organzaIona! asscmb!agcs posscssng an
auIhorIy sIrucIurc. 1 was argucd IhaI Ihc rsc o! such asscmb!agcs
IV
IVd
LNA
was d!Lcu!I Io cxp!an bccausc o! Ihc acIvc mcans IhaI hunIcr-
gaIhcrcr and agrcu!Iura! communIcs had | and somc sIi!! havc) Io
prcvcnIIhccrysIa!!zaIono!auIhorIy. dsspaInggradcnIs o!sIaIus
orprcsIgcIoprcvcnI!cadcrshp !rombccongpcrmancnI, ordss-
paInggradcnIso!cncrgy |surp!us agrcu!Iura!producIs byburnng
Ihcm or rcdsIrbuIng Ihcm Io prcvcnI Ihc crcaIon o!a sIock. BuI !
Ihcsc mcchansms o! prcvcnIon wcrc ncccssary I was bccausc Ihc
dagramo!Ihcsccommuna!asscmb!agcs,Ihcrsoca!Iopo!ogy,a!rcady
conIancdccnIra! auIhorIy as a possb!Iy. Communa! asscmb!agcs
wcrc IcrrIora!zcd Io d!!crcnI dcgrccs. morc H Ihcy had scII!cd Io
pracIcc agrcu!Iurcand!css!Ihcyrcmancdmob!candrangcdovcr
awdcrIcrrIory Bynco!IhcImcsboIhposscsscd!anguagcandIhs
gavc Ihcm Ihc mcans Io codc Ihcr own dcnIIy. Whcn Ihc mcha-
nsms o!prcvcnIon!a!cdandchc!doms andarchacsIaIcs cmcrgcd
Ihcsc smp!cr communIcs wcrc noI !c!Ibchndas a prmIvc sIagc
o!dcvc!opmcnIbuIcocxsIcdwIhIhcmorccomp!cxoncs asparIo!
an cxp!oIcdpcrphcry. As such Ihcy wcrc sub|ccIcdIo an cvcrccdinj
as sIaIc organzaIons supcrmposcd Ihcr own codcs o! conducI on
Ihcm wIhou rcp!acing Ihc naIvc oncsI oIhcrwords, communa!
asscmb!agcsbccamcacomponcnIparIo!!argcrasscmb!agcsrcIanng
somc o!Ihcr own dcnIIy buI bcng !orccd Io cxcrcsc capacIcs n
Ihc scrvcc o! Ihc producIon and manIcnancc o! Ihc dcnIIy o! a
!argcrwho!c.
Thc sub| ccI o! ccnIra!zcd auIhorIy s comp!cxbccausc Invo!vcs
scvcra!!cvc!so!IhcparI-Io-who!crc!aIon.ThcrcsIhc !argcsIasscm-
b!agc nwhch a ccnIra! sIaIc occupcs a corc surroundcd by !argc
chc!doms and apcrphcry o!sma!! chc!doms and agrcu!Iura! com-
munIcs.ThcnIhcrcsIhcasscmb!agco!IhccapIa!cIyIsc!!,madcup
o!avarcIy o! communIcs and organzaIons |Icmp!cs, monumcnIs,
workshops, rcsdcnccs, and Ihc roya! pa!acc Isc!!) as wc!! as Ihc
physca! in!rasIrucIurc g a!! Ihcsc IogcIhcr. Ina!!y, Ihcrc s Ihc
indvdua!organzaIonsacIngasinsIIuIona!agcnIswIhIhcrown
goa!s and Ihc rcsourccs Io achcvc Ihosc goa!s. Thc IcrrIora!zaIon
paramcIcr can havc d!!crcnI va!ucs !or cach !cvc! o! Ihc parI-Io-
who!crc!aIonbuIngcncra!Ihcrsco!accnIra!sIaIccanbcconccvcd
asIhcproducIo!apowcr!u! dcIcrrIora!zaIon. AIIhc !argcsIsca!cI
nvo!vcd achangcn Ihc way Ihc dcnIIy o!a soca!who!c src!aIcd
IoIsphysca!IcrrIory.communa!asscmb!agcsoccupyaIcrrIoryand
LNA
Ihcpccc o! !andon whch Ihcy arc scII!cd or around whch Ihcy
roam s parI o! whaImakcs Ihcm who Ihcy arc sIaIc organzaIons
conIro!!ng a vasI IcrrIory, on Ihc oIhcr hand, dcIach Ihcmsc!vcs
!rom Ihc !and Io bc ab!c Io makc ob|ccIvc comparsons abouI Is
agrcu!Iura! producIvIy, assgnng d!!crcnI va!uc Io d!!crcnI p!oIs
!or adsIraIvcpurposcs. AIIhcsma!!csI sca!c anndvdua!orga-
nzaIoncan bc sad Io bc dcIcrrIora!zcd!Ihc rcsourccs I conIro!s
arc !nkcd Io an o!Lcc as opposcd Io Ihc ncumbcnI o! IhaI o!Lc.
TadIona! !cgImacy, whaI madc Ihc auIhorIy o! an gypIan
pharaoh ow down drccI!y !rom Ihc gods, !c!I p!cnIy o! room !or
pcrsona! caprcc so o!Lcc and ncumbcnI wcrc noI scparaIc. BuI
Ihc !cgImacy dcrvcd !rom Ihc capacIy o! an organzaIono so!vc
prob!cms o! sccurIy, !ogsIcs, monumcnIa! consIrucIon, and oIhcr
acIvIcs m whch Ihc ouIcomc dcpcnds on Ihc succcss!u!maIchng
o! mcans Io cnds, was morc mpcrsona!. Thc cocxsIcncc o!d!!crcnI
sourccs o! !cgImacy mp!cs IhaI Ihcrc wcrc Iwo po!cs dcLnng
organzaIona! asscmb!agcs n archac sIaIcs, Ihc magca! and Ihc
!cga!-raIona Thcsc Iwo po!cs can bc sccn maIcra!zcdmIhc Cza
pyramd, a monumcnI IhaI was aI oncc a rcsurrccIonmachnc and
a!caI o!sIrucIura! cngnccrng.
I ChapIcr I I wc conccnIraIcd on Ihc sccond po!c aI Ihc sma!!csI
sca!c. Ihc !aws, chano! command, Iask dcLnIons and assgnmcnIs,
and conIro! o! ncar and !ar rcsourcc gradcnIs, Ihrough whch
govcrnmcnI organzaIons so!vcd Ihc prob!cm o! pyramd bu!dng.
Wc saw IhaI Ihc dagram o! Ihcsc organzaIona! asscmb!agcs cou!d
bc sIudcd wIhIhc hc!p o!graphs o!d!!crcnIIypcs, somc Io capIurc
Ihc hcrarchca! rc!aIons machano! command oIhcrs Iodsp!ayIhc
dvson o! !abor wIhn cach o! Ihc ranks. Thc LrsI graph rcvca!s
whcIhcr, !or cxamp!c, Ihcrc arc possb!c concIs o!auIhorIy mIhc
casco!arankwhoscmcmbcrsmusIobcycommands!rommorcIhan
onc drccI boss. Thc sccond graph dsp!ays Ihc nIcrdcpcndcnccs
among Iasks IhaI spcca!zcdworkcrs musI Iakc nIo accounIasIhcy
carry ouI Ihosc commands, inIcrdcpcndcnccs IhaI mark ponIs o!
possb!c con!cI m schcdu!cs orin Ihc usc o!rcsourccs. ThcscpoIcn-
Ia!auIhorIy or pcr!ormancc con!cIs arc !!c sngu!ar cvcnIssIruc-
Iurng Ihc spacc o! possb!Ics !or Ihc asscmb!y proccss o! ganI
!uncrary monumcnIs and oIhcr !argc-sca!c govcrnmcnI pro| ccIs.
ThcyparI!yspcc!yIhcprob!cmIhaIanorganzaIonmusIso!vcmuch
IVV
Z
LNlA
as oIhcr dagrams dcnc prob!cms !oroIhcr asscmb!agcs. Ihc sIaIc
pacc o!coup!cdmo!ccu!arpopu!aIonsdcncsaprob!cm!oragrad-
cnIhcprob!cmo!whaI!!owpaIIcrnIogcncraIcIodsspaIcasmuch
cncrgv as possb!c: Ihc coup!cd spaccs !or possb!c gcnoIvpcs and
phcnoIvpcs dcncprob!cms!or cvo!vngorgansms, prob!cms so!vcd
bv scarchng Ihcsc spaccs !or so!uIons: Ihc opporIunIcs and rsks
a!!ordcdbvIhc cnvronmcnI dcncprob!cms!or!carnngorgansms
Ihc so!uIons o! whch nvo!vc Ihc dcvc!opmcnI o! assocaIons o!
sImu!, goodhabIs, orcvcnsk!!s: andIhc soca!d!cmmasgcncraIcd
bv Ihc chocc Io coopcraIc or chcaI dcnc prob!cms !or anma!s or
humansnvo!vcdnndcnIc!vrcpcaIcdsIraIcgcnIcracIons.
Our characIcrzaIo o! asscmb!agc Ihcorv nccds on!v onc morc
!caIurc Io bc comp!cIc. Thc dcnIIv o! an asscmb!agc s noI on!v
cmbodcd n Is maIcra!Iv buI a!so cxprcsscd bv I. Ths dsIncIon
corrcsponds Io IhaI bcIwccn maIIcr-cncrgv on onc hand and n!or-
maIon onIhc oIhcr, noI Ihc scmanIc n!ormaIon convcvcdbv Ihc
mcanng o! words or scnIcnccs, buI raw phvsca! paIIcrn. A !vc
gradcnI, !orcxamp!c,mavconIanIhcsamcamounIo!cncrgvasonc
IhaIhas bccomc dsspaIcdbuIIconIansmorcn!ormaIonbccausc
Iscomosngmo!ccu!csarc ordcrcdandhavcpaIIcrn. lnaddIonIo
dcnng Ihc dcgrcc o! ordcrn an asscmb!agc phvsca! n!ormaIon
can cxprcss Is dcnIIv. To usc a smp!c cxamp!c, aIoms can cxprcss
Ihcr chcmca! dcnIIv bccausc onc o! Ihcr propcrIcs |a ccrIan
dsIrbuIon o! c!ccIrons) gvcs Ihcm Ihc capacIv Io nIcracI wIh
radaIon and !cavc a unquc ngcrprnI n I. cach c!ccIron shc!!
absorbs somc wavc!cngIhs o!IhcradaIonbuI noIoIhcrs crcaIng an
n!ormaIon paIIm IhaI canbc uscdIo dcnI!v Ihc chcmca! spcccs
o! Ihc aIom. AsIrophvscsI usc IhcscpaIIcms |Ihc paIIcms IhaI Ihc
ngcrprnIs!cavcnphoIographcp!aIcs) IodcnI!vIhc componcnIs
o!!arawavsIars,buIcvcn!nooncuscdIhsn!ormaIonIhccxprcs-
svc paIIcrns wou!d sI!!bc Ihcrc. lI wasbccausc Ihcwor!dmwhch
!vngcrcaIurcs cvo!vcdwas !!cdwIhn!ormaIon-!romIhc nuIr-
cnI gradcnIs IhaI a bacIcrum can c!mb: Io Ihc odors, co!ors, and
shapcsIhaIahoncvbcccanuscIondnccIar: IoIhc s!houcIIc orgaI
paIIcrn IhaI a prcdaIor can usc Io dcnI!v Is prcv-IhaI Ihcv wcrc
ab!c IosIarI!ormngnIcrna!modc!s o!IhcrcnvronmcnI.
Wh!c Ihc dsIncIon bcIwccn Ihc maIcra! and Ihc cxprcssvc,
bcIwccn maIIcr-cncrgv and n!ormaIon, s mporIanI Io Irack Ihc
LNlA
para!!c!hsIorcs o!bodcsanmnds, Isa!sorc!cvanIhcrcbccausc
Ihc compuIcr smu!aIons dscusscdIhroughouIIhsbookarc cmcr-
gcnI who!cs composcd o! n!ormaIon cxsIng abovc Ihc compuIcr
hardwarcIhaIprovdcs Ihcr maIcra!and cncrgcIc subsIraIum. Thc
Icchno!ogv IhaI makcs smu!aIons possb!c had Io undcrgo scvcra!
Irans!ormaIons IhaI can a!sobccxp!ancdwIhnIhc !ramcworko!
asscmb!agc Ihcorv. Thc mosI mporIanI dcIcrrIora!zaIon was Ihc
Irans!ormaIon IhaI convcrIcd a spcca!-purposc Turng machnc,
rgd!vdcncdbvIsnIcma!sIaIcsandIhcsvmbo!sIuscsIowrIcon
IsmcmorvIapc,nIoa unvcrsa!Turngmachnc capab!c o!smu!aI-
ng anv spcca!-purposc onc. Ths smu!aIng capacIv dcrvcs !rom
Ihc corrcspondcncc bcIwccn auIomaIa and !anguagcs, a corrcspon-
dcnccIhaIa!!owsa spcca!-purposc auIomaIonIobc gvcnasvmbo!c
cxprcsson and bc p!accd on Ihc mcmorv Iapc o! Ihc unvcrsa!
auIomaIon. A!Ihough Ihcsc auIomaIa arc conccpIua! cnIIcs Ihcv
canbcgvcna maIcra!!ormIhaIc!car!vdsp!avs Ihc dcIcrrIora!za-
Ion nvo!vcd. whcn a spcca!-purposc Turng machnc s maIcra!!v
cmbodcd Is dcnIIv s xcd bv Is spcca!zcd Iask buI a unvcrsa!
TurngmachnchasandcnIIvIhaIsrrcjrummublc.dcncdbvwhaI-
cvcr spcca!-purposc machnc IscurrcnI!vsmu!aIng. ThusIhc rsI
dcIcrrIora!zaIon brngs Ihc mcIa!cvc! |opcraIons on daIa) nIo
drccI conIacI wIh Ihc ob|ccI !cvc! |daIa) crcaIng Ihc possb!Iv o!
usscmblujcsmudccurc]crcrurcrsundduru.Thc componcnIso!agcncIc
a!gorIhm, !orcxamp!c, canbcdvdcdnIoopcraIors-muIaIonand
crossovcr, Incss!uncIon, sc!ccIon!uncIon-anddaIa,Ihc popu!a-
Ion o! svmbo! sIrngs consIIuIng Ihc smu!aIcd chromosomcs. As
wIhanvasscmb!agc IhcsccomponcnIsmusInIcracI,IhcnIcracIon
rcducngmIhs cascIoIhcapp!caIono!anopcraIorIoIhcdaIascrv-
ngasIsnpuI.
ThcscnIcracIons,onIhcoIhcrhand,musIbcchorcographcdsncc
IhcordcrmwhchopcraIorsarcapp!cdorIhc numbcr o!ImcsIhcv
arc app!cd musI bc carc!u!!v spcccd. Ths nvo!vcs an addIona!
componcn !or Ihc asscmb!agc. conIro! sIrucIurcs !kc !oops and
branchng nsIrucIons. A!! IogcIhcr, opcraIors, conIro! sIrucIurcs,
and daIa a!!ow an auIomaIon wIh xcd dcnIIv Io bc codcd n Ihc
mcmorv Iapc o! a unvcrsa! auIomaIon Irans!ormng I nIo a pccc
o! so!Iwarc. anapp!caIon!kc awordproccssor, a sprcad shccI, ora
wcb browscr. Ths mp!cs Ihc possb!IvIhaIIhc dcnIIv o!a codcd
ZI
ZZ
LNA
auIomaIon can undcrgo changcs Ihrough dccodng. a proccss IhaI
dcpcnds on Ihc Iypc o! programmng Ianguagc uscd Io cxprcss I
symboIcaI!y. OIdprogrammng Ianguagcs |Iikc orIan. asca!. or CI
conIroI compuIaIona!proccsscs m a rgd hcrarchcaIway. amasIcr
program. cmbodyng Ihc bascdcnIIyo! an appIcaIon, ycIds con-
IroI whcncvcr I caI!s a subrouInc Io pcr!orm a parIcuIar Iask buI
Irccovcrs I Ihc momcnI Ihc subrouInc nshcs Is Ias Thc IaIIcr
may n Iurn surrcndcr conIroI Io an cvcn morc basc subprogram
buIcvcnIua!!yconIroI o!Ihc proccss movcsbackup IhchcrarchyIo
Ihc masIcr prgram. I oIhcrprogrammng Ianguagcs. rc!crrcd Io
as cb]cc|-cricn|cd. Ihcrc arc no masIcr programs or subrouIncs.
ConIroIsaIways dcccnIraIzcdas so!Iwarc ob|ccIs cncapsuIaIngascI
o!opcraIors arc caI!cdnIo acIon by paIIcs nIhc vcrydaIaIhcy
opcraIc on. Ths mpIcs IhaIIhcdcnIIyo!an appIcaIons dcncd
dynamcaI!y mnIcracIonwIhIhc daIa. H Ihc IaIIcrchangcs sowI!
wIhn !mIs Ihc appIcaIons own dcnIIy. I! dcIcrrIoraIzaIon
Irans!orms a pccc o! hardwarc nIo so!Iwarc. dccodng cIiminaIcs a
rgd masIcr program n !avor o! a popuIaIon o! auIonomous and
cxbc so!Iwarc ob|ccIs. Thcsc Iwo Irans!ormaIons provdcd Ihc
cnvronmcnInwhchIhc smuIaIons dscusscdnIhsbook couIdbc
bo and Ihrvc.
On Ihc oIhcr hand. wh!c a dcIcrrIoraIzaIon may cxpIan Ihc
cxsIcncc o!asscmbIagcs o!opcraIors anddaIaI docs noIaccounI !or
Ihc !acI IhaI Ihc bchavor o! Ihosc opcraIors as Ihcy acI on daIa can
mmc Ihcbchavoro!maIcra!proccsscs. I ChapIcr ! I was argucd
IhaI cxpIanaIon o! Ihc somorphsm bcIwccn a maIhcmaIca!
modcI and Ihc proccss I modcIs can bc gvcn H Ihc ob|ccIvc cxs-
Icncc o!Ihcdagrams o!asscmbIagcss acccpIcd.Thcunvcrsa!sngu-
IarIcssIrucIurngIhcscdagramsarcbydcIonrca!bunoIacIua!,
aIIhoghIhcycanbccomc acIuaIzcdwhcnIcndcnccsarcman!csIcd
or capacIcs cxcrcscd.Thc unvcrsaIIy o!IhcsIrucIurc o!possbIIy
spaccsmcansIhaIIwo cnIrc!yd!!crcnIproccsscs cansharcIhc samc
dagram. or IhaI hcr dagrams can ovcrIap so Ihc proccsscs sharc
somc o! Ihc amc snguIarIcs. In Ihcsc Icrms. Ihc somorphsm
bcIwccn mo

cIs and whaI Ihcy modcI can bc cxpIancd as a cc-

uc|uulizu|icn o!Ihc samc dagram. or o!d!!crcnIbuI ovcrIappng da-
grams. 'IIhaIchapIcr wc wcnI on Io argucIhaIIhc man dangcr
o! Ihs accounI s makng unvcrsaI snguIarIcs nIo IransccndcnI
LNl
cnIIcs,cnIIcscxsIngcnIrc!yndcpcndcnI!yo!IhcmaIcra!wor!
BuI Ihs poIcnIa!pI!a!!can bc avodcdby a!ways IrcaIng dagra
asmmancnIIomaIIcr. cncrgy, andn!ormaIonwh!cIhc ob|ccIvc
cxsIcncc o! dagrams may noI dcpcnd on any parIcu!ar maIcra!
cncrgcIc, orn!ormaIonamcchansm, I docs dcpcn onIhc acIua
cxsIcncc o!somcmcchansmoranoIhcr. Ihs accounIIurns ouIIo
bccorrccIIhcnIw!!ponIIoannImaIc bcIwccn onIoIogy and
cpsIcmo!ogy. AndIhc cxsIcncc o!sucha nIu,w!!consIIuIc
a powcr!u! argumcnI !or brcakng wIh Ihc onIo!ogy wc nhcrIcd
!romIhc c!assca! Crcckph!osophcrs, an onIo!ogybascdonIhc gcn-
cra!and Ihc parIcu!ar, andannccnIvc Iodcvc!opancwoncbasc
onIhcndvdua!sngu!arandIhcuvcrsa!sngu!ar.
ZJ
Z+
NO|O5
!n/I0dNt/l0n
' JohnSIuarIN!!.ASysrcmc] Lcjic.RuriccinurivcundInducrivc. |Iondon.
Iongmans, Crccn,andCo. , I06) . p. 243.
Ccorgc Hcnrv Icwcs. Prcblcms c] Lifc undMind. Vo!. 2. |Iondon.
Tbncr Co. , I 875) . p. 4I 5.
Samuc!A!cxandcr.Srucc, 1imc,undDcin.Vo!. 2. |Iondon.NacIan,
I 20) . p. 46-47. Scc a!so. C. I!ovd Norgan. Emcrjcnr Evcluricn.
|NcwYork HcnrvHo!I, I 3 I ) . p. 8.
LhuD/0I n0
' IcIcr W. AIkns. Dc Scccnd Luw. |Ncw York. SccnIc Amcrcan
Ibrarv, I 84) . p. 73.
IcIcrW. AIkns. lbd.,p. 38.
rc . Schncdcr and oron Sagan. Inrc rhc Cccl. Encr Flcws,
1hcrmcdynumics, und Li]c. | Chcago. UnvcrsIv o! Chcago Ircss,
2005 ) . p. I I 2-I 3.
Crcgorc Nco!s andlIva Irgognc. Exlcrinj Ccmrlcxin. |NcwYork.
W.H. Frccman, I 8) . p. I 2.
` JoscphN. NoranandNchac!. Norgan.Mcrccrclcjy. 1hcArmcsrhcrc
und rhc Scicncc c] Wcurhcr. |Ncw York. NacIan Iub!shng Co. ,
I 86) .p. 287.
JoscphN. Nora andNchac!. Norgan. lbd., p. 7I .
' JoscphN. oranandNchac!. Norgan. lbd.,p. 3 0I and3 I 5 .
Crcgorc Nco!s and lIva Irgognc. Exrlcrinj Ccmrlcxin. Op. CI.
p. I 8-20.
A!an Carnkc!. Fcrms c]Exlunuricn. |Ncw Havcn. Ya!c UnvcrsIv
Ircss, I 8I ) . p. 5862.
NUTb5
'

IcIcr SmIh. Exluininj Chucs. |Cambrdgc. Cambrdgc UnvcrsIv
Ircss, I 8) . p. 72.
` W!!am J. Kau!an andIarrv I. Smarr. Surcrccmrurinjundrhc
1r]crmuricn c] Scicncc. |Ncw York. SccnIc mcrcan Ibrarv,
I3) . p. I4-I 5. CrcdI !or Ihs !andmark smu!aIon s gvcn Io
KobcrI W!hc!mson. H madc Is dcbuI aI SlCCKAIH 8, and was
nomnaIcd!oranAcadcmvAward!oranmaIon.cIa!sandgraph-
cscanbc!oundnIhcwcbsIco!IhcNaIona!CcnIcr!orSupcrcom-
puIngApp!caIons.
'' W!!am J. Kau!!man andIarrvI. Smarr.lbd.,p. I 3.
' ' ugcncI.Wgncr. 1hc unrcuscnublcE]ccrivcncssc]Murhcmurio in rhc
Nururul Scicnccs. m SvmmcIrcs and KcBccIons. |Woodbrdgc.
Ox BowIrcss, I 7) . p. 222-37.
'" Norrs nc. Murhcmuricul Dcujhr]cm Ancicnr rc Mcdcm 1imcs.
Vo!. 3. |NcwYork. Ox!ordUnvcrsIv Ircss, I72) . p. 882.
` Norrs K!nc. Murhcmuricul Dcujhr]cm Ancicnr rc Mcdcm 1imcs.
Vo!. 2. lbd. ,p. 732.
'
JuncBarrow-Crccn.PcincurcundrhcDrccBcdyPrcblcm. |Irovdcncc,
V. AmcrcanNaIhcmaIca!SoccIv, I7) . p. 32-3.
'' lan SIcwarI. DccsCcdPluy Dicc.1hcMurhcmurics c]Chucs. | Ox!ord.
Bas!B!ackwc!I, I8) . p. I 07.
'

A!cxandcrWoodcoandNonIcavs. CurusrrcrhcDccty. |NcwYork.
. I. uIIon, I78) . p. 42.
LhuD/0I 0
' W!!am IoundsIonc. 1hc Rccursivc univcrsc. |Ncw York. W!!am
Norrow, I85) . p. 26-3I .
' W!!am IoundsIonc.lbd. ,p. 82.
' W!!amIoundsIonc.lbd. ,p. 38-40.
" W!!amIoundsIonc.lbd. ,p. I05.
W!!amIoundsIonc.lbd. ,p. 202.

Thc mp!cmcnIaIon o! Ihs machnc was pcr!orcd n Ihc
vcar 2000 bv Iau! Kcndc!I, and can bc sccn n opcraIon aI hs
wcbsIc.
' SIcphcnWo!!ram.Unvcrsa!IvandComp!cIvnCc!!u!arAuIomaIa.
mCcllulurAurcmuruundCcmrlcxin. |Kcadng.Addson-Wcs!cv, I 4) .
p. I40-55.
Zb
ZW
NUTb5

SIcphcn WoI!ram. o imcnsionaI CcIIu!ar AuIomaIa. H Ccllulur
AurcmuruundCcmrlcxin. lbid., p. 2I 3.
' ChrisIophcrC. Iangton.caIIhcdgco!Chaos.lnArri]ciulLi]c 1
diIcdbvCrisIophcr C.Iangton, CharIcsTavIor, J. ovncIarmcr,
and SIccnKasmusscn. |KcdwoodCiIv.Addison-WcsIcv, I 2) .p. 4.
'

CrisIophcr C. Iangton. lbid., p. 83.
' ' SIcphcn WoI!ram. A Ncw Kind c] Scicncc. | Champagnc. WoI!ram
Ncdia, 2002) . p. 845.
'
z
BrosIHassIachcr.iscrcIcIIuds.lnFrcmCurdinulsrcChucs.R#ccricns
cnrhcLifcundLcjunc]Srunisluvulum.diIcdbvNccia CranICoopcr.
|NcwYork. CambridgcUivcrsitvIrcs, I 8) . p. I 8I .
' ' Tomasso TooIi and Norman H. NargoIus. lnvcrtibIc CcI!u!ar
AuIomaIa. a Kcvicw. ln Ccllulur Aurcmuru. 1hccty und Excrimcnr.
diIcdbvHowardCuIowiIz. |Cambrdgc.NIrcss, I I ) . p. 23I .
' BrosIHassIachcr.iscrcIcIIuds. Op.CiI.p. I 88.
' SIcphcWoI!ram. Thcmodvnamics and Hvdrodvamics o! CcIIu!ar
AuIomaIa.lnCcllulurAurcmuruundCcmrlcxin. CcllccrcdEssuys. |Kcad-
ing.Addison-WcsIcv, I 4) . p.260.
'

Tsumu Shimomura, Carv . ooIcn, BrosI HassIachcr, and CasIor

Iu. CaIcu!aIions UsingIaIIicc Cas Tcchniqucs. lnFrcmCurdinuls rc
Chucs. Op. CiI.p. 202-I0.
'' TomassoTooIiandNomanH. Nargo!us.CcllulurAurcmuruMuchincs.
ANcwEnvircnmcnr]crMcdc/inj. |Cambrdgc.NIrcss, I 87) .p. I53.
'

Tomasso TooIi and Noman H. Nargo!us. lnvcrtib!c CcIIu!ar
AuIomaIa. Op. CiI. p. 244-5.
'' IcIcr CaIison. CompuIcrSimu!aIionsandIhcTading Zonc. ln 1hc
Disunin c] Scicncc. diIcd bv IcIcr CaIison and avid J. SIump.
| SIand!ord. SIand!ordUivcrsiIvIrcss, I6) . p. I 38-44.
LhuD/0I 1hI00
' KonaId . Iox.Encrjy undrhcEvcluricnc]Lifc. |NcwYork. W.H. Ircc-
man, I 88) . p. 8-.
' SIuartKau!an. 1hc 0rijinsc]0rdct. Sch0rjunizuricnundSclccricnin
Evcluricn. |

cwYork. Ox!ordUivcrsiIvIrcss, I3) .p. 20-6.

' avidrcss!crandHunngtonIoIcr.DisccvcrinjEnzmcs. |NcwYork.
ScicnIihcAmcricanLbrarv, I I ) . p. I 68-72. And. N. C. VciIchand
J.I. WiIIiams. Thc NoIccu!ar Basis o! !cctron Tans!cr in Kcdox
NUTb5
nzvmc SvsIcms. H Frcnricrs c] Bicrruns]crmuricn. Vo 7. |BcrIin.
AkadcmicVcrIag, I2) . p. 283-0.
" SIuarIKau!!man.ThcOriginso!Ordcr. Op.CiI.p. 302-3.
SIuartKau!!ma. lbid., p. 30-I0.

SIuartKau!!man.lbid., p. 320.
' KichardJ. BagIcvandJ. ovncIarmcr. SponIanco usmcrgcncco!
a NcIaboIism.lnArrihciulLi]c H. diIcdbvChrisIophcr C. IangIon,
Char!cs TavIor, ovnc Iarmcr, and SIccn Kasmusscn. |Kcdwood
CiIv.Addison-WcsIcv, I2) . p. I I2.

KichardJ. BagIcvandJ. ovncIarmcr.lbid.,p. I I8-26.
' WaIIcr IonIana. AIgoriIhmic ChcmisIrv. lnArri]ciulLt]c . Op. CiI.
p. I68.
!v
WaIIcrIonIana. lbid.,p. I7.
! !
WaIIcrIonIana. lbid.,p. I 84.
LhuD/0I 10NI
!
Nan!rcdigcn. Srcrs1cwurdsLifc. | Ox!ord. Ox!ordUnivcrsiIvIrcss,
I 2) . p. 28.
Nan!rcdigcn.lbid.,p.65-7.
' IcIcrSchusIcr. HowoKNA NoIccuIcsandViruscsxpIorcThcir
WorIds?lnCcmrlcxin.Mcrurhcrs.Mcdcls.undRculin.diIcdbvCcorgc
A. Cowan, avidIincs, avidNc!Izcr. |Kcading. Addisson-WcsIcv,
I 4) . p. 400.
Nan!rcdigcn. Srcrs1cwurdsLifc. Op. CiI.p. 2-5.
SIuartKau!!man.1hc0rijinsc]0rdcr. Sch-0rjunizuricnundSclccricnin
Evcluricn. |NcwYork. Ox!ordUnivcrsiIvIrcss, I3) . p. 34-5.

SIuartKau!!man.lbid.,p. 45-62.
' Nan!rcdigcn.SIcpsTowardsIi!c.Op. CiI.p. 28-.

Nan!rcdigcn.lbid.,p. 82-6.
' NcIanic NiIchcI! and SIcphanic IorrcsI. CcncIic AIgoriIhms and
ArtihciaI Ii!c. H Arrihciul Li]c. diIcd bv ChrisIophcr IangIon.
|Cambridgc. NIrcss, I7) . p. 272.
!v
avid . Co!dbcrg. 1hc Dcsijn c] Inncvuricn. Lcsscns ]cm und]cr
CcmrcrcnrCcncricAljcrirhms. |BosIon. KIuwcr Acadcmic IubIishcrs,
2002) . p. 3.
! !
avid . CoIdbcrg. Ccncric Aljcrirhms in Scurch. 0rrimizuricn, und
MuchincLcuminj. |Kcading. Addisson-Wcs!cv, I 8) . p. I25-30.
Z
Zd
NUTb5
'' IcIcr SchusIcr. CompIcx OpIimizaIion in anArIiciaI KNA WorId.
mArrihciulLifcII.diIcdby ChrisIophcr C. IangIon, CharIcsTayIor,
oync Farmcr, and SIccn Kasmusscn. |Kcdwood CiIy. Addison-
WcsIcy, I2) . p. 28I-3.
'' IcIcrSchusIcr. lbid.,p.287.
'" John K. Koza, ForrcsI H. BcnncII, avid Andrc, NarIin A. Kcanc.
CcncIic Irogramming. BioIogicaIIy lnspircd CompuIaIion IhaI
xhibiIs CrcaIiviIy in Iroducing Human-CompcIiIivc KcsuIIs. ln
CrcurivcEvcluricnurSysrcms. dIcdby IcIcrJ.BcnIIcyandavidW.
Cornc. |Sanicgo.AcadcmicIrcss, 2002) . p. 283-0.
'' JohnK.Koza. CcncricPrcjrumminj. 0nrhcPrcjrumminjc]Ccmrurcrs
byMcans c]NururulSclccricn. | Cambridgc.NIrcss, I2) . p. 80-8
'

JohnK.Koza.lbid.,p 32.
'' John K. Koza, ForrcsI H. BcnncII, avid Andrc, NarIin A. Kcanc.
CcncIic Irograming. BioIogicaIIy lnspicd CompuIaIion IhaI
hibiIsCrcaIiviIyinIroducingHuman- CompcIivcKcsuIIs. lnOp.
CiI. p.284.
'

Jon K. Koza, NarIin A. Kcanc, NaIIhcw J. SIrccIcr, WiIIiam
NydIowcc, Jcsscn Yu, and Cuido Ianza. Ccncric Prcjrumminj
RcurincHumun-CcmrcririvcMuchincInrcllijcncc. |Ncw York. Springcr,
2003) . p. 22-30.
'' JohnK.KozacIaI.lbid.,p. 268-7I .
'

JohnK. ozacIaI.lbid.,p. 237.
'' WaIIcrFonIana. Thc TopoIogy oIIhc IosibIc. 1cumulc]1hccrcricul
Biclcjy. VoI. 2I 3, No. 2. |Ncw York. Acadcmic Ircss, 200I ) .
p. 24I-74.
'' SIuarIKauIan. Dc 0rijins c]0rdcr. Op. CiI. p. I I 5.
'' JohnHoIIand.AdurruricninNururulund Arri]ciulSysrcms. |Cambridgc.
NIrcss, I2) . p. 68.
'" CoIin K.KccvcsandJonaIhan. Kowc. CcncricAljcrirhms. |BosIon.
KIuwcrAcadcmicIubIishcrs, 2003) . p. 65.
'' avid. CoIdbcrg. 1hcDcsijn c]Inncvuricn. Op. CiI.p. I 22-5.
'

NichacI C. Kossmann. lnIroducIory ComcnIs onIhcFuncon oI
omains in IroIcin SIrucIurc. m Inrcr|cninj Scuucnccs inEvcluricn
undDcvclcrmcnr. diIcdbydwin N. SIonc and KobcrI J. SchwarIz.
|NcwYork. OxIordUnivcrsiIyIrcss, I 0) .p. 5-7.
'' S. K. HoIIand and C. C. 1 BIakc. IroIcins, xons, and NoIccuIar
voIuIion.lnlbid. p. I I-I 3.
NUTb5
LhuD/0I 1lv0
' J. WiIIiam SchopI, J.N.Haycs, and NaIcoIm K. WaII. voIuIionoI
arIhs arIicsIcosysIcm.lnEurrh's EurlicsrBicsrhcrc. diIcdby J.
WiIIiam SchopI. |IrinccnIon. IrinccIon UnivcrsiIy Ircss, I83) .
p. 375-8I .
' FrN. Harrod. 1hcrulFcrcc.ASrudyc]Biccncrjcrio. |NcwYork.
W. H. Frccman, I86) . p. I 87.
' Ccorgc WaId. Thc Origin oI IiIc. ln 1hc Chcmicul Busis c] Lifc.
|SanFrancisco. W.H. Frccman, I73) . p. I6-I7.
" Jan Sapp. Iiving TogcIhcr. Symbiosis and CyIopIasmic lnhcriIancc.
ln Symbicsis us u Scurcc c]Evcluricnur Inncvuricn. diIcd by Iynn
NarguIis and Kcnc FcsIcr. | Cambridgc.NIrcss, I I ) . p. I 6-I7.
' John Naynard SmiIh. A arwinian Vicw oI Symbiosis. ln lbid.,
p. 30-6.

J. Van NcArIhur. Micrcbiul Ecclcjy. |AmsIcrdam. Iscvicr, 2006) .
p. I 34-47.
' NanIrcd IcschcI and Wcmcr Ncndc. 1hc Prcdurcr-Prc Mcdcl.
|Vicnna. Springcr-VcrIag, I86) . p I 2I .

SIuarI I. Iimm. 1hc Buluncc c] Nururc. | Chicago. UnivcrsiIy oI
ChicagoIrcss, I I ) . p. I 05.
' SIuarII. 1. lbid.,p. I 5-3I .
' FiIippoNcnczcrandKichardK.BcIcw. IaIcnIncrgnvironmcnIs.
mAdurrivcIndividuulsundEvclvinjPcruluricns. diIcdbyKichardK.
BcIcw and NcIanic NiIchcII. |Kcading. WcsIvicw Ircss, I6) .
p. I 4.
' ' FiIippo Ncnczcr and Kichard K. BcIcw. voIving Scnsors in nvi-
ronmcnIs oI ConIroIIcd CompIcxiIy. ln Arrihciul Lifc. diIcd by
Kodncy A. Brooks and IaIIic Nacs. |Cambridgc. NIrcs, I6) .
p. 2I5.
'
FiIippo Ncnczcr and Kichard K. BcIcw. From CompIcx nviron-
mcnIs Io CompIcxBchaviors. mAdurrivcBchuvicr. VoI. 4, No. 3l4.
| Sagc JoumaIs OnIinc, I6) . p. 3I 7-63.
' ' SIuarIKauIIman. 1hc0rijinsc]0rdct. Sc/[-0rjunizuricnundSclccricnin
Evcluricn. |NcwYork OxIordUnivcrsiIyIrcss, I3) . p. 243.
'" SIuarIKauIIman.lbid., p. 263.
' ' John H. HoIIand. Hiddcn Ordcr. HcwAdurruricn Builds Ccmrlcxin.
|KcadiDg. AddsonWcsIcy, I 5) . p. I 04-7.
ZV
ZI
NUTb5

John H. Ho!!and. Hiddcn 0rdcr. Hcw Adurruricn Builds Ccmrlcxin.
|Kcading, Addson-Wcs!cy, I5I . p. I I I-I3.
' ' IcIcr. T. Hrabcr, Tcrry Joncs, and SIcphanc IorrcsI. Thc co!ogy
o!cho. InArrihdulLifc.Vo!. 3,No. 3. | Cambrdgc.NIrcss, I7I .
p. I I-l.
'

Sorn Sonca. BacIcra!vo!uIonWIhouI SpccaIon.In Symbicsisus
uScurccc]EvcluricnurInncvuricn.Op. CI. p. l00-2.
'' JohnH.Ho!!and.HddcnOrdcr. Op. CI.p. I l5-20.
4v
JohnH.Ho!!and.Ibd.,p. l22-23.
i
NarkA. Bcdau, mi!c Snydcr, NormanH. Iackard. A C!asscaIon
o!Iong-Tcrmvo!uIonaryynamcs. InArrihdulLifc V. dIcdby
ChrsIophAdam, KchardK. Bc!cw,HroakKIano, andChar!cs,
Tay!or. |Cambrdgc. NIrcss, l8I . p. 23 l-2.
LhuD/0I Sb
' John H. Ho!!and. Hiddcn 0rdcr. Hcw Adurruricn Builds Ccmrlcxin.
|Kcading.Addson-Wcs!cy, I 5I . p. 32.
4
Kona!dJ.NacCrcgoranddwinK. Icws.NcurulMcdclinj. |NcwYork.
I!cnum Ircss, I 77I . p. 2l-30.
' John T Bonncr. 1hc Evcluricn c] Culrurc in Animuls. |IrnccIon.
IrnccIonUnvcrsIyIrcss, I80I . p. I l 3.
" JamcsI.Cou!d.Iho!ogca!andComparaIvcIcrspccIvcsonHoncy
Bcc Icamg. In Insccr Lcuminj. dIcd by anc! K. Iapa| and
A!cnda C. Icws. |NcwYork. ChapmanandHaI!, I 3 I . p. 3 l .
JamcsI . Cou!d.Ibd.,p. 3 I-8.

IzabcIh A. Bcmays. Avcrson Icamg and Iccding. In Insccr
Lcuminj. Op. CI. p. l0.
' N. J. NackinIosh. Ccndiricninj und Asscdurivc Lcuminj. |Ox!ord.
C!arcndonIrcss, l83I . p. l 8-2.

N. J. NacknIosh.Ibd.,p. 52-5.
' Wi!!amBcchIc!andAdc!cAbrahamscn. CcnnccricnismundrhcMind.
AnImrcducricnrc PurullclDisrriburcdPrcccssinjinN c/wcrk. |Camrdgc.
Basi!B!ackwc!!, I l I . p. l06-7.
' Kar!Sms. vo!vng 3 Noho!ogyandBchavor by CompcIIon.
In ArrihdulLi]c . dIcd by Kodncy A. Brooks and IaIIc Nacs.
| Cambrdgc.NIrcss, l 4I. p. 33.
NUTb5
I
Jamcs J. Cbson. 1hcEcclcjiculArrrcuchrcvisuulPcrccrricn. |BosIon.
HoughIonN!!!n, I 7I . p. I 5-I6.
J
Thc anmaIon, ca!!cd vo!vcdVrIua! CrcaIurcs,maybcwaIchcd
aIIhc auIhors wcbsIc.
' avd.Kumc!harIandJamcsI.NcC!cI!and.IINodc!sandCcn-
cra! Issucs m Cognvc Sccncc. m Purullcl Disrriburcd Prcccssinj.
Exlcruricns in rhcMicrcsrrucrurcc]Ccjniricn. Vo!. . dIcd by avd
Kumc!harIandJamcsNcC!cI!and. |Cambrdgc.NIrcss, I86I .
'" avd . Kumc!harI, Cco!!rcy . HnIon, and Kona!d J . Wi!!ams.
LcuminjInrcmulRcrrcscnruricnsbyErrcrPrcrujuricn.InIbd., p. J22.
'' Nci! Ccrshcn!c!d. 1hcNururc c]MurhcmuriculMcdclinj. |Cambrdgc.
UnvcrsIyIrcss, Cambrdgc, UK, lI . p. l57-
'

Wi!!amBcchIc!andAdc!cAbrahamscn. CcnnccricnismundrhcMind.
Op. CI. p. 32.
'' Iau!1 N.J. Vcrschurc andAnIhony C. C. Coo!cn.AdapIvc Ic!ds.
sIrbuIcdKcprcscnIaIonso!C!assca!!yCondIoncdAssocaIons.
In Nc/wcrk.Ccmrururicn in Ncurul Sysrcms. Vo!. 2, No. 2. |In!orma
Hca!Ihcarc, OnIncJourna!, ll I . p. l8-206.
'
Cco!!rcy HnIon, Jamcs NcC!c!!and, and avd . Kumc!harI.
sIrbuIcdKcprcscnIaIons. InPurullclDisrriburcdProcssinj. Op. CI.
p. 86-7.And.1VanCc!dcr.cningsIrbuIcdKcprcscnIaIon.
In Ccnnccricn Sdcncc. Vo!. 4, No. 3. | Ox!ordshrc. Car!ax, l2I .
p. l76-87.
'' Wi!!amBcchIc!andAdc!cAbrahamscn. CcnnccricnismundrhcMind.
Op. CI. p. l26-8.
' I!ppoNcnczcrandKchardK.Bc!cw.IaIcnIncrgynvronmcnIs.
InAdurrivcIndividuulsundEvclvinjPcruluricns. dIcdbyKchard K.
Bc!cwandNc!ancNIchc!!. |Kcadng.WcsIvcwIrcss, l6I. p. I6
i
JamcsI.Cou!d.Iho!ogca!andComparaIvcIcrspccIvcsonHoncy
Bcc Icamg. mInsccrLcuminj. Op. CI. p. l.
Chcp/cr Scvcn
anc! I. SchacIcr and ndc!T!vng. WhaI Arc Ihc NcmorySys-
Icmsm I 4?InMcmcrSysrcms 1 VV4. dIcdbyanc! I. SchacIcr
andndc!T!vng. |Cambrdgc.NIrcss, l4I .p.28.
J
anc! I. SchacIcrandndc!T!vng. Ibd., p. .
ZII
ZIZ
NUTb5
>
SIcphcn WaIkcr. Animul 1hcujhr. |Iondon. KouIIcdgc and Kcgan
IauI, I985 ) . p. 243.
4
Howard ichcnbaum. 1hc Hirrccumrul Sysrcm und Dcclururivc
Mcmcr in Humuns und Animuls. In Mcmcr Sysrcms. I994. Op. CiI.
p. I 52-5.
SIcphcnWaIkcr.Animul1hcujhr. Op.CiI.p. 257-8.

IauI N. ChurchIand. 1hc Enjinc c] Rcuscn, rhc Scur c] rhc Scul.
|Cambridgc.NIrcss, I 996) . p. 40-5.
7
IauIN.ChurchIand.Ibid., p. 4 7-5 I .

crck BIackman. 0rcrunr Ccndiricninj. AnExcrimcnrul Anulysis c]
Bchuvicr. |Iondon. NcIhucn, I 974) .p. 48.

crckBIackman. 0rcrunrCcndiricninj.Ibid.,p. 35and97.
!v
avidC.Naycrs. ExlcrinjPschclcjy. |NcwYork.WorIhIubIishcrs,
2005) . p. 345.
l \
A. Harry KIop!, Jamcs S. Norgan, and ScoII . Wcavcr. NodcIing
Ncrvous SysIcm IuncIionwiIhaHicrarchicaINcIworko! ConIroI
SysIcms IhaI Icarn. In Frcm Animuls rc Animurs 2. diIcd by
JcanArcady Ncycr, HcrbcrI I. KoiIbIaI, and SIcwarI W. WiIson.
| Cambridgc.N Ircss, I 993) . p. 255.
!
Icdcrico Cccconi and omcnico Iarisi. NcuraI NcIworks wiIh
NoIivaIionaIUniIs.InFrcmAnimulsrcAnimurs2. Ibid., p. 3467.
!>
SIcphcnWaIkcr.Animul1hcujhr. Op. CiI.p. 35-64.
!4
Kogcr C. Schank and KobcrI I. AbcIson. Scrirrs. Pluns, Cculs, und
undcrsrundinj. |HiIIsidc.Iawrcncc arIbaum, I 977). p. I 8.
!>
KogcrC. SchankandKobcrII.AbcIson. Ibid.,p.42.
!
KisIo NiikkuIaincn. Subsmbclic Nururul Lunjuujc Prcccssinj. An
ImcjrurcdArrrcuch rc Scrirrs, Lcxiccn. undMcmcr. |Cambridgc. N
Ircss, I993) . p. 5-6.
!7
Jc!!rcyI. Iman.Findinj Srrucrurc in1imc. Ccjnirivc Sdcncc. VoI. I4.
|Norwood. AbIcx IubIishing, I 990) . p. I 79-2I I .
!
Tcuvo Kohoncn. Sc/f-0rjunizinj Murs. |BcrIn. Springcr, 200I ) .
p. I 06-I 9.
!
BcmdIriIzkc. CrowingScI!- OrgamzingNcIworks.InKchcncnMurs.
diIcdbykki O|a andSamucIKaski. |AmsIcrdam.Iscvicr, I 999).
p. I 3I3.
v
KsIoikuIaincn. SubsymbclicNururulLunjuujcProcssinj. Op. CiI.
p. I 9-29.
!
KisIo NikkuIaincn. Ibid. ,p. I245.
NUTb5
22
IhIip . Agrc. CompuIaIionaIKcscarchonInIcracIionand Agcncy.
InCcmrururicnul1hccricsc]InrcrucricnundAjcnn. diIcdbyIhiIipA.
Agrcand SIanIcyJ.Koscnscn. |Cambridgc.AIrcss, I 6). p. I2.

' Yvcs Icspcrancc and HccIorJ. Icvcsguc. IndcxicaIKnowIcdgc and

KoboI Acon.InCcmrururicnulDccricsc] InrcrucricnundAjcnn.Ibid.,
p. 443.
" Iokcndra ShasIri. Advanccs n ShruIi. A NcuraIIy NoIivaIcd
NodcIo!KcIaIionaIKnowIcdgcKcprcscnIaIionandKapidIn!crcncc
Usng TcmporaI Synchrony. In Arrlicd Inrcllijcncc. VoI. I I, No. I .
|AmsIcrdam. SpringcrNcIhcrIands, I) . p. 7I08.
' IaoIa BaIdasari, IaoIoIuIiIi, AnnaNonIcsanIo, andCuido Tascini.
ScI!-Organizing Naps Vcrsus Crowing NcuraI Cascs n a KoboIic
AppIicaon. m CcmrururicnulMcrhcds in Ncurul Mcdclinj. VoI. 2.
diIcd by Josc Nira and Josc AIvarcz. |BcrIn. Springcr-VcrIag,
2003) . p. 20I3.

NcIson Coodman. Scvcn SIricIurcs on SimiIariIy. In Prcblcm und
Prc]ccrs. |IndianapoIis. obbs-NcrrII, I 72) . p. 445.
Chcpcr Ltgh|
` IransB. N. cWaaI. SocaI SynIax.ThcI!Thcn SIrucIurc o!SocaI
IrobIcm SoIving. InAnimulScdulCcmrlcxin. diIcdbyIrans B. N.
c WaaIandIcIcr I.ack. | Cambridgc. HarvardUnvcrsiIyIrcss,
2003) . p. 237-46.
2
uancuiaII.IanguagcandvoIuIioninIhc NiddIc SIoncAgc.In
Lunjuujc Evcluricn. diIcd by Cabor Cyori. |Irank!urI. cIcrIang,
200I ) . p. 2-34.
' SamucI S. KomoriIa andCraig . Iarks. ScdulDilcmmus. |BouIdcr.
WcsIcrvicwIrcss, I 6) . p. I2-I 3.
" KobcrI AxcIrod. Thc voIuIion o! CoopcraIion. |NcwYork. Basic
Books, I 84) .p. 5.
KobcrIAxcIrod.Ibid.,p. 42.

Susan . KcchcrI. In Cumc 1hccty und Animul Bchuvicr. Ic A.
ugaIkn andHudsonK. Kccvc. | Ox!ord. Ox!ordUnvcrsiIyIrcss,
I8) . p.77.
' N.KciIhChcnandNarcHauscr.NodcIngKcprocaIionandCoop-
craIionnIrimaIcs.vidcncc !oraIunishing SIraIcgy. In1cumulc]
1hccrcriculBiclcjy. VoI. 235. |AmsIcrdam. Iscvicr, 2005) . p. 6-7.
ZIJ
ZI+
NUTb5

Sarah1 BrosnanandFransB. dcWaaI. AProxmaIcPcrspccIivcon
KcciprocaI AIIruism. In Humun Nururc. VoI. I 3, No. l . |NcwYork.
WaIIcrdc CruyIcr, 2002) . p. I48.
' KobcrI AxcIrod. 1hCcmrlcxinc]Cccrcru/icn. |PrnccIon. PrnccIon
UnvcrsiIy Prcss. I 7) . p. I72I .
' " KobcrI AxcIrod. Thc voIuIiono!CoopcraIion. Op. CiI. p. 54.
ll
MarInA.Nowak.EvcluricnutyDnumio. | Cambridc.BcIknapPrcss.
2006) . p. 86-0.
' KcnBnmorc. Cumc1hcctyundrhcScciulCcnrrucr. | Cambridc. N
Prcss, I 8). p. 3 I4-2I .
l >
Joshua M. psIcn and Koss Hammond. In Joshua M. psIcn,
CcncrurivcScciulScicncc. |PrnccIon.PrnccIonUnivcrsiIyPrcss,2006) .
ChapIcr 3.
' KcnBinmorc. Cumc1hcctyundrhcScciulCcnrrucr. Op. CiI.p. 3 I.
l J
Brian Skyrms. Thc ynamics o!KaIionaIcIibcraIion. | Cambridc.
HarvardUnivcrsiIyPrcss, I 0) . ChapIcr6.
l
KcnBinmorc. Cumc1hcctyundrhcScciulCcnrrucr. Op. CiI. p. 403.
l7
avid W. SIcphcns and KcvD C. CIcmcnIs. Camc Thcory and
Ica. InCamcThcoryandAnmaI Bchavior. Op. CiI. p. 244.
l
MichacI Macy. NaIuraI ScIccIion and SociaI Ic B Prisoncrs
iIcmma. m SocioIoicaI McIhods and Kcscarch. VoI. 25, No. l .
|SacPubIicaIions, OnIncJoumaI, I6) . p. I 22-4.
lV
MarInA. Nowak.EvcluricnutyDnumio. Op. CiI. p. 7.

MichacI Macy. NaIuraI ScIccIion and SociaI Ican n Prisoncrs
iIcmma. Op. CiI. p. I 25.
' KcnBnmorc. Cumc1hcctyundrhcScciulCcnrrucr. Op. CiI. p. 24-6.

MarIn A. Nowak. EvcluricnutyDnumics. Op. CiI. p. I45-5.

John W. PcppcrandBarbara B. SmuIs. Thc voIuIion o!Coopcra-
Iionn an coIoicaI ConIcxI. an AcnI-BascdModcI. In Dnumics
in Humun und Primurc Sccicrics. diIcd by TmoIhy P KohIcr and
Ccorc J. Cumcrman. | Ox!ord. Ox!ord UnivcrsiIy Prcss, I 8) .
p.63-4.
omcnico Parisi, Fcdcrico Cccconi, and AnIonio Ccrni. Kin-
irccIcdAIIruismandAIIachmcnIBchaviornanvoIvinPopuIa-
Iion o! NcuraI NcIs. m Arrihciul Sccicrics. 1hc Ccmrurcr Simuluricn
c]ScciulD]c. diIcd by NicI CiIbcrI and Kosaria ConIc. |Iondon.
UnvcrsiIyCoIIcc o!IondonPrcss, I 5) . p. 238-48.
NUTb5
' ChrisIphHaucrI. CoopcraIion. CoIIccIivcsFormaIion, and SpcciaI-
zaIion. In Advunccs in Ccmrlcx S,srcms. VoI. , No. 4. |Snaporc.
WorIdScicnIicPubIishD, 2006) . p.3 I6.
Thomas C. SchcIIn. Micrcmc/ivcs und Mucrcbchuvicr. |Ncw York.
W.W. NorIon, I 78) .p. I I0-4.
' KobcrI AcIrod. 1hcCcmrlcxinc]Cccrcruricn. Op. CiI. p. 52.

KobcrIAcIrod.Ibid., p. 52-5.

" TomohisaYamashiIa, KoshiIzumi,andKoiiKurumaIani. !!ccI

o!MuIuaI Choicc McIanormn Croupynamics !or SoIvn SociaI
iIcmmas. m Acnr-Buscd Simulu/icn. Frcm Mcdclinj Mcrhcdclcjics
rc Rcul Wcrld Arrlicuricns. diIcd by Takao Tcrano, Hamc KiIa,
Toshiuk Kancda, Kiyoshi Arai, and Hiroshi cuchi. |Tokyo.
Sprncr-VcrIa. 2005) . p. 38-46.
Kcn Bnmorc. Cumc 1hccty und rhc Scciul Ccnrrucr. Op. CiI.
p. 344-58.
Chcpcr tnc
' Mary. Icakcy. CuIIuraI PaIIcHIhc OIduvai Scqucncc. InA]cr
rhc Ausrrulcrirhccincs. Srrurijrurhy. Ecclcjy. undCulrurc Chunjc in rhc
Middlc Plcisrcccnc. diIcd by KarI W. BuIzcr and CIynn lsaac. |Thc
Hauc. MouIonPubIishcrs, I 75) . p. 477.

S. A. Scmcnov. PrcHsrcric 1cchnclcjy. |ToIowa. Bamcs and NobIc

Books. I85) . p. 3-55.
CIivc CambIc.xchanc, Foran,andIocaIHomnidNcIworks.m
7rudcund Exchunjc in Prchisrcric Eurcrc. diIcdby Chris Scarrc and
FrancisHcaIy. | Ox!ord. OxbowBooks, I 3) . p. 36.
ChrisScarrc.InIroducIion.InTudcundExchunjcinPrchisrcricEurcrc.
bid., p. I .
' MarshaII Sah!ns. Srcnc Ajc Eccncmio. |Chicao. AIdncAIhcrIon,
I 72). p. 277-80.

MarshaII SahIns.bid.,p. I0I2.
' HaroIdI. ibbIc and NicoIas KoIIard. OnAsscmbIac VariabiIiIy n
Ihc MiddIc PaIcoIiIhic o! WcsIcm uropc. m 1hcMiddlcPulcclirhic.
Adurruricn. Bchuvict, und vuriubilin. diIcd by HaroId I. ibbIc
and PauI Mc!Iars. |PhiIadcIphia. UnivcrsiIy Muscum, UnivcrsiIy o!
PcnnsyIvana. I2) . p. I -I I .
ZIb
ZI
NUTb5

Iawrcncc Cuy SIraus. vcn Ihc NoIon o! TansIona!IdusIry
Is a SuspccI Typo!ogca! ConsIrucI. m NmArrrcuchcs rc rhc Srudy
c]urrcrPulcclirhic DunsiricnulIndusrrics"in WcsrcmEurusiu. dIcd
byJu!cnKc!-Sa!vaIorc andCco!!rcyA. C!ark. | Ox!ord.Bar, 2007) .
p. I I-I 5.
' Iconard Joy. Onc conomisIs Vcw o! Ihc Kc!aIon BcIwccn co-
nomicsandAnIhropo!ogy.m1hcmcsinEccncmicAnrhrcrclc.diIcd
byKaymondFrIh. |Iondon.KouI!cdgc, I6 7) . p. 34.
i
" Pau!WonnacoIIandKona!dWonnacoII.Eccncmics. |NcwYork.John
W!cy Sons, I 0) . p. 33-8.
'' Thomas N. Humphrcy. Thc a!y HsIory o! Ihc Box agram.
|Fcdcrul Rcscr|c Bunk c] Richmcnd Eccncmic 0uurrcrly. Vo!. 82l I ,
WnIcr I 6) .p. 37-5I .
i
Icgh Tcs!aIson. AgcnI-Bascd CompuIaIona! onomcs. A Con-
sIrucIvc Approach Io conomic Thcory. m Hundbcck c]Ccmruru-
ricnulEccncmics. Vo!. 2. |AmsIcrdam. !scvcr, 2006) . p. 845-50.
'' an K. Codc, SIcphcn . Spcar and Shyam Sundcr. Ccnvcrjcncc c]
DcublcAucricnsOPurcrc 0rrimulAllccuricnsinrhcEdjcwcrrhBcx. |Ya!c
Schoo!o!NanagcmcnI, workngpapcr, 2004) .
'" Jc!!rcy S. can, Ccorgc J.Cumcrman,JoshuaN. psIcn, KobcrII.
AxIc!!, A!an C. Swcd!und, N!cs T. Parkcr,and SIcphcnNcCarro!!.
UndcrsIandng Anasaz Cu!Iurc Changc Through AgcnI-Bascd
Nodc!ng.mJoshuaN.psIcn. CcncmrivcScdulSdcncc. |PnccIon.
PnccIon UnivcrsIy Prcss, 2006) . p. . Joshua N. psIcn and
KobcrI AxIc!!. Crcwinj Arrmdul Scdcrics. |Cambrdgc. NIT Prcss,
I 6) . p. 23-6.
'' Joshua N. psIcnandKobcrIAxIc!!.Ibd., p. 30-2.
'

JoshuaN. psIcnandKobcrIAxIc!!.Ibd.,p. I 06.
'' JoshuaN. psIcnandKobcrIAxIc!!.Ibd., p. I 0-I I .
'

JoshuaN. psIcnandKobcrIAxIc!!. Ibd.,p. I 20.
' ' SIcvcnNIhcn. Smu!aIngPrchisIorcHunIcr-CaIhcrcrSoccIcs.m
SimulurinjScdcrics.diIcdbyNgc!C!bcrIandJmoran. |Iondon.
UCI Prcss, I 4) . p. I 8I-3.
2
Jc!!rcy S. ca cI a!. UndcrsIandng Anasaz Cu!Iurc Changc
ThroughAgcnI-BascdNodc!ng. Op. CI. p. 8.

i
SIcvcn NIhcn. Smu!aIng PrchisIoc HunIcr-CaIhcrcr SoccIcs.
Op. CI. p. I 7.
22
SIcvcnNIhcn.Ibd., p. I 6-70.
NUTb5
'' NarkWnIcr Iakc. NACICAI CompuIcr Smu!aIon o! Ncso!Ihc
Foragng. m Dnumics in Humun und Primurc Scdcrics. Ajcnr-Buscd
Mcdclinjc]Scdul und SruriulPrcccsscs. diIcd by TmoIhy A. Koh!cr
andCcorgcJ. Cumcrman. | Ox!ord. Ox!ordUnvcrsIyPrcss, 2000) .
p. I I 56.
'" NarkWinIcrIc.Ibd. ,p. I I8.
' NarkWnIcrIc. Ibd., p. I 30.
'

NarkWnIcr Ic. Ibd., p. I 37-8.
'' Cramc C!ark. WcrldPrchisrcty. |Cambrdgc. CambrdgcUnvcrsIy
Prcss, I 77) . p. 64.

Jc!!rcy S. can cI a!. UndcrsIandng Anasaz Cu!Iurc Changc
ThroughAgcnI-BascdNodc!ng. Op. CI. p. 4-I 00.
'' Jc!!rcy S. cancIa!. Ibd. ,p. 103.
'

Jc!!rcy S. cancIa!.Ibd.,p. I 06-7.
Chcp/cr Tcn
' Nark . Hauscr and W. Tccumsch FIch. Whur Arc rhc uniuucly
Humun Ccmrcncnrsc] rhc Lunjuujc Fuculnm Ianguagc vo!uIon.
diIcdbyNorIcnH. ChrisIanscnandSmonKrby. | Ox!ord.Ox!ord
UnivcrsIyPrcss, 2003) . p. I 6I-8.
' Sona Kagr. 1cwurds un undcrsrundinj c] rhc Rcluricnshir bcrwccn
Bircdul Wulkinj, Enccrhulizuricn, und Lunjuujc 0rijins. m Ianguagc
vo!uIon. diIcd by Cabor Cyry. |Frank!urI. PcIcr Iang, 200I ) .
p. 845.
' KchardY. Kan. Aurcmuru 1hccty. |Ncw York.Ncgraw-H!!, I 72) .
p. 84-0 |Turng machincs), I 22--4 |!ncar-boundcd auIomaIa),
I42--4 |push-downauIomaIa) .
" Iuc SIcc!s and Frcdcrc Kap!an. BooIsIrappng Croundcd Word
ScmanIcs. mLinjuisricEvcluricnrhrcujhLunjuujcAcuuisiricn.diIcd
by Tcd Brscoc. |Cambrdgc. Cambrdgc UnvcrsIy Prcss, 2002) .
p. 56-7.
' IucSIcc!sandFrcdcrcKap!an. Ibd., p. 5.

IucSIcc!sandFrcdcrcKap!an. Ibd., p. 7I .
' dwn HuIchns and Bran Haz!chursI. Lcuminj in rhc Culrurul
Prcccss.mArIca!I!cH. diIcdbyChrsIophcr C.IangIon, Char!cs
Tay!or, oync Farmcr, and SIccn Kasmusscn. |Kcdwood CIy.
Addison-Wcs!cy, I 2) .p. 63.
ZI
NUTb5

dwinHuIchnsandBrIanHazIchursI.Ibd., p. 67-8.
' dwInHuIchinsandBrIanHazIchursI.Ibd.,p. 702-4.
'

Noam Chomsky. Asrccrs c]rhc Thccr c]Synra. |CambrIdgc. N
Ircss, I 65) . p. 66-73.
'' NaIaIa I. Komarova and NarIn Nowak. Lunjuujc, Lcuminj. und
Evcluricn. mIanguagcvoIuIon. dIcdbyNorIcnH. ChrIsIanscn
and SimonKrby. Op. CI. p. 326-8.
' ' SIcvcnInkcr.LunjuujcusunAdurruricnrcrhcCcjnirivcNichc.mIbd.,
p. 25-8.
'' SmonKrby. Icamg,BoIIIcnccks,andIhcvoIuIono!Kccursvc
SynIax. In LinjuisricEvcluricn rhrcujh LunjuujcAcuuiricn. Op. CI.
p. I 77-8.
'" SImonKIrby.Ibd. ,p. I 84.
'' SImonKrby. Ibd., p. I 2.
'

Jamcs K. Hur!ord. Ercssicn/InducricnMcdcls c]Lunjuujc Evcluricn.
Dimcnsicns und Issucs. In Linjuisric Evcluricn rhrcujh Lunjuujc
Acuuisiricn. Op. CI. p. 306-I I .
'' SImonKrby. Icamg, BoIIIcnccks, andIhcvoIuIono!Kccursvc
SynIax. Op.CI. p. I 8.
'
J. NarkBaIdwIn. ANcwIacIorInvoIuIon.mAdurrivcIndividuuls
in EvclwPcruluricns. Mcdcls und Aljcnrhms. dIcd by Kchard K.
BcIcwandNcIacNIchcII. |KcadIng.WcsIvcwIrcss, I 6) .p. 64.
'' NorIcnH. ChrsIanscn andNckChaIck. InIcNodcIs o!InnIc
Ianguagc. A ConnccIonsI Approach IoKccurson. In Ccnnccricmsr
Psyhclinjuisrio. dIcd byNorIcnH. ChrsIanscnandNckChaIck.
|csIporI. AbIcx Iub!ishing, 200I ) . p. I 47.
'

Kchard K. BcIcw, John Nncmcy, and NcoI N. SchraudoIph.
voIvIngNcIworks. UsIngIhc CcncIcAIgorIIhmwIh ConnccIon-
sIIcamg. mArrihciulLifcH Op. CI. p. 527-30.
'' Cco!!rcy . HinIon and SIcvcn J. NowIan. How Icang Can

Cudc voIuIon. In Adurrivc Indiwduuls in Evclvinj Pcruluricns.

Op. Ct. p. 45I .

ZId
'' Zc!Iig HarrIs. A Thccr c]Lunjuujc und In]cmuricn.A Murhcmuricul
Arrrcuch. | Ox!ord. CIarcndon Ircss, I 8I ) . p. 367.
'' Zc!IgHarrIs.Ibd. ,p. 33-7.
'" Zc!Iig HarrIs. IbI,p. 3 3.
'' Jamcs K. Hur!ord. Exrcssicn/InducricnMcdcls c]LunjuujcEvcluricn.
Op. CI. p. 320-I .
NUTb5
Chcp|cr Llcvcn
' NaxWcbcr.ThcThccrc]ScciulundEccncmic0rjunizuricn. |NcwYork.
IrccIrcss, I 64) .p. I 24-32.
' KrisIanKrisIanscn. Chc!doms, SIaIcs, andSysIcmso!SocaISIraI-
caIon.InChic]dcms:Pcwct:Eccnmy,undIdcclcjy.dIcdbyTmoIhy
arIc. |Ncw York. CambridgcUnvcrsIyIrcss, I I) . p. 24.
' KaIhIccnN.CarIcyandIccCasscr. Ccmrururicnul0rjunizuricnThccr.
InNuIIagcnI SysIcms. dIcdbyCcrhardWcss. |Cambridgc.N
Ircss, 200I ) . p.300.
" NchacIWooIdrdgc. InIcIIgcnIAgcnIs. Ibd., p. 54-6.
' NchacIWooIdrdgc.InIcI!gcnIAgcnIs.Ibd.,p. 8.

ancIC.cnncI.ucBc!cvcrs.InThcInrcnricnulSruncc. |Cmbridgc.
NIrcss, I 0) . p. I 5-22.
' Jim . oran. a|cIorics Io CompIcxIy in ArIcaI SoccIcs.
In Dnumio in Humun undPrimurc Sccicrics. Ajcnr-BuscdMcdclinj c]
ScciulundSruriulPrcccsscs. dIcdb TmoIhyA. KohIcrand Ccorgc
J. Cumcrman. | Ox!ord.Ox!ordUnvcrsIyIrcss, 2000). p. 4-5.

MchacI N. Huhns and Iarry N. SIcphcns. Mulriujcnr Sysrcms und
Sccicricsc]Acnrs.InNuIIagcnI SysIcms. Op.CI.p. 85-5.
' Jim . oran and Nkc Ia!mcr. Thc OS Iro|ccI. InIcgraIing
oNcIso!IaIco!IhcSocaI Chngc.InArrt]ciulSccicrics.ThcCcm-
rurcrSimuluricnc ]ScciulLifc.dIcdbyNgcICiIbcrIandKosara ConIc.
|Iondon.UnvcrsIyCo!Icgc o!IondonIrcss, I 5) .p. I 06-I I .
1
Jim. oran andMikcIa!mcr. Ibd.,p. I I 3-20.
'' Icrrc CIasIrcs. SccicnAjuinsr rhc Srurc. |Ncw York. Zonc Books,
I 87) . p. 30.
'' NaxWcbcr. Thc Thccrc]ScculundEccncmic 0rjunizurtcn. Op. CI.
p. 35-66.
'' Crahamc CIark. WorId IrchsIory. | Cambridgc. Cambridgc Unvcr-
sIyIrcss, I 77) . p. I 05.
'" !zabcth N. BrumcI and TmoIhy K. arIc. InIroducIon. In
Srcciulizuricn, Exchunjc, undCcmrlcxSccicrics. dIcdby !zabcIh N.
BrumcIdandTmoIhyK. ac. | Cambrdgc. CambrdgcUnvcrsIy
Ircss, I87) . p. 3-4.
'' CaIhy A. SmaII. Thc Io!IcaI ImpacI o! Narragc n a rIuaI
IoIyncsan SoccIy. In Dnumio in Humun und Primurc Sccicrics. Op.
CI. p. 228-3 I .
ZIV
ZZ
NUTb5
'
e
CaIhy A. Sma!I. Ibid., p. 234-.
'' Barbara Iricc. IrrigaIion. Sociopo!iIicaI ynamics and Ihc CrowIh
o! Civi!izaIion. In ThcAsiuricMcdcc]Prcducricn. diIcdby Annc N
Bai!c and Joscp K. LIobcra. |London. KouIIcdgc Kcgan IauI,
I 8I ) . p. 2I 8.
'

AIcxandcrBadawy.AHisIoryo!gypIianArchiIccIurc.VoI. l. |Ciza.
SIuoNisr, I 54) . p. 4.
'' KichardNcIsonandSidncyWinIcr.AnEvcluricnurThccrc]Eccncmic
Chunjc. | Cambridgc.Bc!knap, I 82) . p. 8-I00.
^
KcvinCrowsIon.voIvingNovcI OrganizaIionaIForms.H Ccmruru-
ricnul0rjunizuricn1hccr.diIcdbyKaIhIccnN.CarIcandNichacI
J. IricIuIa. |Hi!!sday. LawrcnccrIbaum, I 4).p. 27-.
^

TmKovacs.oVicwso!CIassicrSysIcms.InAdvunccsinLcuminj
Clussihcr Sysrcms. diIcd by Iicr Luca Lanzi, WoI!gang SIoIzmann,
SIcwarIW.Ison. |Bcr!in. Springcr-VcrIag, 2002) . p. 77.
'' Larry BuII andTm Kovacs. InIroducIion. H Fcunduricnsc]Clussihcr
Sysrcms. diIcd by rry BuII and Tm Kovacs. |Bcr!in. Springcr-
VcrIag, 2005) . p. 8.
'` Zahi Hawass. Mcunruins c] rhc Phuruchs. |Ncw Yor. oubIcday,
2006) . p. 7.
'" NarkLchncr. Thc CcmrlcrcPrumids. |London. ThamcsandHudson,
I 7) . p. 203-I 0.
'' NarkLchncr. Ibid., p. 224-5.
'
e
KciIh S. cckcr.TasknvironmcnICcnIcrcd SimuIaIion. InSimulur-
inj0rjunizuricns. IcdbyNichacIJ.IricIuIa, KaIhIccnN. CarIcy,
andLcsCasscr. |Cbridgc.NIrcss, I 8) . p. I 08.
'' C.I. 1 Van crBoom. ThcDuricsc]rhczicr. |London. KcgaIauI,
I 88) . p. 3 I0.
'

C.I. 1 Vancr Boom.Ibid., p. 3 I 820.
'' ZahiHawass.Mcunruinsc]rhcPhuruchs.Op. CiI.p. I 60.
`

avidKrackhardI. CraphThcorcIicaIimcnsionso!In!ormaI Orga-
nizaIions.InCcmrururicnul0rjunizuricnThccr. p. CiI. p. I-5.
`' KciIhS. cckcrandVcIorLcsscr. csigningaFamiIy o!Coordina-
Iion AIgoriIhms.H Rcudinjs inAjcnrs. diIcdby NichacI N. Huhns
andNnindarI. Singh. |SanFrancisco. NorganKauann, I8) .
p. 453.
` KciIh S. cckcr. Task nvirnmcnI CcnIcrcd SimuIaIion. Op. CiI.
p. I07.
NUTb5
'' Yoav ShohamandNoshcTcnncnhoIIz. OnSociaILaws!orArIiciaI
AgcnI SocicIics. O!!-Linccsign. In CcmrururicnulThccricsc] Imcruc-
ricnundAjcnn.diIcdbyIhi!ipA. Agrc and SIanIcy J.Koscnschcin.
|Cbridgc.NIrcss, I 6). p. 58-603.
'" NichacI N. Huhns and Larry N. SIcphcns. NuIIiagcnI SysIcms
and SocicIicso!AgcnIs.InMulriujcnrSysrcms. Op. CiI.p. 87.
'' KciIh S. cckcrandVicIorLcsscr. csigning aFamiIy o!Coordina-
IionAIgoriIhms. Op. CiI. p. 455.
`

ToshiharuSugawaraandVicIorLcsscr.LcarningIoImprovc Coordi-
naIcdAcIions BCoopcraIivcisIribuIcdIrobIcm-SoIvingnviron-
mcnIs. In Muchinc Lcuminj JJ. |AmsIcrdam. KIuwcr Acadcmic
IubIishcrs, I8) . p. I 36.
`' Toshiharu SugawaraandVicIorLcsscr. Ibid., p. I 33.
'

IaIIic Nacs and Kobyn Kozicrok. Lcamg InIcr!acc AgcnIs. In
Irocccdings o! P 3 Con!crcncc. | ScaIIIc. P Ircss, I 3) .
p. 45-65. N. S. NiIIcrandK.. rcxIcr.NarkcIsandCompuIaIion.
Agoric Opcn SysIcms. In Thc Ecclcjy c] Ccmrururicn. diIcd by
Bcmardo Hubcrman. |AmsIcrdam.NorIh-HoIIand, I 88) . p. I 50.
Appcn4b
' NanucIcLanda.ANcwIhiIosophyo!SocicIy.AsscmbIagcThcory
and SociaI CompIcxiIy. |London. ConIinuum, 2007), p. 8-I I .
' NanucIcLanda. Ibid.,p. 26-32.
' NanucIcLanda. Ibid., p. I 3, 50, 58,74,0, 8, I 0I-2.
" NanucIcLanda. Ibid., p. I 5, 5,62,75,I .
' NanucIcLanda.AThousandYcarso! Non!incar HisIory |NcwYork.
Zonc Books, I 7), ChapIcr 3.

Ci!IcscIcuzcandFcIixCuaIIari.AThousandIIaIcaus|Ninncapo!is.
UnivcrsiIy o!NinncsoIaIrcss, I 87),p. 43I .w
' Ci!IcscIcuzc andFcIixCuaIIari.Ibid., p. 427-8 and435.

Ci!IcscIcuzcandFcIixCuaIIari. Ibid.,p. 44I .
' Ci!IcscIcuzcandFcIixCuaIIari. Ibid.,p. 424-6.
l
NanucIcLanda.InIcnsivcScicnccandVirIuaIIhiIosophy. |London.
ConIinuum, 2007),p. I 47-8.
ZZI

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Z
dD0 SCIIDIS Z~, V, V9
SclOI2dDI2ID2 DdDS 9~,

IIdIDID2 O 3, , VZ, ,
93, V~Z, 3
IWOldVcIc0
(DcICcDIIODS| 93, ,
Z 3
ODDOIIUDIIIcS dD0 IISKS , ,
V, V9, VV, , , V,
Z, Z, Z, V9
DdIlS dD0 WHOlcS , , V, Z, ,
Z3,93,9V3, 3,, 9,
3V, ~, V
DOSSIDlIIV SDdCcS
CODIIDUOUS 3, ZV, 9Z, 9
lNL
COUDlc0 , 9, VZ
0ISCIcIc, CODDIDdIOIdl 3,
Z~V,9, 3,,9, 9V,
33
DdDDD2 DcIWccD V~, 3
DcSIc0 9V
ODcD-cD0c0 ,9Z
OVcIldDDID2 V, 9, 9, ,
ZZ
ScdICH O Z, 9, 3, 3, ~,
, , Z, 3, 9
SIIUCIUIc O 3, ~Z, ZV,
, , , 9, ,
V,,V,VV,ZZ
SUDjcCIIVc, IDIcISUDjcClIVc,
dD0 ODjcCIIVc ~9, ,
Z9, Z,
DIODlcD-DOSID2, DIODlcD
SOlVID2 ~9, 393, 33V,
Z~,,,~9,V,
, V, V, , ~Z,
Z3, 9, 3Z, 393, ,
9, , V~Z, VV~Z
IcCUISIVc, IcCUISIOD 3~,
93, 3~, 9, 39, 33,
3V,
SI2DICdDCc, IclcVdDCc
ID cXDldD8IIODS 9, V, Z
ID DcICcDIIOD, DcDOIV ,
, V9, , , , V9
SIDUldIIODS
dD0 cDDO0IDcDI 3, 9, ,
Z, 3, , V, V, ,
, V, 3, 3, 3,
3
dD0 DdIHcDdIICdl DO0clS 3,
Z9,,,V
ZZ

smulatOns ( C0nl1)
and spatal tclatOns
(ptOXmty. stuatcdncss) 45.
4. 70-1. 87. 91. 109. 121-2.
135. 151. 153. 157
stan ntctactOns n 21. 44.
117 120 137 150 155
and tcmpOtal scalcs 139. 141.
143. 171. 174-5
ttackn ptOccsscs n 34.
42-3. 53. 70. 73. 11. 118.
125. 127. 138. 5
snulattcs
cOmpcttVc
cqulbtum 134-5. 197
cttcal thtcshOlds.
ttanstOns 53. 1 90
lObal and lOcal Optma 50~3.
1. 74
lmtn pattcns.
statcs 29-30. 8
maXma and mnma 2 5.
Nash cqulbtum 113. 118.
120. 12-7. 195-
tOpOlOcal atttactOts.
nVatants 18-20. 42-3.
4. 70
sttatcc ntctactOns amOn
nOn-human ptmatcs 111.
115-1
amOn humans 112
mathcmatcal mOdcls
O 113-15. 118-19
tcciptOcal alttusm n 112.
11. 117. 121. 123
publc OOd dlcmmas 12 3-
sOcal dlcmmas 113-14.
123
tcndcnccs 4. 8. 12. 17. 20. 23.
2. 30. 3. 49. 54. 70. 123.
125. 129. 19. 185