The Energetic Paradox of Human Running and Hominid Evolution'
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CURRENT ANTHROPOLOGY Vol. 25, No. 4, August-October1984 K 1984 by The Wenner-GrenFoundation for Anthropological Research, all rightsreserved 0011-3204/84/2504-0005$2 00 The EnergeticParadox of Human Runningand HominidEvolution' byDavid R. Carrier THE FEATURE THAT DIFFERENTIATES HOMINIDS fromother mal thesize of man shouldconsume roughly 0.10 ml of oxygen primatesis notlarge brain size, but theset ofcharacters associ- per grambody mass per kilometertraveled (Taylor, Schmidt- ated witherect bipedal postureand a stridinggait. Long before Nielsen, and Raab 1970), but the measuredvalue forman is the rapid increasein relativebrain size thattook place during over twice this amount (0.212 ml). A recentanalysis of 64 thePleistocene, early australopithecines possessed the postcra- speciesof runningbirds and mammals(Taylor, Heglund, and nial morphologyof an erect,striding biped (Lovejoy, Heiple, Maloiy 1982) confirmsthe initialobservations that the cost of and Burstein 1973). Evidence for the bipedal gait of early transportis relativelyhigh for human runners. hominidsis provided by the morphologyof the 3.5-million- It is somewhatsurprising, therefore, that among cursorial year-oldpostcranial material from the Hadar Formationof mammalsman is one ofthe best distance runners. While game Ethiopia (Johanson, Taieb, and Coppens 1982, Lovejoy, animalsare fasterover short distances, they generally have less Johanson,and Coppens 1982), earlyPleistocene material from endurancethan man. Aside frompublished accounts of endur- eastern and southern Africa (Preuschoft1971), and the ance performancein thedomestic horse and camel,which have Pliocenetrackways (3.6 to 3.8 millionyears old) discoveredat been bredfor endurance racing, very little is knownabout the Laetoli in northernTanzania (Leakey and Hay 1979, White stamina of mammals. What littleinformation is available is 1980). Together these suggest that early australopithecines usuallytaken fromanecdotal accounts of animals performing were relativelysmall-brained creatures, possessing structural in a varietyof climatic and topographicenvironments (see adaptationsfor upright walking and runningthat in a broad Howell 1965), and consequentlyinterspecific comparisons are sense are remarkablysimilar to those of modernman. The hazardous.Comparisons between man and a numberof curso- aspects of locomotionthat unite the hominidsas a groupare rial species can be made, however,in the contextof a contest also uniquelyhominid in characterand distinctlypeculiar for betweenpredator and prey. mammals. Consequently,the studyof human locomotion,in Huntersof a numberof differentcultures are knownto run additionto explainingmuch of the biologyof the one remain- down prey by dogged pursuitoften lasting one or two days inghominid, may proveto be one of themore powerful induc- (Krantz 1968, Watanabe 1971). Bushmenare reportedto run tive approachesto the studyof hominidevolution. down duicker,steenbok, and gemsbokduring the rainy season The energeticcost of transport (oxygen consumption per unit and wildebeestand zebra duringthe hot dryseason (Schapera body mass per unit distancetraveled) for running humans is 1930). TarahumaraIndians chase deer throughthe mountains relativelyhigh in comparisonwith that for other mammals and of northernMexico untilthe animalscollapse from exhaustion runningbirds. Early comparativestudies showed that a mam- and thenthrottle them by hand (Bennettand Zingg 1935,Pen- nington1963). Paiutes and Navajo of theAmerican Southwest are reportedto have huntedpronghorn antelope (one of the 1 This contributionarose out of conversationswith D. M. Bramble fastestof all mammals)with this same technique(Lowie 1924; and S. C. Carrier.Further development resulted from discussions with Foster 1830, citedby Lopez 1981:111).Furthermore, Aborigi- C. Gans and M. H. Wolpoff.For criticalreview and additionalim- nes of northwesternAustralia are known to hunt kangaroo provementof this manuscript,I am gratefulto S. L. Beck, D. M. Bramble,S. B. Emerson,J. R. Fetcho, D. B. Jenkins,R. G. North- successfullyin thisway (Sollas 1924,McCarthy 1957). In these cutt,P. A. Pridmore,and R. 0. Whitten. examplesthe staminaof preyanimals generallyconsidered to be specializedfor running and presumablyputting forth their best effortproves to be less than thatof man. Energeticallyinefficient transport and endurancerunning DAVID R. CARRIER is a doctoralcandidate in zoologyat the Uni- would seem to be incompatible.The combinationof thesetwo versityof Michigan (c/o Division of BiologicalSciences, University of Michigan,Ann Arbor,Mich. 48109, U.S.A.). Born in 1955, he attributespresents obvious morphologicaland physiological was educatedat the Universityof Utah (B.S., geology,1980; B.S., problems.For instance,there must be a way ofdissipating the biology,1980; M.S., biology,1982). He views himselfas a func- highlevels of metabolicheat producedduring long-term mus- tionaland evolutionarymorphologist and is particularlyinterested cle action. In addition,extended periods of high energyde- in vertebratelocomotion and the ontogenyof the musculo-skeletal mand requirelarge body stores of glycogenand fattyacids and system.His publicationsinclude "Postnatal Ontogeny of the Mus- an effectivemeans of mobilizingthese stores.It is clear that culo-SkeletalSystem in theBlack-tailed Jack Rabbit (Lepus califor- nicus)" (Journalof Zoology [London] 201:27-55) and, withD. M. duringthe sequence of evolutionarysteps that led frompre- Bramble, "Running and Breathing in Mammals" (Science hominidsto Homo sapiens these problemshave been over- 219:251-56). The presentpaper was submittedin finalform 9 I 84. come. How this was accomplishedand what factorsmight Vol. 25 No. 4 August-October1984 483 This content downloaded from 155.97.178.73 on Fri, 27 Nov 2015 17:24:13 UTC All use subject to JSTOR Terms and Conditions have initiatedthis sequence of specializationis the subject of ward flow of heat fromthe hot environmentthan sweating the presentanalysis. animals.Taylor and Rowntree(1974) have shownthat in a hot environment(47?C) heat flowsinto sweating animals at about twicethe rate thatit flowsinto panting animals. For the same DISSIPATION OF METABOLIC HEAT reason,when environmental temperature is low thecooler skin temperaturein sweatinganimals resultsin a smallergradient The normal core body temperatureof eutherianmammals fornonevaporative heat loss (heat loss by radiationwhen skin ranges from36?C to 38?C (Morrisonand Ryser 1952). The temperatureexceeds ambient temperature).Thus, sweating lethalcore temperatureof thesemammals is only6?C higher, animalsgain moreheat froma hot environmentand are capa- rangingfrom 42?C to 44?C (Adolph 1947). Using empirical ble of losingless heat by nonevaporativemeans than panting measuresof the cost of transport,Taylor (1974) has calculated ones. that the maximal exerciseheat loads generatedby medium- 2. Because pantingrestricts evaporation to the nasal, buc- sized and large animals (greaterthan 10 kg body mass) are cal, and pharyngealcavities, pantinganimals lose littlesalt. nearlyan order of magnitudegreater than the maximal en- Sweatinganimals, on theother hand, mustdeal withlarge salt vironmentalheat thatthey are likelyto encounterin nature.If losses (Schmidt-Nielsen1980). metabolicheat is produced more rapidlythan it can be dis- 3. Pantingprovides forced convection across the evapora- sipated,body temperature rises. Thus, adequate thermalregu- tivesurfaces, whereas sweating requires animals to relyon free lationis criticalfor animals that run for extended periods. For convectionor wind. (This should not be a problemfor a run- example,cheetahs sprint at speeds exceeding100 km/hr,and ninganimal, particularlyif it lacks hair.) extrapolationof measurementsmade at lowerspeeds suggests Evaporative coolingfrom the skin offersthe followingad- thatat 100 km/hrheat production would be morethan 60 times vantages: greaterthan at rest(Taylor and Rowntree1973a). The cheetah 1. Sweating provides an additional evaporative surface. storesmost of thisheat, and thereforeas it runsits bodytem- Taylor and Rowntree(1974) measuredthe maximumrates of peraturerises. Once the rectaltemperature reaches 40.5?C the evaporativeheat loss froma groupof pantingand a groupof animal refusesto run. Taylor and Rowntreeestimate that if a sweatinganimals in responseto exercise.They foundthat the cheetahterminated a sprintonce its bodytemperature reached ratesof evaporation from the respiratory surfaces were similar, 40.5?C, it could run for only 1 km. This approximatesthe but the sweatinganimals had much highermaximal rates of actual distancethat cheetahs pursue their prey in the wild. It evaporationbecause of the evaporationthat occurredat the seems that the durationof the cheetah'ssprint may be deter- skin surface. mined not by fatiguebut by the amountof heat it can store 2. Sweating is independentof the respiratorycycle. As a beforeit reachesa limitingbody temperature. runninganimal increasesspeed, its oxygenconsumption in- Mammals dissipateheat by evaporativeor nonevaporative creasesin a linearfashion (Taylor, Schmidt-Nielsen, and Raab means. The Africanhunting dog is exceptionalin that it ap- 1970). However, as runningspeed changesit is unlikelythat pears to dissipatelarge exerciseheat loads, forextended pe- therespiratory air flowrequired for metabolism will matchthe riods, throughnonevaporative means (Taylor et al. 1971). flowneeded forheat loss, since the rate at whichheat can be Most medium-sizedto large mammals rely on evaporative dissipatedwill vary