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    Semana 6 Stanford Et Al. (2009) CAP 15

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    Naaomy Duque

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    © All Rights Reserved
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    of 29
    walked trough the surgical ‘wars ofthe Bere hompal in ane An erin scart ard surgeon and Ste member of he French Senate, Broce wakes ike 2 man we war zed 0 aeope geting aut afi way He was tre tt merning 7 so meet paler. ramed Laborge no wie gay Lana Orme infection. Rather he wae Imerestes in Leborgne 2 & nrcogial pater with 2 long sary of bnormal banaver: [At he tre of roca’ meeting wth Laborgne, sets irtretd nthe hua bran were erbroied na Lncimera| dette about the nei of bran rion Some aus ‘atthe unctions of he oan ware evenly ributad twoughout the bran they Deleves th there ware no agovs oft bra het were spec er ry pa Deravar or fncion Others. suchas Brace, believed ta ar rome ofthe uncon ofthe bran ware base in, or localaed t,cartain spec areas, = Unerunaey forthe adiocste of localeaon the pzbudceiene of phranlogy The ilar ewpoie atnaugh he prvenoopss baleved they cule def Jocalied funcional area of he bran based on the exeral morgholegy—of ‘ye sc Tht the phrenloge slime were not bee or erin sues ch 1 prevent ahrealogy Fam Becaming a popu fa, anus troughout te wor * Wren Soca eamine Leborme, he found So-yearcld man wha was ery weak an cule no longer wae Hs sion was oor bu he Pear vas sl 2000 He Cay inden ha was beng sai ohn, ut he Pad eny one response te any queson aed of hin" 2s Broce ka the coepver (Leborgne tac beer uncer cre for mare tan 20 yea is parents in cher patents onthe ward ne eames bet Leborgre nad sere fom sates tem At age 70, rower Laberge lox the ait 2 ‘me he vas fom are lounge pee ah eg whch evertualy confined rim to hs bes | Laborgie var not seni o ane atough he ster patie ner corsicered tie 2 be epost anc re Because mos he ony ware he cou say was tn Ne became noun aTan to Te re ofthe espa The cher wor he cold sy was an ‘epic th he uternd when apts or angry Sroc inacvertenty ected this ‘epletie while rneztng 2 ts the: Labor fund sesame Labora des ony cayeatermestng Brom ante morning of Api 7. 26 hours rec hd perfor an autoay on the sate and ont ame Dvn some src of urganoe dicused Leborges case 2 meting ofthe ‘Chapter 15 + Evolution of ha Bran and Lange 439 FIGURE 15.1 Enccae rom South Arca aatiacee ‘Mort of te important quertions concerning hocinid brain evolution address ‘ways in which the human brain i diffrent fom the brains ofothes primates and rmameals. But there ate many ways in which our brains are similar to chose of other mammals, We use the same neurochemical, share a basie microscopic and macroscopic architecture, and have some basic sulci and ye around which funcional ‘epions are organized BRAIN SIZE AND ENCEPHALIZATION One of the defining features of the genus Homo, and especially of our own. species, i large brain sta. Bue what do we mean by “large"? In absolute tezms, the human brain weighs in at about 1,300 g, and human cranial eapacities usu ally are reported tobe in the region of 1,300 to 1,400 ce. These are average fig lures, and there is mach variation in brain size, Howeven, for purposes of rost-species comparisons, he 1,350-c estimate for the volume ofthe typical hhuman brain is good enough, “Look at the cranial capacities of various primates listed in Table 15.1 on page 4440, As you can see, humans have the largest brains among primates, The sec tnd largest brains belong tothe gorillas. Among the Old Warld moakeys, baboons appear to bave relatively large brains. As discussed in Chapter 6, among the ‘New World monkeys, spider monkeys have substantially largee brains than their close celacves, howler monkeys. To put these daca in a broader zoological con- text, catte have brains of abou 486 ce and horses of about 609 ee—somewhat larger than that seen ina great ape (Figure 15.2 on page 441), The botle-nosed dolphin has a brain volume of about 1,118 cc, which is neacly human-sized (Hofman, 1988), Encephalization Quotients Many scientists find absolute brn size values to be of limited usefulness in understanding brain evolution or the relationship between brain size and behavior. After al, i comes as no surprise that bigger nimals have bigger brains chan smaller animals, buc just because big animal hat ‘The mouse lemur may possess the smallest pri- mate brain—e welghs In at 1.8 grams. 438 _-Part¥ + New Frontiers in Biological Anthropology neurons The clr an of the nerus stad aren cot ‘acl boy td specnand process ‘aed ders wich cae pee fom other pron) nd sone (ourgrow oh wich hurt end mptes sor eros) cerebral cortex. The lit fray ‘Sater Sacco Ow sree fe ‘er hegre Sd ne ‘Gren rape corespond cel pater of euro orton ‘cerebellum Thee bn ‘Sched under be carb nd ‘porare nha colo aes porns an enary mone cerebrum Th bgt arf te Foran rar wen ete 2d ih hrphees Sea of “ahe" ban cons paleoneurology The suey othe ALTHOUGH THE HUMAN spcis possesses many features that help to make us sigue, ‘tis our complex behavioe and extraordinary traditional and material cultures oat fetus apart from all other animals, Our behavior is ultimately the product of an SSnatomicel feature the human bea. Complex cultural behavior is made possible by 2 specific behavioral adapration—language—which has evalved since the hominid lineage split fom the great apes. The stdy ofthe evolution ofthe beain ‘nd language highlights the relationship berween our bebavior and our biology. tn this chapter we wil eve the evluson ofthe human bain and language. The ‘human bean sa srucare of great complex andi produces bebaviors that are of ‘unparalleled sophisication inthe animal worl. Atsome point in homiaid evluton, Changes inthe bral t che appearance ofa spcis that behaved more lke ws and tess ike ou ape cousins, Commpared with the bens af our closest relatives, che human bin is larger and it exhibits imporeantdiferenees ints funcional organization ‘Some ofthese organizational itferences inthe human bran reflect he evolution of language. When we consider the fundamental importance of language to human Social existence; is not suprising that ia behavior tac is well represented in the onganization ofthe brain. Language has also helped shape the anatomy ofthe ‘hoa, leading to che development ofan organ of speech capable of producing an ‘extraordinary caage of sound’, ‘We wil ever know exactly when our hominid ancestors started to communi- cate with one another using a eytem of commenicaton thar was mor like language {nd less like the forms of communication used by other primates, Language, and the soft snus that produce st do.notfosiiz. But with a greater understanding ff brain function and of the natural history of language, anthropologists linguist, Derchologsts, and other sciensis in rcene years have turned tothe peoblem of Tenguage origins with inczeasing enthusiasm. Issues in Hominid Brain Evolution “The anatomy ofthe brain is rather complex. Athe microscopic level the brn is composed of billions of specialized eels called nearoas (nerve cells, which comm Imunicate with one another to form funesonal nenworks (see Appendix A). At the ‘table level, the cerebral corte, the surface ofthe brain composed of aeuron cell ‘odies is divided into complex partern of grooves and sdges called sult (sing Sule} and gyn (sng, slau), some of which can be used as landmarks co divide the brain into functional regions. Two of the major parts of che brain are the {Cerebellum and the cerebrum. The cerebellum, oF “Le brain,” sits cocked under the cerebrum, and is important in the control of balance, posture, and voluatacy ‘movements If alo plays an important rolein “higher” cognitive functions chat were {ace thought tobe solely under the contol of the cerebrum. The ceebrum itself ‘which i where most complex cognitive functions ar located, isthe part of the Train that most people recognize as being “the bral.” In humans and other pric ‘ates the cerebrum forms most of the brain's volume, andi is eneclly thought {hat the expansion of the cerebrum in human evolution has occurred asa direct = su of selection for more complex forms of behavior. ‘Recent technological advances have provided us with some extraordinary tools for examining the bran, bus the std ofthe evolucion of brain sruceure and fancton, or paleonearology, mains for she most part dependent onthe study of tndocast. Endocaste ate impressions of the interior part ofthe cranium, from which we ean make inferences about the size and structure of the brain (Tobias, 971) [Figure 15.1), Scientists make endocasts from fossil skulls, or in rae cases ndocasts form natuzally during fosszation (eee Chapter {1). Unforrunately, the brain s separated from the inside of the cranium by several protective tissue layers fod cerebrospinal fluid, so endocast ars inevitably a poor election ofthe brain's fbatomy, Nonetheless, they provide us with che only souzce of direct information tve have about th brain strucare of extinc species. Chapter 15 + Evolution ofthe Brain and Language 429 ‘South Alin antrpibecres Mort of the important questions concerning hominid brain evolution addcees ‘ways in which the oman brain is different fromthe brains of other primates and mammals. Bue there are many ways in which our brains are similar co those of ‘ther mammals, We use the same neurochemicals, share 2 basic microscopic and macroscopic architcrre, and have some bas sui an gy around wach funcional ‘epions are orgaaized. BRAIN SIZEAND ENCEPHALIZATION (One of the defining fertures of the genus Homo, and especially of our own specie, large Bean size. But what do we mean by "large"? In absolute terms, the human brain weighs in at about 1,300 g, and human cranial capacities us ally are eeported tobe in the region of 1,300 to 1,400 cc. These are average fig lures, and there ie much variation in brain size. However, for purposes of ‘rost-species comparisons, the 1,350-cc estimate forthe volume of the typical hhuman brain is good enough ‘Look at the cranial capacities of various primates listed in Table 15.1 on page 440.'As you ean see, humans have the largest brains among primates. The sec- ‘ond largest brains belong tthe gorillas, Among the Old World monkeys, baboons appear to have relatively lacge brains, As discussed in Chapter 6, among, the ‘New World moakeys, spider monkeys have substantially larger brains than their close relatives, howler monkeys. To pu these data in a broader zoological con- text, cattle have braine of about 486 ce and horses of about 609 ee—somewhat larger than that seen in a geese ape (Figure 15.2 on page 441). The botle-aosed dolphin has a brain volume of about 1,118 cc, which is nearly human-sized (Hofman, 1988), alization Quotients Many scientists find absolute brain size values to be of limited usefulness in understanding brain evolution othe relationship between brain size and behavior. Afterall i comes as no surprise that bigger animals have bigger brains chan smaller animals, but jst because a big animal has The mouse lemur may possess the smallest pri- ‘mate brain—it weighs in ‘at 1.8 grams. 440° Part V + New Frontiers In Biological Anchropology ‘encepalization quotient (EQ) ‘Fiera ome acual bran sof spaces fe gute ra sk Teton mscea rgresion of ‘neo s aes on ae umber of ese ee Se eee ees Cranial Capaciy (ce) Body Weight (ke) EQ Ho abi ml heats ne an ore pen eae 12052 2 re) an rile gra). sara 195 as an get gr omale wu ns tas Pon res (hime) eas ar Le ‘Pegs pence erngn, male 38 or a Feng ons (eeu). ale ys gor ‘OLD WORLD MONKEYS pana (boon). ale ass as hie Paani tabn). ers va hg us aah ier m3 10 le (Grateced marten itn ten ne 14 908 ut NEW WORLD MONKETS Aussi der ke) ra 40 as Act pts ove monk) ae 455 oe Saint sctrea egtrl monkey) aA 068 2 | _Sstet st eel penance lene Mee | ‘Noe ue to Koen 98 rH (98) orm or EO Nan ar monty it Senefetaresteiaen ta ncenie big brain does not mean thatthe animal is more intelligent. For many years, fciencnts have tied ta determine ways to measure bin sie relative to body siz. [Researchers suchas Facey Jeon (1991) and Robert Martin (1983) have shown thatthe tlationship between brain size and body size is somewhat more compli {ated chan a simple linear relationship. By looking at lage numbers of mammal Species, they derived equations tat allow us to calculate the expected brain ize {Sra mammal of any size, The encepbalization quotient (EQ) is s ratio of the factual bra size tothe expected sre. Thus mammals that have EQs greater than 4100 have brains that are larger than expected for a mammal oftheir size; an EQ Jess than 1.00 means that ts smaller chan expected, Returning to Table 15.1, me see that humaas have the largest brains not ont in ble bt alo ig elaine ey at ened bys FT ea Sacheopoid primates have EQs greater than 1,00, indicating tha their brains are larger than would be expected for mammals oftheir size. So even though cat- tle and horses have brains that are ape-sized in absolute terms, theie EQ are Chapter 15 + Evolution ofthe Brain and Linguge 441 HORSE: weigh, 40 ey cen capiy 600 ce (Pipa sigh 0 ig eral copy, 400 ce FIGURE 15.2 Encefaliaion a FIGURE 15.2. Chinpsreee suhores have ran ht resin #8 seul than those of pe cae of thes agr by ses ei geerly assumed thas the larger brain ie fn anchcopoid primates fag evolve in conjuction sth the evluon of complex soci eheor sad sdaptaon tote boreal (Can we say hat mammals wth higher EQS arin some sense “smarter han those with lower Qu? Yes at no Teneace Bescon (1997) plas ou thatthe ‘neepllaation quotients dsved from Bok brain se and ody se and at thant iw enc o reload the fae ta scl cng elecion pases ‘har hz bd as swear rain is. Among dog bees or explana are more caceplined than Genta spherical sccm on chiuahuns its driven boy se dowa at ater ste than bins gure 15) But to one (except sihsabon ances) woul arp that a chia sare than Gessan ‘hepherd In anhropotdy smal oc even dwarfed speci uch ate quel mow keyin the New Wor or the eslapin monkey sn te Olt Worl, have hgh ECs ‘Agi, ater han interpreting this a4 ign of are brain ie, we cul alo soe ‘es sn example of section fc sal body sat wich probly sore comet Golobine monkeys rend to have lower Er than cercopiiccine mocks (see she mangubey vers the colo in Table 15-1. There ao evden that Colobine behavior isin sme sense les sophisticated than cexcopidecne beat ioe (Figure 15-4, As cuca in Chapter 6 colobine moakeys ate adapted 0 2 leafy ds thi igenve cequtement bas diver slction for gener gut and boa sz, eslng in ower Qe. Calobites ar stil more encephalad than a typi mammal. Coils, who Bare arge brains in abvole sis aso have low Gs Again thes low-quality dc (aswell nother cron such on pro tection rom predation) may have doiven selection for larger body sine, earings them with Ee ower shan thet slovnfltves the ott apes Howevee i compaion of two cowl tlted species shating sparta: earronsen, Such athe spider and howler monkeys ri eetonale to hypornesze thie irae enn te ih ict monic uy bve red av ace of ewe opniive demands of fraicbused det, ln summary, the EQ is porewally EES valablelnittor of opine airy bur weneedofanembe tht visa ine. RCUNEIEA, Ta mei sion of both brain and body size, Ses conebutar 1 slower 6 447 PartV + New Frontiers In Biologieal Anthropology Sex Differences in Primate Brain Size In almost all primate species, males ave large brain than do females In che three highly sexualy dimorphic species Haced in Table 13-1 (orangutans, gorilla, and baboons, the absolute brain size {dilfences ae lige, a8 ate the body siz diferences, In each case, EQs forthe flmmales are substantally larger than for males. The EQ for male gorillas is below 1.0, indicating that thei brains are smaller than we would expect for « fammai ghee sive. There is strong selection for ineeased male body size in Highly sexuslly disocphie primate species, but there is no ceason to suppose Thar there are profound differences in behaviocal sophistication berween the ‘Even in less sexually dimorphic primate species, suchas rhesus macaques and humans, males have larger eins thaa females, This is tru after we correct for Pia ste Elolloway, 1980; Falk etal, 1999) Although we could speculate on Teo elecion forces on behavior or other biological proosses that might drive such eer iffereace, one conclusion is thatthe sex diffrence in brain size observed Juthuipans is nota function of recent evolution for higher cognitive function in hominids bu aems to reflect a general primate trend (Falk et al. 1399) BRAIN SIZE AND THE FOSSIL RECORD In previous chapters, you read that increasing brain sie is a characteristic of onus Homo. A compilation of average cranial capacities of different hominid Eesal caxa i presented in Table 152. (Please note thatthe H. sapiens values ix Tables 15-1 and 152 differ because they are based on different samples.) As you caa sc, the different groupe can be sorted to some extent according to theis resnial capacities and EQ OF coues, this comes as no surprise because cranial [Suck fone of the marphrlnpieal Features Wwe use co classify specimens into GiiGrene axonomie groups, Brain evolution in hominids can be divided into three phases (Followay etal, 2004). Phase 1: Early Hominids and Robust Australopithecus Brain sie increases from the early auszlopithecnes (A. afarensis and A africanus) to the "robust tustalopithecines," or Paranthropus. The early austalopitnecines have cranial exon ange (2) — 2 19 10-509 a Ader 7 4s 105-500 Ie bu on ose 7 so ss Hee , a1 sos Hance 2 ion 50-136 ch Here aw 1 on.1se6 onto ” tas 90-150 ee i aroha ort ide Dem i pon wlborOH Nu Eng Oy rer devel ing olf pchs cee cr cline . (Chapter 15 + Evolution ofthe Brain and Language 443 ‘eapacitis in the range of 400 to $00 ce, whereas the later A. robustue and A, boised aren che 475 t9 530 ce range. The exey australopthecnes have cranial capsciis Similar in size to those seen ia chimpanzees, orangutans, and femnale gorillas, ‘whereas the cranial capacities ofthe pacanthropines are more similar to those seen in male gor, ‘Ae che robust australopithecines species more encephalized than the ealies auseralopithecines? Ace grailes and robusts more encephalized than the con temporary great apes? Answers to these questions depead on estimates of body sass and bein sae. As we have already sea, gol have large brains, but they als0 hhave large bodies, especialy male gorilas, So they are not impressively eacephal- ined, (They are impressively big) Estimating boy mass of fossilized individu. als is very dificult and depends on how well sizes of available pacts of the skeleton correlate to overall body size. EQs calculated for any individual fossil Specimen theefore should be caken with a grain of salt. Heney MeHenry (1992; see also Kappeiman, 1996) estimates that A. afarensis, A. africanus, and A robuetus hhad male body sizes of 40 t0 45 kg and female sizes of 30 to 32 kg; A. Boiseh ‘was about 10% lager. These estimates indicate the these hominids were smalet than contemporary great apes given that theic cranial capacities wers a least as large, we can conclude that gracile and robust austealopthecines were indeed ‘oce encepbalized than the great apes. In addition, che brain sie increase seen in the robust forms relative tothe earlie: focms is also likely ro reflect an increase in encephalizacion, Phase 2: Early Homo and Homo erectus Hominid fossils assigned to Homo Inabilis oe early Homo have cranial capacities substantially laeges on average (by 25-20%) chan those seen in Australopithecus othe great apes (sce Chapter 4), Although the smallest early Homo specimens (for example, KNM-FR 1813, which has a cranial capacity of $09 ce) and the largest gorillas may overlap in cranial size che rlatvey small habiline body size, estimated by McHenry (1992) {0 be 52 kg for males and 32 kg for females, combined withthe larger brain sie, represenss an inczease in encephalization over earlier hominids, As you read taclies che appearance of H. hablis roughly coincides with the appearance of scone tool in the archaeological record, providing evidence ofa least one kind Of cognitive evolution “The average cranial capacity of fossil assigned ro H. erectus shows an even ‘more profound jump than H.babilis in both celative and absolute size compared ‘with earlier hominid taxa. Although botk brain and body sie increased in H. rectus, brain size may have increased relatively more quickly leading to an in- ‘tease in encephalization (Kappelmaa, 1996). As discussed in Chapter 12, H. rectus was widely distributed geographically and exhibited gradual change over Its more than 1 malion years in existence, On average, the ealest H. erectus spe=- itpens (such as KNM-ER 3883 and KNM-ER 3733) have smaller cranial apace then do later specimens, Thus the range of cranial capaciaes een in erectus speciens is quite lage from 650~1,250 ec), which is one reason char some investigators have justified spliting the taxon ito two or mote species, The recently discov. cred Dmanis cania from Georgia, dating to 1.75 million yeas ago, have cranial capacities of berween 600 and 780 ec; the smallest ofthese isa subadule(Vekua tal, 2002). Thee cranial capacities aze well within the eange of H. habilis and 7. erectus, but their cranial anatomy links them with HE. erectue, Phase 3: Archaic Homo sapiens, Neandertals, and Modern Homo sapiens CCranial capacities in the modern range are found in both archaic H. sapiens and Neandertal specimens. Indeed, one of the apparent paradoxes of the ater hominid fossil ecord i that Neandecal cranial capacities often exceed the average canal ‘capacity of modeen bumans (se Tables 15,4 and 15.2) Even the archaie H eapiens ‘ean is within the range of modern F.apiens. The increase in average anal capacity from H. erectus to the later Homo species is quite profound and undoubtedly 444 Part + New Frontiers in Biological Anthropology FIGURE 15.5 Athoush Neanéeral rants al wal we (or enced) (Se movera nae angeso £0 [ower oder paras bees ‘hoy ht trp soe, exceeds sny increase in body size, Thus the hominid trend for increasing brain size and encephalization continucs—and even accelerates through the appearance of archaic F. sapiens and Neandertals. “What about the apparent decline in brain siz in modecn humans compated with Neandertals and even with ester modern humans? ‘We should keepin mind that cere may be some kind of sampling bias (for example, rowaed larger males) alter ale have only small ‘numbers of fossils avalable to compare with large numbers of moder humans. More ertiealy, John Kappelman (1996) points tut that che larger body sizeof archaic H. sapiens and Neandertals felative co modern humans, often is overlooked or underemphaszed {Gee Chapter 13). Thus modern humans are more encephalized than. ‘Neandertals because their bodies are much smaller but thei brains fae almost as large as Neandertal brains (Figure 15.5) ‘Brain sz increase and increased encepbalization have chacacter- ized hominid evolution over the past 3 4 milion years (Figure 15). ‘These trends have become more marked over the past 2 milion yeams, as absolute bran size has neaslywipled. Dusing che past 2 Trillion yacs, increases in brain size have outpaced increases in body ‘ne, dhs leading to increasingly encephalized hominids. Although brain size and encephalization ate not everything, expanding brain Size in che hominid lineage claey elects an adaption, given how Sxpensive” beaindssue is (ace Insights and Advances: The Ten- Percent Myth: Evolution and Energy). BRAIN REORGANIZATION ‘As the brain has expanded, ts functional organization has also changed. We {Know ths by comparing ou brains with those of our closest relatives, suc asthe ‘Shempanate, oF ofthe ess macague, an animal often used asthe primate standard rani capciy lee 1a v0 es Howat on = 9 Os 10 1S 20 25 30 as Arche & med mans Ail of yoo FIGURE 156 Crna! pay has crane pproninatay fovea sien ers ftom olin f: ave at hoard the oye stozarnt ta not ony re may people heard bu hey bere ayehologt ary Beyerstein (1999) na pene ary ‘en des Althouse cnot arpa “gr wit praca, be ha sown ea Pasbor aura gut one ara On of Chef poupe techs ra pd (he yeh ae hu ay salampronrent (sti tng’ nase For exam {out7EN PEIGENT of ur bran power” 2c tt by aroling ie course Beng Seren panen ate sone Iran ae te IO gre hehe are conmon krowedge Tine dae Sega ot na mye matt safe nay acer an 1929 Alrough Bayer Seah rid oct hes Peychelogs who may tae sald some thing i ha havo fa let ed specie reararc ot t ercre. nthe 10 are cae en ‘ocnyth ac of of eareees tee wo aay nig Chapter 15 + Evolution ofthe Sin and Linguage AAS sens rom neces pyoegy ‘hae the 10% fares wl unanatla, Ravarch method ha dreey mans repose cane other ins vi reer of eure hat dawn mle Ore the ose ompaligapunents agains the 10% myeh comes from the Bran ues aloe of anergy le hua count for about 2 ofthe boy mast eur sou 1-208 fe oe eer {nd ony consumed yb body ei a ive save" (Alo & Whose, 1995)Te rain canoe wore snare ergy reves se cro varaile ihe yen sip ee From an erohtorry sanding mine ‘ling sch a experi organ on se [oid notre ry rhe a ed wih rg ch ra ‘eraee Gaper 6) f aed 108 he ban er wl are bons ‘als brat rece bein os orf cry se Th Bu ot apoan of course, bran exprsion ft chew evn ng ear cues om ran otto pre 2 rnch sega eae Hm oa Se epee ora mal usa Chen Pe tr cee cay ‘he stmach nd ns he tna inte Thate erations pesmaly teva btn congue yo retoton Te sal oh Siig eters oa ath The conplx sont Sarwtn be navontran stand pictur vane ct Sermon rang rosin inh sy hana cso dAtough ts erent toseroranszean git exprein Soest outst ne sandon eth 4 bemor conpios dont Nonny ‘Ato and Wher ae er ta me fre copy for wie we hve 8 ye src bh in in experimental neurological research. For example, there are pats of he rain that are essential for aormal language production. Because other primates do noc have language, obviously some recganizacion ofthe brain has accompanied the evoiton of language able. Although scientists debate the relative importance of organ zation and expansion ia hominid brain evolution, itis quite reasonable to assume that hoth processes have been crucial 446 Part V+ Now Frontiers In Biological Anthropology actor bbs Kaelin seu, ‘tet on enfant (oe erm reason of ‘jon he ba reontal replan The secon (Sint e pray motor ron (tbe pacer yar ae the ‘Noplomanal meta fe vestgators have tried to trace the evlusion of othe aspect of brain organi ation via both comparative anatomy and the examination of fossil endocast [Reorganization can accu in three ways: Am anatomical cepion of the brain a0~ (Gated with a speife function can become large ov smaller compared withthe fest of the brain of functional regions ofthe brain can shift o change position, [Eckepeadenty of tgional expansion or contraction. Alternately, new behaviors ‘maylead othe evolution of new functional Selds, which would supplant or enbance pesviously existing functional associations in chose areas. “We wil discuss examples ilustrating the first vo kinds of reorganization ia this section but wil save the third forthe secion on language late in this chapret ‘Several srades ave show that wen we look at large numbers of mammal species, ‘he anatomical organizations of their brains are semarkably uniform in terms of the relative sie of one structure compared wich another or withthe whole brain Jee oo, 1991; Finlay Be Daclingron, 1995). This i true whether the brains ae big or ital As with any scatistcal gencalizaton, there are exceptions. What we want £0 Ienow i, What exceptions are present in the humaa bran and when did they evolve? (Olfactory Bulbs In the human brain che olfactory bulbs, which control ous ease of svell, ave small, knoblike structures found on the bottom ofthe Fronal {bes in each hemisphere Figure 15.7). Compaced with other mammals, anthropoids have olfactory bulbs that are small for their overall brain size ([erison, 1991), meesurng only about D.1 coin volume (Seephan etal 1981) In contrast, wolves fave olfacrory bubs that are about 6 c in volume, 2 60-fold advantage over the bhaman-sized clfactory bulb. Humans have olfacrory bulbs that are about the sme Size as those found in strepsihine species whose brains ace only 1-2% the sizeof hhuman brains. "Humans rect (ia more extreme form) basi rend in olfactory bub reduction that we san ie inal living anthropotds. We presume hat this eduction occurred Sr other sensory domains (uch ae vision) and higher-level cognition became more important, reducing tliance on the sese of smell. Studies of endocests of the Oligocene primate Aegyptopithecus may indicate tha olfactory bulb reduction ‘vas slcendy present inthis easly antheopoid (Radinsky, 3979). Frontal Lobes Olfactory bulbs area good example of reorganization by size ‘eduction it the human brain, Atthe other end ofthe spectrum, many brain inves- figators [ouch a8 Deacon, 1997, but see also Holloway, 1968) have argued that ‘See ofthe largest regions ofthe bran, the frontal lobe, has expanded over the {Sours of hominid evolution, relative fo the ses of the brain (see Appendix A) Scientists believe thatthe prefrontal region—che parts of the frontal lobe that do fot inclade the primary motor gegions—hat showin a marked relative expansion FIGURE 5.7. View fhe The ctr bare ‘he ndrie fe rel Chapter 15 + Evolution ofthe Grain and Language 447 ye may havea larger prefrontal region (and frontal lobe) the size ofthe human brain In other words, would expect ‘Why should we otal lobe ela to overall bein i frontal lobe seem to coincide with many ofthe higher functions cae we associate with ineligencs specially wich the king 3 neligeace that we sem to have more of than any other animal sich a forcing goals and devising plane to aan them, It is not uatsazonable to price that given ‘ur apparent reliance on these functions, our prefrontal region should be large. Hloweres recent MEI studies of human, ape, and money brains conducted by Katerina Serendeferi and hee colleagues (2006, 2902) indicate thatthe frontal lobe is not proportionally larger in human brains (Fguce 15.8). They found that the frontal lobe makes up about 36-37% of the hemisphere in humans, orang. tans, chimpanzees, and goilas. Ie is proportionally large i hurmaas aed geet apes than in gibbons (25%) and in a combined sample of rhesus macaque sad ‘ebus monkeys (31%). These results indicate that we andthe geeat apes may share 4 small gceeae in elie frontal lobes. ‘The high forehead of modern hurmans compared wit che sloping foreheads of close relasves such as Neandereals aad acchaic H, sapiens might seem to bean ob- vious indication of frontal lobe expansion. As you recall in Chapter 13 Neander- tals and archaic H sapiens difer substantially fom moders humans inthe anatorty ofthe forehead and eye orbits: In general, ther foreheads slope backward from laege beowidges. Looking at profiles of frontal bones in crose-secion, Fre Book stein and his colleagues (1939) found chac despite differences in external mogpholo- 27 of the fronal region, the incernal morphology was remarkably smilayindeating that the shape of the frontal lobe probably was also similar in these groups Primary Vienal Regions The primary visual region isthe pare of che brain ‘where visual information from the eyes is nitally proceed, Although itis preseat In the ocipzal lobes (a the rear ofthe cerebrum) in both humans and other pe ‘ates in humans the primacy visual region is located ina sulcus onthe nner sur face ofthe lobe, whereas in primates the primacy visual cortex encompasses most » @ FIGURE 15.8. The fol ibe (ges) Bounced bye Svan Fesure low lin) a cones (es. 8) Pama) ‘hinge nd) eto 44a Part + New Frontiers In Biological Anchropotogy hunate sulcus A prominent tne el sie of ch epere STorenetra primase heh ‘Sele the pay ual or of ‘heel ob fom he reo the coeur ofthe lobe's outer surface. Furthermore the visual corex i smalle than we would kere fora pemate brain tsi: I i ony about 15 cies larger than the visual ExEts of a chimpanzee or gorilla, whereas the braia asa whole is about 3 times Iacpe (Stephan eta, 1981). The reduction and shi ofthe visual region in primaces prcrupably has allowed the expansion of the pasial cortex. The pacietl contr Eiiepion where sensory information from different sources is processed and Synthesized te alio important in tool usage ‘Controversy bout reorganization ofthe visual region in hominid brain evo tution has beea-not about whether bus about when it ocurred, The beginnings OF ebe controversy go ack to when Raymond Dare (1925) published his intial Aescripcon ofthe Taung child (A. africans) skull and endocast, In nonbuman painter, the primary visual zegion of the occipital lobe is reliably sepasated Fram the vest ofthe besin by the lanae slew, well-defined sulcus that i almost flways peesent. [a contrast in humans the lunate sulcus often is absent or very ‘ootly developed and it does not mack the primary visual region, which is poked by the calerine sulcus located on the interior surface of the occipital lobe (Figure 15.) “When Dact analyzed the Taung endocat, he confidently marked the lunate sulcus n'a posterion “human-like” position. He interpreted this to mean that ‘Sect the apelke size ofthe Taung bran, it showed evidence of human-Lkebeain oreo uation. This conclusion wae accepted for many years, but in the 1980s woiEsrous debate about the location of the lonate sulcus in Taung and other 1 eefopitheciaes broke out between Ralph Holloway and Dean Falk, ewo of ‘Ro aow experienced paleoneurologits working inthe field (Falk, 1980, 19836, {oe5s, 19836, 1989, 1991; Holloway, 1981, 1984, 1988, 1991; Holloway & ‘Kimbel, 1986), Falk argued that Dac’ positioning ofthe lanate was incorrect and ‘(Ririe va in'a more apelike posiion. Holloway, who initlly accepted Dares blavemene, argued that Falk's positioning of the lunace on the endocast was cise anatomically untenable. Currently the weigh of evidence, including new ‘xtovertes and fucther reassessments of older specimens, indicates that rhe Tunstesulews was located more posteciorly in australopichecines compared tits [ofan in the great apes, and thus the primary visual epion was ina more human- like poston Fioloway etal, 2004), This change in postion marked the begin hing of the extensive reorganization ofthe visual regions of che human brain, Compared to great apes and other primates. » FIGURE 15:9 Frinay vital roceing srs) in) burn tera an aleve) ane) compan. furans te priary ial ares urea the ‘lear sn hinges dent cs forms arterir Bound ‘Chapter 15 + Ewcucon afeha Grin and Langage 449 Language: Biology and Evolution ‘Much of what makes human behavior more complex and moce sophisticated than the behavior of other animals depends on our posession of spoken language, tis one thing to possess sophisticated cognitive ablities—to make plans o draw complex cause-and-effect relationships between the things you see inthe env. fonment; and to think in terms ofthe pas, present, and fatare—~bat without the ability to convey these thoughts to other members of the socal group, their ‘usefulness for enhancing survival and reproductive access would be limited. ‘Language isan adaptation. eis easy fo imagine that a social group of hominids who possess language would have an advantage over social group Of hominide who did not. Language ably, oweves is as much an anatomical asa behavioral adaptation. Modern humans are designed by natural selection-—in the anatomy oftheir throats and respiatory system and in various aspects ofthe strucire and function of theis braine—eo produce language, ‘Whats language? Language isthe system of communication wed by members ofthe human species. Alchough linguists differ on which features ere most eritcal in defining language, they all cend to agees on cecain critical aspects that make language a unique form of animal communication. Language is spoken and we ae anatomically specialized to produce language and to process language-orienred Sound. Language is semantic: The woeds we use when speaking have meanings that epeeset real-world objects, events, or actions. Language is phonemic. Words are ‘made fom small sound elements called phonemes; thece is no biological limit to the number of words that can be foemed from phonemes and there iacisic association berwein a word ad the objector concept it represents Finally, language 4s grammatical. All languages have agrammas an implicit set of rules that governs the way word dase are defined and ued. Although thre may bea lation te number of words a person can know, there is no limit on the ways the may be srammatically linked together. As a child acquires its fist language, he or she assimiares the grammatical rales of language subconsciously. ‘THE EVOLUTION OF GRAMMAR “The place of grammae in defining language and studying its evolution has been 4 point of controversy over the years. One school of linguistic thought, ed by Noarn ‘Chorasky (1967), placed grammar atthe center ofthe linguistic universe, Chow sky and his followers (such as Jackendoft, 1994) argued that by studying the gen- eral grammatical rules of language, we cn find a "deep steucture," which in arn isa reflection ofa “mental grammar” found in the brains fall people. Evidence cof the existence of mental grammar comes from language acquisition in chldzen ‘With litle effort children master th rules of grammar of any language to which {hey are exposed, despite their complexcy. Linguist Steven Pinker (1994) has alld his ability the language instinct: Children appear co be genetically specialized to leara language. ‘An intecesting piece of evidence ofthe relationship between children and a possible deep structure of language comes from the study of pidgine and eects. Pigs ae simplified, nongrammatical communication syetems that have aise it aceas where speakers of diferent languages need to communicate with one at ‘other but do nor spend enough time around each other t learn each other’ lan {uages (aew colonial situations, fishers from different countries mecting on the Seas) [nconras,ceoles are grammatical languages that have arisen and developed, typically in colonial situations (such a in Hawati or New Guinea) in the context ofan ongoing situation of linguistic change or instability (Figure 15-10 on page 450). te has long been notied that creole languages around the world converge on 4 similar grammacieal secure. Linguist Dezek Bickerton (1983, 1990) suggests English-speakers recognize the phrase “Colorless green ideas sleep furiously” as grammatical, even though It is nonsensical 450 pare + New Frontlers in Biological Anthropology ee Hewaion Gace Fin Engi Tulding Tigh place wal] Gear thre cn eee port inertia sed | sgn high us on cowl of Roncpomengoraure | dabuting show you wot ary tine ae os Sine on temperature gett FIGURES.10. © Ss ociereene parson) pn rat Seproincese ® thatthe source ofthis convecgence is nota common language of origin but he fact that creoles are invented by children who share a common, biologically based, deep ‘Ratcrue for language. ‘The first generation of children growing up in these grapeed linguist environments will not tolerate a nongrammatical sytem of Communication, and they impose Linguistic structure on the language around them, chs leading t the developmens of ceoes ‘aay advocatss ofthe deep grammar point of view believe that language repre- sents a cognitive process that is fundamentally diferent fom that underiying Se cokove ofaimal communication. Hower, several recent evolutionary ‘Rbrst of langeage have argued aginst the exseace of universal menzal rama (Gostge Rumbsugh 8¢ Rumbaugh, 1993; Hurford, 1991; Schoenemana, 1999). Seose Rumbaugh and Rumbaugh believe that syatax and grammar must develop tuct anyone tiesto go beyond a two-word urerance; rules have to exist co let the [ener know what the speaker is aang about. They write, "Whatever commonsl- fhe bee ae among grammars may well exist because onlya limited number of so- Jusoas to the same problem are workable, given the constrains placed on the probless Int” (1993, p- 106-107), Thus grammars inevitably emerge, but there sno uni- ‘opal gramme Sach a posiion ie consistent with te view that man language ex- Fibs voluconary continuity with other forms of animal communication, because it tioes not posta zoologcally unique cognitive mechanism, such as a deep mental grammar forthe evolution of language (Figure 15.11. LANGUAGE INTHE BRAIN {We can define language area ofthe brain as any par of the brain thats activa ‘ei daring the produerion or comprehension of perc, The casa language regions ‘re found around the lft Spvian Ssure, or persian language are (Figure 1512) In the froatal lobe, there is Broca’s area, As we sav eazlie, « lesion in Broca’s trea causes a disreption in speech production (an aphasia), ye comprehension re- fiains inet. At the posterior end of the Sylvian fisare, spanning the top of the temporal lobe and the bottom of che parietal lobe, another language area was leaded by German physician Carl Wernicke in 1874. Werncke's area lesions ‘Guse a person to have difficulties in speech comprehension, People with Wer, ‘icke' ates aphasia produce fluent but nonsensical speech, subsitting one word for another or producing incomprehensible strings of words, Wericke predicted ‘het because i is likely that his azea and Broca's area are in communication, {Efferene lesions in the white matter joining the two should produce aphasas with ‘ferent symproms. These conduction aphaas have been obsecveds for example, Chapter 15 + Eioluton ofthe Grin and Language UNIVERSAL GRAMMAR VIEW | Origin Men . Cranmer aly ——pe aren Discontinuity | 7 Teen orn Barer os ‘elton ‘Asin Communion Eat ‘ - Continvy Fam ceed onmunicaton % rors See Humen aqvene EMERGENT GRAMMAR VIEW. FIGUNEI5.11 Th nvr ranma ard amersnt rma views leno very Cioran sensi aha ecu of agus 4 lesion inthe projection feom Wernick's area to Broca’s area causes someone to produce fluent, aoasensical speech wile taining compreheasion (Damasio & Damasio, 1989 ‘Wernicke's insights bout conduction aphasias taught us think about language se de product of inecactve nerwarks Inthe beain rather than Of ust one oF Oo areas, In adizion to Broca’s and Wernicke's area, the perisylvian language atta include several ocher regions important for speech, Inthe fatal lobe, Brocas ace sits justin fron ofthe motor strip controling the tongue and mouth, which are ‘obviously involved in speech production. Along che rop of the temporal lobe lies the primary auditory cortex, where sound signals from the ear are initially processed, which is essential for speech perception. The angular gyrus in the Parietal lobe is imporant forthe comprehension of written language. This is not Surprising because projections from the primary visual caeex isthe ociital lobe passthrough the angular gyrus on the way eo Wernicke's area, Language Lateralization When a function of the brain typically and consis tently occurs in only one of the hemispheres, we sy that function is lateraized FIGURE 15.12. Th ajar rage arts fh eft emp th bran. The nnn eee Waichasan Brc ara pase ough fe area ys 452. Part¥ + New Frontlers in Biological Anthropology In general, chimpanzees show no strong hand pref ference at the population level although Individuals may have a preference for certain tasks. 1n.95% of people, the persplvian language area is inthe left hemisphere. Most people are alo right-handed, and because motor contzol of one side of tke body Er petsed in the opposite side of the bran, ii very Ukely that right-handedness vou language ably evolved in tandem. The classical view that both language and {ihhandeness ate associated with the left hemisphere has led 0 the notion of [eft kenisphere dominance over the cight hemisphere (excep in about balf of the Iefchanders—who make up about 10% of the population—who have eight hemisphere dominance) "ARhough fis easy to focus on the classical lef pesisylvian cegions asthe seat of lnguage Keepin ind that lesions in other parts ofthe brain also disrept coral sticch, Lesions inthe right hemisphece (of people with left hemisphere Jhaguage dominance) disrupt the matical or prosodic elements of speech. Prosod peeaabtal for speech co sound nocmalothervise, c would have the Hse sound of ‘Stteatersynthesized speech, Lesion inthe right inferior frontal lobe (opposite Groc's are) lead to deficits in the production of normal prosody in speesb and isons in the sight hemisphere opposite Wernicke's area lead to deficits in the Spechension of prosod} in specch (see Innovations: Musi, the Brain, and Evo~ eereage 454). Neatoimaging studies have shown thatthe numerous pars Up bead dedicated co the control of the lips, tongue larynx, and voluntary con- Clot ot the dapheagm ar active during speech production [Wise eral, 1999) LANGUAGE INTHE THROAT [Although thete is itl evidence that evolving language capabilities has cos us avthiag in terms of Brain function—jost the opposite in fact—itis quite cleat adr the eartangement of the anatomy of our throats fr language purposes bas fneroduced sew risks in everyday life that our ancestors did not have to worry {5S0Eadaman, 1984; Lieberman, 1991}, To ose thee cs, cece must have best a Strong selective advantage for the developmeat of language abilses over the course of hominid evolution, “The supralermigeal veay is more precise way to describe the pars of the throat ‘and head that have undergone changes during hominid evolution (Figure 15.13. $i the ante sugges, iis that pat ofthe airway thatis above the ary, oF voice ibe SThe larynx sts at the top ofthe rachea and has vocal folds (vocal cords) Rich can modulate the passage of air through the caches to produce different Sounds: The eavity above the lary, atthe back of the mouth is known as the ‘ham. The posterior part ofthe congue, the epglots, and the soft palate form the boundaries ofthe pharynx. FIGURE 15.13 The supeabrynes say ns chingarate anda aman. Nasr rae Saarrmntas hava nd Nu bc of te hand aed sora tng erm Grtroe par of para. Chapter 15 + Evolution ofthe Brain and Langinge 453 When we compare the supralaryngeal airway of « human with that ofa more typical mammal, suchas a chienpanzee, we can see several differences that have profound functional implications (Figure 15.13). Fis, the laryns ia humans is ‘much lower than in other mammals. The nay postion of the larynx leads fo an expansion ofthe pharynx. This expanded pharynx, whose aateriog walls formed ‘uniguely in humans bya shortened and rounded tongue, ie much more efficent fot ‘modifying the stream of air passing tarough the larynx to generate a greater varie of sounds, leading to flly articulate speech. In other mammals the seal phatyns has vey lle capaciy for modifying the sounds produced by the lacy, Supralaryngeal modification of sound can be done only by alteation of the shape ‘of the oral cavity and ips (Laieman, 1984) (se isights end Advances: Ape Lan ‘guage Soles on page 456), ‘These changes in anaromy havea profound cost, however: hey greatly increase the risk of choking on food or liquid. There is too much distance becween the hhumaa larynx and nasal cavity foc a sealed connection to form between the cwo, as ic does inthe typical mammal. The epiglatss and sft palate are separated by the rear part of the tongue, Everything we swallow must pass over the income pletely sealed opening of the lacyax, which greatly increases the csk of choking and Suffocation, Interestingly, human babies less than 1 year old have a supralacyngeal Anatomy that more closely resembles the mammalian norm, This lows them to sik, swallow, and beeathe a the same ime, which greatly enhances their suckling bility. Dusing the second yeas, the larynx begins the sife to the adult positom, which increases ther isk of choking while increasing thei ability to produce articulate speect, Darwin noted in On the Origin of Species thatthe postion of fhe eachea in the human throat was an example of natuel selection warking with wat history makes available toi LANGUAGE ABILITY AND THE FOSSIL RECORD The bran and supealaryngealteact—anatomical steuctures chat demonstrate most cleaey our adapeations associated with the production of spoken language—are ‘composed primarily of sof dssues that do aot fosilie. Howevey, we do have en

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