Approaches to the SrudyofBehavior

Behaviorcanbc dctued rs drewayar arimrl responds to srimuli in its enviromrcnt.A stimulus might bc assimpleasdetection ofthe presencc of f<x)din the enlironmcnt.In rhis sense, a bacterial cell "behaves" llv movingtowardhighcr conce,rcm tions of, sugarin its surrouDding mednun.This lxhavior is verv simplernd is well suitcdto rhe life ofbactcria, alloving theseorganisms to tiveand reproduce. \" rnir|lrl,erohrd.rhr1 uccupied diFercnr.r\iron,nenrs problcmsthar affected rnd ficed diverse thcir surrivalrDd reproduction. Their ne*ous slstems andbehavnrr corconitandy becrDre more complex. Ncnrus sr,stems percive and process information and trigger adxptivenrocorresponses, rvhichIe seeaspatterns of behavior that i tinct and experience both phy significant roles,olicn interactiry in complr war,sto producc drc final beh;rrior.

Innate bebavior does not require learning

Earlvresearch in the 6eld ofanimalbehrvn,rfncused on behaviord patterns that areah-avs exhibited h mcnlrcrs ofr species ir rcsponsc to a particrlar stirnuluqthat is, thel'appered to be instinctivc, or innate, behaviors. Bccausebehrvior is olicn stereog?ed(appenring in the sameway in differentindnidu, alsof r qrecics), theseearlyresearchers arg"ued thrr ir nrst be b$ed on prcsctpaths in rhe nenoussystcn.In their !.ie$,thcse pathsarestructured liolr genetic blucprintsand cause animals to showcssentinlly the samebchavnrr fron the first time it G produced throughouttheir livcs. Theseresearchers based tieiropinions on bchavn,rs such ascgta retieval bygeese. Cecscincubate their eggs in a nest.lf a gooscnoticesthrt rn egghasbeenknockedout of thc nest, it will cxtendits Decktowardthc cgg,get up, and roll the egg backinto the nestvith a side-to-side morior ofits ncck vhile thc cgg is nrckedbeneath its l,ill (figure 5+.1).Evcn i{the egg is rcmovedduring retrieval, the goose cornpletcs the behar-ior, rs if driven b1'a program releasedby dre initirl sight of the egg Egg retrieral behrvnr is triggered bl, i sign stimulus, the rppcarance ofrneggoutoftie nest.A componertofthe gooset ncFous svstem,the innatc rcleasirg mechanism, providesthe neural inscructionsfor the motor progrrn, or ixed action pattem, involvedir eggrcnieul. \'lore generally,the sign srimulus is r sigu.l in the cnvironDrent that triggcn a behavior. The irnarerelersing mechanisnl is the sensorlmechanisn rhatdetects the signal, mcl the tucd actionp:rftemisthc stcreorvped act. One interestingaspectof sign srimuli is th:t $ey lrc often not !ry specific;in sornesimations, a vide varietv of rill nggr r rrrrd p,ncrn. acrion f,,r errrrrple. g,*. ',brrcr. $-ill attenpt to roll basebrlls andevenllcer cansbacLilto their nesrs. Moreovcr,once rhe objectsarc in rhe nest,thc g<rxe recogDizes thrt thevare not eggs andremoves thernl A similar cramp)eis providedby nrle sticUebicLfish. During the brccdnrgseason, maiesdcvcl<,p bright red col<,r ation on their undcrsides. Teritorirl malesreactaggrcssirclv to the rpprorch ofolher nales, perfi,rnringm rggressivc rlis

Behaviot's two components are its immediate qauseand its eyolutionary origin
Itrt crn talk aboutanimil beh:rvn,r in nvo diflerclt ways. Iirst, rvcminht ask/r-o ir all vorls-that is, how the animal! senses, ncivc Detworks, or internalstatcprovidea physiolosical b:rsis for thc behavior. the "how" of behavioris a questionabort praximat ,trilti|n. To anih'zc rhe proimdte cause of behavio1 se nighr lreasLrre hormonelevelsor record the inpulse activiF of neurons in the animal.).or onrnple, a malesongbird nlay snlg during dre breedingscason because of an increased lere1of thc steroidsexhormonc testosterone, rvhichbinds to receptors in the br:rinrnd triggtrs the prodnctionofsongi rhis erphmtion r.ould describcthe pro:rimate causeof the male br ,1,*'ng. llrZr a behalior evolved an investigation ofits adaptilc value is a question conccming Lbj,Mte u!L!.!tian.To studv thc ultimatecause of I behavior, r'e attemptto dcternine horv it inllucnc.d the ani'n 'ssunnd or reproductivc success. Arnrle bird siDqs to defendr territorv FrD orhermrlcs and to actract a lcmilc with \drich to reproducei this is the ultinraer or evol:Li,,rrr. <rpLn:riortbr rhc,,,d1(\ !o, Jli/J i,,n. The sudy of behar nrrhas had alonghistorvofconnoveny. Onc sourceof controvenyhasbeen the quesrion of whethcr behrvnlr is dercrninedinorc bj.an indilidurlls genes or bl its lellrningxndexperieDce.In othcrs.ords, is bchariorthe resultof mturc (nlsnnct) or nurnrre(cxpdence)? In thc past,rhisquestion wasconsidered an eithcr/orproposition, rrurrr now roon

ssn

**l
Jigure\4.1
TNNATtr r.GG-RoLIINGRtrsloNsF rN .;F tssE. Thc series ofnryorents rscd bv r goose ro retncvcN eggls r iixedact,onprnern. ()ncc It detects (in thh case, rhc signsdlDulus an eggou6 c ihe !esr),the sooscsocsdrough the ortirc scrofnovelnenrs:Irrilleltenditsneck td!d the egg,ecr nP, dnd n)llthc ceg backinto thc ncsrvith r slde-to-sidc morion ofirs neckwhilc the eggn tuckerlbcncafi ns bill. .,,i h.ht,,i1116 t.nVIII-"1"y

othef IN sTIcKlEBAcr FISH-I{ale sticllebrclslre lery teffitoridl and .ggrcssir-ely defen.lthei. ter.it ries dgdinst SIGNSTIMULUS (in shich hehavn)r. ln l,bo.rtorv studies malcs.Itnitori,l nales havered belLiesi rs r result,red is the sign stnnllns ihat eLicits aggrcssive incswhc.hcrthe rn,le .eact\rggressivelv ti sh arc usually presenterl onlr one nodel dt d rirre),the presence of . red stripedeterm

plal andcrer attacking. When researcher Niko 'linbergen ob a malc sticlleback nr a laboratory aqurrium displaling sened p.rssed when r red maii truck by the rnrdor', he aggcssivcly rcalizcdthrt the red coloratiol r'.rs the sign stinulus (fignre 5,+.2). Subsequent enperirnents rerealedthrt m es rrculd re spondto nany unlishtike nodels rs long m the modelshad a redstripc.Tinbergen the 1971Nobel Prizein nedicineor '{rcn $ort n,,1,'St lo fi. fl,\ This phenomcnonis taLcnone stepfurther bv ruhatare termed supernormal stimuli. Civtn a choice betweent$o sisD stinuli, onc of normal size and the other rnuch larger, manl animrls will respondto the larger of the nvo. 'l'hus, givcn a choice of a normal gooseegg and one the size of a volleyball,a goosewill atterrrpt to roll the bigger one backto

stimuliexistis not alw|vsclexr. One as\\hy supernormal pectto keepin miDd,horc'cr is that in man), cases, supemornay prefereggs rnalsrinruli do not occurin mturc. Thus, geese but they never encountereggsof that fie size of volleyballs, haveevolved to respond to the hrger size.Ttmay bc thrt gecse object so that the,vnill rncnd to eggs,rather than to sm.rller, rnayhavefarrcred circuiarrocks.As a result,naturaiselection n,r hrgcr objccts. This general $e evohrrion of a preference rn.r-v leadto urexpected outcomcs in crpcdntcrts,but response probrblldoe+i ofr.rrl.. l ,,' nJlrJatli!{ bcha' I
imate behaviors tbat Edly research in nitul betiyiot mdEized in the nNoB systm md thus .rc the 6ult of preset patls,. are [rely to be smticlly.tntbled.

BehavioralGenetics
sith thosecomponents The studyofbehrvioral genetics de:tls of bchavior that arehereditq' governed bv genes andprssed h'om onc genention to the nerlt.A wide vrriety of drtl, liom artificialselecflon experiments to modernmoleculargenetics, indicrtcsthat behruioraldifferences among individuals often rcsulti orn genetic differences. Sone ratsquicklylerrncdto zip throughthe mazeto the food, nraking fev incorrectturns, but othcr ratstooknuch longerto learnrhe correctpadr. Researchers llred thc fast lcarncrswith one anotherto establish a "maze-lrright"colony,end the do* lerrners with a "mrzc-dull" colony.Oft:pring in one anorherto establish eachcolonvwere then testedto scchow quicLlythey learned the maze.'Ihe offspring of nlaze-bright rats lcarncd cven [rore quicklv than their parentshad, and thc offspring of maze-dull parentswere even poorer at mazc lcarning. Repeating this selection for several generationslcd to no behaviorallydisrincr qpes of rat s'ith vei diffcrcnt mazelearning abilities(fisure s4.3).
,b"tt* 5+ b,t-'i--t bi"t"st lll7

Rats can be artificially selected for learning capacity

A famouseraerimcrt in tie 1940sstudiedthe rbilitl' of rats of rmny blird al|o find their way through r rnazeconsisting Ieysand only one cxit, wherer rewardof fbod rwrited them.
{L v w r u r c n Dl o '.3 f.o r

fgare 54.3

I I
| ..r;iisl!dE8r.**..
-njil li -r:r..3a.ia;:::;: .': :

I
|

THE GENETICS or LEAR\ING. The f.srest rlis (rhose dr.r na.le rhe fewest errort in the p.rent.l poprl.tior lvereusedto estdblish one population (qr.ar, rnd the slo"esfudts to establjsh the other ('"/). ln subsequnt generadoN, the sane procedure wdsusedto selecr for fast(q,"d) drd sLo\'.&4 uts.

wbat uoatd happet 6 attu tbe lde"th gcneruno\ ratr ute ra".lmlr ^igherl hates reganles! oftheir abilitr to kdn 9 39 84 114 214

(Fou eenTrials) Toral Number ot E(orsln Negol alinglhe Maze

Clerrh, thc xbilit! ro lcxrn the Dr.rzewas to so[re degrcc hcrcdita4,: Furthcrmore, those genes appeared ro lre specific to this behevior, becausethe nvo groups of rets dicl not difTer in their abiliq to perfonn other behrviorel trsLs, such as mrning a completeln different kind of rnaze. This rcscarch dcmonstratcs horv a study can releal that behavior has a hedtrble componcnt.

Human twin studies reveal similarities independent of environment

The role of genetics crn also be seennr humrns by compar ing the behavn)r ofidentical trl'ins.ldenrc twi11s..rrc, rs thcir nx c inptics, genetically irlenrical, and most setsof iclcnticxl twins arc reisedin the sameenrironnrent, so it is nor possiblc to determiDe vhether similarities in behalior resultliom their geneticsn laritl or fronr environmental expericnccs shared as they gren up (the classic Drture versus-rurturc dcbate). Lr honerer, $ins hrve beenseprratcdar birrh and somecases, raised in differentfarnilies. A rccert studv of 50 such setsof twins rcvealcdmany similaritics in personalig', temperirnent, endevcnlcisuretinre actilities,e1-en thoughthe rs'inslud often l)ccnraiscdin verv. .,,,Jkarr Inr !<nel JrfrrrenL en$rorn,enr".I he.e.ir,.lbfl,i, ics pl.r-rs r role in determiningbehxvbr evennr hunans, al,l-orreh therehr:\e rrI:oflarce.l gfnfri., \ etru.er\ ironr enr is still hodv debated.

Some behaviors appear to be controlled by a single gene

just described The nrzeJearning and identicelt$-in' srudies phy e rolc nr behavior, suggest that genes bur recentresearch hrs provided much greatcrdetajlon .he generic brsrs ofbehavior. In both Dfor,p,ft/ andmicc,manvnrurarions h..l!.e beenas.oc.rred prn ruhr hrhrr.o rlrl,n,rrrulrrie,. "irh rrrt8 !'*\'lII-"1,,rr,,,,t t,h,',,,

In iuit flies,for cximplc, nldividurlsdret possess rlrer nativeallclcstur a singlc gcnc dif{er grearlvin fieir feedillg bchrvior aslanac: Lrme vith one allelemor.e around.r grert dcal asthq'e:rt, whereas indiridu.als widr e altermtne allele norc hardl,v at all. .{ r'itle varieq of nrutatjons at other genes arc rorv klom n Droiop,/, drat affecrxlDrosr elery rspectof courtship bchaYior Thc rays nr r.hich qeneticdifferences affect beharior havc irccn Rlrkcd out for sereralmousegenes. lor eurnple, somemicewith onemuta*rn havetroubleremernbering informatnrnlerrnedtso da,vs eirlier aboulwhereobjecrs erclocetcd. This dilTerence appears to resultbec.rrse the mutmt rlicc ilr not producethe en4me a crlcJumcrluodulin-dcpcndcrtki nase II, which p1at5 an irnportantrolein the function F ofrhc irippocampus, e p,rrtof rhe brrir importantfbr spNtiLrl lei ing (iescribedir chapter +6) \Iodern moleculr biology techniques alli,r. the role of genetics h behariorto be investigated \rith crcr greaterpre cision.lbr erllnrple, male mice geneticallv cngineered to lack rherbilrrv o.rnrhe.i,errirrIunJe. |.'H,n n{, r',rrts,,,,1er. shownrcreased.rggressive behrrior. A prrticularlvliscinatingbre throughoccurred nr I996, vhen scientists discorered r ne$ genc,lrj4 thxt seems to de terminewhetherfemele nicc nunurc theirl)ung in parricuhr wavs.Femdeswith bothrjB a elesdisxbledinili:rllt'irrestigatetheb newborr b|bics,bur then ignorethem, in sr.rrk contrrst to the crring rnd protcctivcnatemal behariordisphyed by nonnrl ftnules (iigxre 5.1.,1). 'l'he cause of this inattentivcncss appcars ro result fro[r r chrin rerctior. \ en mothcn ofncv tabiesinitiJly inspect then, inlbnmtion from thcil auditory olf.;rctort',ard t.rctile senses is tnnsmittc.l ro rhe hpothahnns, where.lbsa le1es are rctivetcd. The rjB allelesprcxluce.r particularprotein, which in tnm xctivatcs other enz\rnes ind genes that alfectthe '1 hesemodifications neuralcircuitlv \rithin the h'J'othrlamus. within the brdn crusethe firnale ni react nrrtern.rllvtovard

this reaction is stopped midway. No protein is actil?ted, tie brain's neural circuitry is not rewired, and matern behavior ofthe geneticbasisofbeexample Anorher fascinating two species ofNorth Americanrodents:thc havior concerns

fer jn their social behavior:Male and female prairie voles form nonogarnouspdr bonds and work together to raise volesmate and go their separnontane their young,whereas thesespecies havebeeDintenThe diftirencesbetween of the sivelystudied.The act of mating leadsto the release neuropeptides vasopressinand orl.tocin in both vole species (as well as in rnany other nanmal species).How th voles respond to the releaseof thesc peptides,however,diffcrs drarnaticalll Injection of either of thesepeptidesinto p rofmating rie volesleadsto pair bondingevenin the rbsence Convenely, injccting a chemical that blocls the action of these prairievoles notto form pairbondsafter causes neuopeptids mating. By contrast, montane voles ,re unnffbctedby either of thesemanipulations. havebcentracedto interspecific These differelt responses The prdrie volehas in brxin structtre (figure54.5). differences many receptors for these pepcidesin a particular pan of the which seenx to be involved in brain, dre nucleus accurnbens, the expressionof pair-bonding behavior.By conuast, few such

Igure )+.+
GENETICALIJ CAUSED DEIECT IN MATERNAL CARE. 4 In

hice, noinal mothers t'Le very good we of their offspring, drieving thcm if $y move dwayand ciouching over them. ,. Mothers wirh lhe mut ntt6B allele perforD ncither of c Anowt of tine thesebehaviors, leavingdleir pdpsexposed. $er c ot r --J .,,,u ,\i n g m a n u Ai n gp o ' rur( ftm,le I nic e over offspring. L Proportion of pups letiieved when they were

Igure )+. )
CENE1'IC BASIS OI DII]IiERENCDS IN PAIR-BONDINC

114! doestbe hek of tosB stleles lead to Mtmal

BEHAVIoR rN Two RoDENT sPEcIEs,a, drd r. The prairie lMidot6 orhtugttzr) and m.ntdne (rU. zo,ta,6) voles differ in the distiibntiotrof one type .f vasopresinreccptorin the with dle prairie-volc brain...'l-ransgedcmicc created ve$ion of the receptorgelcs respondto injectionsoi levelsof pair-bondins by erhibiting hcightened vasopressbn to a behaviorin 5 nin ffials conparedwith their response nonal witd t1,pe rniccshowno conrol injecion. By contrast,

rcceptors occurin the s ne brain regionin the llrontme vole. In laborato4 expe;nrentswith prririe voles,blocLingthese rr'(t'Lorsrend,r,, prelenrF,FLonding.uherers.rimutarirg them leads ro purhondJng bchav,or. Recently,, the genetic basis lbr these diflcrenceshm bcen uncovered. Scientistsha1-e identified the geDe that codcsfor the peptide receptorsrnd have discolercd that a diftcrenceexistsin the DNA structure be$-eenthe species. To test the hypothesisr.hrt this genetic differencewas responsiblefor the diflerencesin behavior,scientistscreared trxnsgenicmice rrith the prxirie volerersion ofthe gene,rnd

sureenoLgh,when injected$-ith rasopressin, thc rrarsgenic nrice exlibited pair-bondingbehaviorverv snnilar to thrt of prairie voles,whereas normal mice shoved no response (sce fi$lre 5,+.5).
The genetic bdis ofbeh.vior is supponed by ariff.ial selection experimna, studies ofidoti.'t teins, ,nd studies d the bharior of gletic hurmB. Reent ddelol,hents in noteol,r biolosy hrve led to rte discdery of specific an3 th.t control bhflioi

-....@r;;;;';;
Man)' of dle behxviorrlpattemsdisplarcdby animalsare not solclvthe result ofinstinct. In mmy cases, :nimals alter their behavioras r result of previousexperiences, a process rermed learning. 'l he role of learning tirst studiedintcnsivelv in '.$ iabofitory rodenrs, bur no.i,! rcsearchers in1srigatc the learningprocesses rnd crpabilities of a ride rangeoforganisms. ,)rcL through the association.'This form oflearning is more complex thrn h..rbituation.The t$-o major qes of associativc leirning are classicai conditionnrgrnd operantconditionhg; they differ in the nay the associations arc cstablished. Chssical conditioning In classical conditioning, the paircd presentatioDof two dif fcrent kindsofsrimuli causes the animatto tbr[ a bctr.eenthe stimuli. Classical conditioDing is alsocdlgd pavlovian condirionins, iker Rxssian ps)'ch;log;.t han i'""Io.,, who first described itPrvlov preseutedmeat po\\(lLr, ai unnndnioned ninal r. to a dog m.l notedthat the dog responded by salivrting, an ?llltunditionelftsl)oLre. It n vn related stinrulls, such asthe rnrghg o{a bell,rcpeatedly waspresented at the same time asthe powder, -eat the dogwouldsoonsaLivate in response to the sound of the bell alonc.'l he dog had lermcd to :rssociate the unrelrted soundstimulus with thc n*rt powdersrimllus. Its response to gas, thereforc,conditioncd, the soundstimulus andthc sounrr of the bell is referred to asa nulnioned ltimut6. Operant conditioning Ilr operant conditioning, an animalleamsro associate its be hrvioralresponse wjdr a rewardor prnishment. Imeric..rn psy chologist B. l: Slirner studied operantconditioning in rrti by plrcnrgthen in rn xpparanrs that cameto be crlled a ,,Skinncr box." As rhe rNt cxploredthc box, it vould occasirmally prex r levcr bv accidcrt,carxinga pellet of food to appear.It finr, the ratwould igrore the lever,eatthef<,od pellet,ind continue to m,r< rbour.soo,,.ho$e\er.i, ,(Jrnedro r*,,ci. re prr* ing the lever(thc behavioral response) obtrining food(the '.ith resrrJl.\\}en i, $J. hungr.). ,, uo rhJ rpe.rd ,rttir- r,rne pre* ine,h, lever. Thi,.on ofrrial ,rnd-emur tern-irL,,. of,nJjur importance to most r.ertebrates. Comprrativepsl,chologists uscdto belicvethxt llny t1,o stimuli could be linled in clrssical conditioning.and that anirnalscould bc conditionedto perfom my lcrrnabtebchavior ir response to an1'stimdus by opcnnt conditioning.As vou

flabituation occurs when organisms respond lessto a stimulus over time

T)re ,.mp[.r Nrreof h aring .lu'. nor re,]uire rn rnrrrt ro l,'nr rn r,{( r. ion brr$c(n|$o.rinUli o- In.."< and a response.One fotrl of an^laci/ttire ttataing is l.abiar ation, which can be dcfinedasa decrease in resp peated stimuhrs that hasno positi\.e or ncgatrve consequerces. In n an)',,+. rhe{imulu. erol,e' r.rrorrU re.por\ uhen:r ;. trr\ren.ounrcred. bUrrh( r'rdgnirude of rher(,pons gradurlh declines with repeated cxposure. As onc example,young birds see rmni qpes of objects n,uvingourh d. \r finr. rhel Inr) re,f,,,,dbr rr"uching ,lo$n:tndk,,u,ning,Dll.sorrc,,t rtreot,iccr.. .", t ,, t Ui"! le:ve,or_ merrrben uf rneiro" n .p<cie, fl1rlg b1.rn senreri lrequen' lJ Ind hr\ e r. po\ir;, e,,r legrLi\c(o \equ, n(e ro rh, nesdings. Over tirr e, the voung birds rnav hrbituate to such stnluli andstoprespondnrg. Thus, habituation canbe thought ofas learningnot to respold to a stimulus. tseingable to ignore udrnportant stinn i is criticrl for an rnnnd confrontinga brrrge of stinuli in i complexenvironnenti animalsthat could not do so rvoLrld fail co lbcus their rttcntion on inportmt activities, suchasEndingfoodand aroiding predators, and probablv *rcull leavelen. offspring nr the nextgeneration.

Associative learning links stimulus with response

A changch beha\,ior that invoh.es al associatron stimulior between a stimulus anda response is termedassociative learning (figlrc 5.t.6). Thc behavbr. is modited, or tanti

ll2o

?ot\IfI".,kst

.,,t tthari0.

roro$1ng o'scusslon! m$ !'es nas cnangeo. T ar',rcsearchers think that instinct gxideslearning bv detemining shat tvpe of infomration can be learnedthrough conditn,ning. Instinct and learnhtg ft is no$-clearthnt someanimalshi'e innrte predispositions For exrmple,if a rrt is to{ard forming certainxssocixtions. offereda food pellct rt thc samctimc it is cr?oscdto X-rap (rtich later produccnausca), thc rat rcmembers the tasteof the foodpelletbutnotits sizc,rnd in thc tutures'ill avoidfood that taste,trut will rcarlily cat pclletsol the samesize if 'idr r r."- h ...- . , t , t f - . - . r hi, .
food !ith colors, Similarh', pigeoff car lerrn to associatc dangcr r.ith but not vith smnds.In contrast, thev can associxtc sounds,but not with colo1lE. These examplcs of lcrrdng prcparedness demonstrate that s'hat aD animal can learn is biologically i.fluenced that is, leaming is possible only vithin the boundaries set by in stiDct. Innate prograns hivc cvolved becausether underscore r Jrl u ,e r 1' r r r . . ll n a ru n . i " ,J -h J , .,,,\,, r,, I r, ,, liLcl,v to hrvc a perticular taste; thus, it is rdaptire to be able to ass)ciatc a taste s ith i feeling of sicLncssthat may develop hours latcr. Thc sccd a pigcon cats may have a disrinctire color thxt thc pigcon can scc, but it makes no sound the An aninalis ccologv is kcv to understanding jts menral caprbilities. Some spccicsofbirds, such as Clark'.snutcracker, feed on sccds. $hen seeds arc rbundinr, these birds store seedsin buric.l cechcsso ticy sill hare food during the vinter. Thousands of seedcachesmay be buried and then l.rrer recov ered, sometimes as nany as rine months later. One rrculd er peccthe birds to hafe an extraordinary spatjalmenon, md this is indeeddrat has been found (fi9rrrf 5.t.7).Clark! nutcrrcher ard o&er seedJrardnrg birds, hare an unumally large hippo campus,thc ccntcr for ncDory storage in the brarn.

Jlgut'e )+. I
THE CIARK'S NUTCRACKIR HAS AN EXTRAORDINARY ErtoRr'. A Cl.rkl nutcrdcler NiLifnly .atxhbrltt. .n renenrber the locrtjons ofu! to 1000 seed crches nx,nths h d h ! r l - e r r . \ l '. , , . 'J . , r p '\ * i , r ,..f .',N ,. ''. r " hdle concluded thdt the birds use fertures ofthe laftlscape ard . 'J , , . ijpd.. rl, r,.' n .n ,,i ,( 'i \ . | {, , , 'u r d , , g the l{,crtn,ns ofthe crches.

H.bitu.tion is . simple fom ofle.hing th.t dos not rquire an ssociatiotr btween stinuli md respoNes. h cortnst, asoci.tive le.mins (classical md opemt conditionins) involves the fonnation of d asociltioD betweetr tqo stimuli or bctseen a behavior and a respons.

-**

{t!:

fitr

Jigure - 11.6

Lr,:ARNING wHdr rs EDIBLT.lssoci.trlc lcdrjng is inlohed in predarorprey interacrions. a. A naire tord is offereda bunblebeeds fbod. &.The toatl is stuns,and (.) subscqneDdv arcidsfeednBon blmblebees or any other insects hrrlng bl.cl .nd \ellow .oloratn,n. Tfie toarlhasasociated the appcarancc of the insectwith paD rnd rnodifies its beh.vior.

The Developmentof Behavior

Beharioralbiologistsnow rccognizcthat behaviorhas both geneticand lcamcd components. Thus fir in this chapter, we havediscrsscd thc influenceof gencsand learningseparately. But es 1'ous,ill see,these factors htcract during develrpment

Parent-offspring interactions infl uence cognition and behavior

As ar animalmaturcs, it nay firm socialattechnrenlt to other preferenccs individuals or develop that sill nrfluence behavior later in lift. This process,ca ed imprintin& is so sidered a gpe ofleaming. In lilinl imprinnng, soci^l attachrnentsform betweenpar enrsand offspring.For cgmplc, y,ung tirds of sonrespecies beginto fbllow their mothcr {ithin a feF hoursafterh.rrching, and thcir follownrgrcsponsc rcsul* in a bond beciveen mother l}ndtoung. However, the youngbirds'initial experience deter mineshorvthis inprint is estrblished. Thc Germanbehaviorist Konrad Lorenz shorved that gccscrvill fi,ll,n rhe firsr objecr thcv sccalter hatchirg rnd dircct their srcial behaviortoward that object.Lorcnz raisedgccscfron eggs,rnd when he of fered himsclf as a model for impr ting, rhe goslingsfferrcd him as if hc wcrc thcir parent,folbling him durililly (figrre i,+.8). The success of imprinting is highestduring.rcriricalpe riod (roughll ll to 16 hoursrkcr hatchinsir geese). Scveralstudiesdcmonstratc that rhe socialinteracrions that occurbctsccn parcnrixnd ofspring arekevto the normal delelopmenr of behariorThe psvchologist I larn Hrrlow grve rhesus monkeyinfants orphaned the opportuniwto fbnn social

Igurc )+.v
CHOICE TRIAL ON INIANT MONKEYS. Giren. choice bet$een a $ire iirme th.t provided food dnd r similir f.anre colered $ith cloth dnd gn en r monkev liLe herd, orphancd rhesusnonkeys chose the monkey like $sure orcr rhc tbod.

Igure )+.6

thev hatcherl, and

Lorenz$on the 1 9 73 NobelP r iz c phrsjology lor dns

attaclnnents with two surrog"te"mothcrs,"one madeof soft cloth coveringa rvirc liamc and the other madeonly ofwire (figlrc 54.9).Thc inhnts choseto spendrinle $-ith the clorh mothcr, clcn if only thc lire nn)therprolided food, nrdicating that tcxturc and tactile coDtact, rather than provisioDof food,maybeamongthckcvqualitics in a motherthatprornote infrnt socirlattac'hmcnt. Othcr studies showthat ifinfant mon kcvsare deprircd ofnormal socialconract,their der-elopment is abnornal. C;reater degrees of deprivrrionlead to greater in socialbe}avior during childhoodand adultabnornralities hood.Snrrlies oforphanedhunan infurs snnihrlysuggest that 'rmotherfigure" is requiredfor nonnal growth md r constant psychok,gical development. Recentresearch has revealed a biologicalneed tbr thc lunng prrenr o|f,pringinremcrior, that occur. 'rrnrrlatror early in life. Fenale rats lick their prps alier birth, and tnis stnnuhtioninhibitsthe relemeofa honnonelikcchcmicalthar can block nomal grolth. lups that receiverormal tactjle sdnuhtion alsoha1,e more brain receptors lbr glucocorticoid hornones,longer tived brainneurons,andr greatcrtolcrance for sress. Prernarure hurnaninhnts who are mrssaged ganr veighcrapidly.'l hese smdies hdicate that r\e necdnlr nornral social interaction is based inthe brain,rnd that touchandother parcnrs:rnd aspects ofcontactbenveen offspringare important sell pl-r.ir al r, r. behl ro rl dcrcl,'prnerrr. 'or Sexurl impdnting is a proccssnr which ar individurl learnsto direct its serualbchariortovard mennrers ofits ovn species. Cross-{ostering studics, in which indivnluals of one species arenised bv parents ofanothcr spccies, revealrhat rhis

rr22

r.rtJ..Ill{,t,,st

t.,t hth'tir

f<,rrl of imprinting also occursearly tu life. In most specics hrve sho*n that r ibsteredbird rvill atofbirds, thesestudies $'hen ir is nith rrernbers of irs lirstcr species tempt to mate

Instinct and learning rnay interact as behavior develops

llature malc rvhite crormed spirrows silg .r sPecies-spccillc Youngmrle birdsacquire courtship solg duringrnrtnlg season. a,on'bin"rion olilrind rnd Ie,nrirtg. r-he,olg 1,1 in soundresearchcrsrearednalebirds In oneexperiment, with sperLers and uicrophones ln proof incubators equipped rhisva'', rhcv couldcontrolwhatr bird heardasicnraturcdand then recordthe songit produccdasan adult. \44rite-cro$ted or thrt thrt heardno songit all during dellopment, sparrows dre song sparoq henrd only the song of a differcnt species, songasrdults (figure5:t.I0).But birds sanga poorly developed or thatheardthe soDgs thc songofiheir o'n spccies, thrt heard andthe songsprrrov,sanga sparros' ofrot the $-hitc-crowrled songasadults. white cros'nedsparaow firlly developed, r1r. |-h-eb;rd'h,'e : genedctemThhc re..rlr..uppe.r plate, or imrateprograrr, that guidesthem to ler the appropnare song. During a critiol period iri develop[rent, the temflatc uiil rcc(prr-hc Thu'. 'o,rg"cqui'iou,, core. r 'ong r' r ,,,"JcI. $e correct bt t onh rl'. 'orrP oepend. or' l, ,"rung. 'pecit'.,rt' "l be lerrned; r\c genetic telrplxre for leafling is setective. But leaming pla)'sr promnrentrole as well. lf a r,nung dcrfrker it hearsits species' rvhicecrorned sprrrov becomes

f.gure 54.11

laythet eggs in thelestsofother Cuckoos BRooDIArL{SITE. (lrrgcbirdro $e ofbirds. Bec.usc rhcpurg cucloos species srrecies .s dis mcadox isho de nisedbl' a dirTcrcnt Guch lean pipit,snuller birdto thclcft),thevh,veno opportnDiwto ihe cncl<oo sonsthel latershg is unatc. rheocko. song;

songduring the criricalpcrnld,it will sing a poorly devel<,pcd the bird must "practice"listcning songasan adrlt. l heref<rrc, to himselfsilg, matchingrvhathe heus to the nodcl his tbmof song devehJ'mcntstood Although this explanation hasshownthat for many years,recentresearch unchallenged song $tite-crowned spaffowmales.l, leam rnother species' under cenrin conditions-If a livc male stra$berry finch is the youngspr. placed in r cage next to a yomg Dile spirrroir', finch'ssong. l his firding ros-will lcarn to sing dre stmv-borry indicatcs that socialsdmuli in dLiscrse,beingableto seethe than a tape recorded sorg other bird-nui, be more effecdvc songdeveloptnent. in alteringdre innateprogranlthat guides 'I 1,"male.ol to 'ornebird sp<rrr'ha.eno opporrunrn herr rh<.rrg,'i rheiro{n \pec(.. l,,.uch ca'e'. ir ,upear. song. thar rhe malesinstinctively"kno$r' their own species' fernrleslav their For erample,cuckoosrre brood parasitcs; ofbird, andthe young drtt eggsin rhe nest of anotherspecies When hatch are rearcd bv the foster parents(figure 5'+.11). the cuckoosbecomeadults,they sing the song of their own Because mrle species rather than thrt of their foster parenrs. would most lihely he:r the songof thcir host brood parasires species during devehpment,it is id:rpti1for them to ignore such "incorect" stimuli. Thev hear no adult malcsof their own species singing,so no correctsorg morleisarc available nrtural selectiorhrs produccda completely In thesespecies, guideds.'l1g. genetically

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t\\\\\lt
1 .0

/,\v\i1fi't,|{'i'i\''
1 .0 2.O

b.

Igure ) +.tu
SONG DEVELOPMEYT IN BIRDS. A. ThC SONOgiADSOI

prcdrcedbv malewhitc-croutd spetows(Zd,,r,rs songs L,.orr']t thrt h.d been erloscd to their o*n speciej song aredi*crent hon (r) thoseofn.le during developnent sparros that hedrd !o sorg during rearing. This ditrerence programnselfis insufiicientto thdt the generic indicale5 a nornn song. Producc
srwra\cnD'or,g!..om

lrtmctions that occu duing striti' phases of de'roPmdt re aitic.l to nom.l heh,vior.l delopmenL Pltsiclr (mbct pLls r ihponant rcl in smrttr ud in the deelopnent of psrcholosic.l wl-hing.

,h"N- 54 b,ht"i",.t bi"t"s

ll23

The degree ro which anirn s "di*" is a subject ofltueil clisputc. Ilary of us hai'e obsen'ed the behaftr of a pet car or clog thrt woukl sugqest t}e ,rnim.rlhrd r degrccof reasoninsrbility or sas cxprblc of dinking. For manv decadcs,horvs'er, studens of aniLrl bchrf ior fladI rejected the notion thrt nonhumrn rntrxtls crn tiinl. In hc! behxliorist l.Lold ruorg:n stared in dre lNre ninctcdldr ccDtua'that one shouLdneler lsnmc I belllior represcnrs conscious dlourhr if rhere is an1 othcr crplanation that precludcs thc as$mptioD of consciousness. Thc pr6'xiling ,pprorch ras m trcet animrls asthoush ther responded r() the enrirornent drouqh h'tnrctirc bcharjoN xnd simple, nrnrtell' prognrnrned lerrning. In recentvcars,scriNs arrentionh:rsbccn gir.enro the topic ofad rd i$xrcness. lIe centrrl qucstion i: viether miDrls shc,w cognitive behavior drat is. do thq' prccess irlbfinati('l and rcslxDil in x manDerrhat n,ggtsts drinking (figurc 5'+.12); \46xr knnls ofbehirior $oulcLdcnloDsrr|te cogniti(n? A nunbcr 01 cases rre suggesrirec,f cognitne c'rpabilirics: . Sonre birds in urbm rrces in the mldtyerticdr cenmn; $+en nrillt deher\, tu ho renrore the foil cNpshon noDhon,ogenizcdmilk bortles ro ge( at dre cremr l)cncithj odrer birds lcrmed the be, harior through obscnarion; Japrnesenlcrqucs lcamed to vash srnd offpotaroes rncl io floal grNh to scpante ir from s,ind; Clhinrplnzees1rllrebccn ol)sened ro pull the le,rles off . rree brlrch Nrd thcn sdck rhe bnnch !,tr, lie el

i
i

54.13 Jigure

PR()aLFtISOLIING BY A ctI|M?A\ZEE. Lrmble n) glr rhc brnrrrs h! i!Dpnrg. the chimprn?cc derises r solutnn.

. .

trance o{a tcrnrire nesrj shelr tcmlitcs clinrb orto thc brrnch, thc chinrpanzeesrerri,vc rhc branch liorn thc 1tn]uDd!nd cat the lermires. Onh' a tev.erpc rncnts have testedth c t hinkinq r bilit1' of nonhurnrn .rnirnals.Some of rhese srirdics snqqesrrhat 'rnirnxls m.rr deiiberrtcll give felse intbrmetitn. Currenrlr. rcscarchers are trring to .lcternrine ii solrlc pdDr:rtesdecele orhers io nanipuhtc thc bellrI-ior of thc othcr rnembers of their lroop. llirnl uccdotal rccounts rppcer ro support thc ,1,| 1,,, de,,J,i . n,..h1,,, r , l- , r r e 1, ci es, such as baboonsand chi mprnzees, bur ir hr s been dif ficult to devbe fieldfiscd cperinrents to tcst thi! idea. r\luch of this wpe of research <)na.imal cogniti(n is in irs xrfi cr, bur ir is re to grow end to rrise contr(ncrs).. In rnv cirsc, nothi ng i s to bc gei ncd b).dogmati crl l vdcnring dr e possibil io oi rninal consciousress. So[re bchrriors. parricrlarll thost irro]rjng pfol,lcnl solving b\ aniDxls, are h.rd tc, erplxin in aDv s.av othcr than as r result of somc so.r of mentrl proccss. l:or er.mplc, in x series oi classic cxpcrnnenrs condudccl jn the I920s, a chjn pinzee { rs lefi in a roonr $jd} binrnas hxnqing frc,1llthc ccil irg out oI rcach. Also in dre ftnm ircre severrl borcs, cach l \ fp on, f l ," ,r. \frer...rr r .u,,r* trl . I I ( . , I r \ | . . I . | |,t r up Nnd grab thc binams. the chimpanzee suddenly kDked rr the boxes aDd ther inrnrediNtel) proceeded ro novc rhcnr un dernerth thc bananrs, stacl r:r]1c on top ofanothcr, and climb up to cl l j rn i ts pfi ze (fi gure i .l .1l ). Rcccnt sfl ies have fbund that anirnals c,thcr thaD prj, tn*es ;Llsoshov evidence of coqnitiof. RNrcns hare Nlsavs bccn c(Dsidefed rmong thc trost intelligent of birds. A recent cxpcrmenr usinq h.nd-rcrrcd ravens that livcd nr xD ourdoor rrian provided eaidencc of reasoning rbilitl: A piece olmert wxs placed or ihe end of ! s$ing and hu g 1lon e brrnch nr thc arian. The birds lited to ear nrertj but hld nerer seen string bclire and lere ulablc to qet it the mcat. Afrer ser-

Jigare 14.12

.\:{InaL TIINKING: 4, I his chnnpdrzee ls fil)fins thc lerles iionr . riliq, \hirh it sill thcn nr t., l,nn,e r tcrDire nest.ThN bch.!ior srlnrgh {elrcsrs rhm the !1"' puzce $ corscionshphnmlg rhetul,lrirh lirll tr1o c,lscof{lut it ntcnds ro do. ,. -lhis \ca oncr N lsing I .ock is .n "anrll." is.rinsrrili.h rt brshes , cl.m ro bredkit o]]cn.A se. otter rvill olicn teep r irvorite rocL rbr a long time, .s drolsh ir hs r .lcr ideaof ns tue uscof ihc ro..k.Bchxrxnsslch rs these mggcsltht dninrls hxlc cognitle rbihies. t, nV I lI t ar ' ( ,t,t ttra tt,,

l r2 1

erd hours, during vhich tjme the birds periodicallvlooked rt the meat but did nothing else,one bird fles to the branch, reached dovn, grabbedthe string with its beak,pulled it up, and pllced the string under its foot. It then rerched dovn rnd grabbedanorherpiece of the string, reperting this ac tion overand over eachtime bringing the melt closer(figrre Erentuallr; thc raven broughr the meat within reach 5.1.14). it. Thrce ofthc othcr 6ve ravens and grasped alsofi$red out hov to gct thc ncat. Prcscntcdvith a novel problem, the ravenhad der-ised a snutn,n.
beh,'ior of mihds Resea(h on the @gtrlr sone ruhples @ onr'tllin& in its inf.ncy, but 's

Igul'e )+.1+
PROBI,]i, 1 SOII'ING BY

a RAvr,N.Coniiorred Fith a problen ir hrs preuond)'frced, Dcver dle rarenfi$res olt ho$ to g.t rhe ne.t rt the erd of thc slrjtrgbr refe.tedh pllltug {p a brt ofstring drd steppngon it.