Physical activity evolution - obesity

SHORT COMMUNICATION

Low physical activity levels of modern Homo sapiensamong free-ranging mammals

M Hayes1, M Chustek1, S Heshka1, Z Wang1, A Pietrobelli2 and SB Heymsfield1*

1Department of Medicine, Obesity Research Center, St Luke’s/Roosevelt Hospital, Columbia University, College of Physiciansand Surgeons, New York, NY, USA; and 2Pediatric Unit, Verona University Medical School, Verona, Italy

Obesity prevalence rates are increasing worldwide and one prevailing hypothesis is that physical activity levels of modernhumans are markedly reduced compared to those of our Paleolithic ancestors. We examine this hypothesis by deriving relativeactivity energy expenditure from available doubly labeled water and indirect calorimetry data in free-ranging non-humanmammals. Our results, given the constraints posed by limited data availability, suggest that a low physical activity level, muchless than that observed in free-ranging non-human mammals or highly active humans, is present in modern adult humans livingwithin advanced settings. Our observations lend support to the hypothesis that low activity-related energy expenditure levelscontribute to the rising worldwide prevalence of obesity.International Journal of Obesity (2005) 29, 151–156. doi:10.1038/sj.ijo.0802842

Published online 9 November 2004

Keywords: energy expenditure; physical activity; doubly labeled water; resting energy expenditure

As living standards rise, countries all over the world are

experiencing an unprecedented increase in the incidence of

obesity.1 Obesity is a consequence of a long-term positive

imbalance in individual energy intake and expenditure

leading to excessive deposits of adipose tissue. One current

hypothesis is that levels of energy expenditure as physical

activity are markedly reduced in modern day humans

compared to conditions under which our Paleolithic ancestors

evolved and that biological regulatory mechanisms have failed

to adequately down-regulate food intake to prevent positive

energy imbalance in genetically predisposed individuals.2,3

A critical missing component in this account is a lack of

data on energy expenditure from physical activity in prior

time periods. It would be of interest to know how active

primitive humans were prior to the time when modern tools,

labor saving devices, and social organization reduced the

needs for food gathering and defensive activities. Our

analysis that follows reveals interesting trends but also

highlights the paucity of information on key measures of

energy expenditure in mammals.

As humans share with other mammals many physiological

and metabolic characteristics,4 we estimated early human

activity levels based on the assumption that primitive

humans and free-ranging mammals today share in common

the same approximate relationship between free-living

energy expenditure (FEE) and body mass. The FEE estimates,

adjusted for body mass, were expressed as multiples of resting

energy expenditure (REE), also adjusted for body mass. We

then examined concordance of our results with human group

data from active individuals living in rural areas of develop-

ing nations and from selected individuals participating in

high levels of occupational and recreational physical activity.

Finally, we compared these results with observations from

representative adults in developed nations.

Free-ranging energy expenditure was evaluated using a

composite database developed from previously reported and

new (six Kenyan yellow baboons) doubly labeled water

human and animal studies (Appendix). REE was estimated in

earlier studies by conventional indirect calorimetry and we

assumed that diet-induced energy expenditure (DEE) repre-

sented 10% of energy intake and thus of FEE under weight-

stable conditions. Activity energy expenditure (AEE) was

then calculated as FEE�(DEEþREE). We expressed energy

expended in physical activity as the ratio of AEE to REE.

Our results for the FEE (in kJ/day)-body mass (M, in kg)

function, based upon doubly labeled water estimates,

indicates FEE¼800�M0.73 (Figure 1). Our results for theReceived 5 May 2003; revised 26 October 2003; accepted 17 November

2003; published online 9 November 2004

*Correspondence: Dr SB Heymsfield, St Luke’s/Roosevelt Hospital,

Weight Control Unit, 1090 Amsterdam Avenue, 14th floor, New York,

NY 10025, USA.

E-mail: [email protected]

Supported by National Institutes of Health Grant RO1-NIDDK 42618.

International Journal of Obesity (2005) 29, 151–156& 2005 Nature Publishing Group All rights reserved 0307-0565/05 $30.00

www.nature.com/ijo

REE–M function, based upon compiled indirect calorimetry

data, indicates REE¼253�M0.75 (Figure 1). As the powers of

the FEE–M and REE–M functions, 0.73 and 0.75, do not differ

significantly, the AEE–M relationship can be derived in

mammals, AEE¼467�M0.75. Accordingly, AEE/REE¼ (467�M0.75)/(253�M0.75)¼1.8. The evaluated free-ranging

mammals thus expended energy in physical activities about

1.8 times their REE. This AEE/REE value of B1.8 can be

equated with the widely used physical activity level

(PAL¼ FEE/REE) of 3.1.5 The AEE/REE and PAL can be

interconverted as: AEE/REE¼0.9�PAL�1; and PAL¼1.11�(AEE/REEþ 1). We also examined the AEE/REE and PAL in 33

species of smaller mammals (body mass, 0.2170.45 kg) on

whom both FE and REE measurements were available

(Appendix). The calculated AEE/REE and PAL in this group

were 2.271.2 and 3.671.3, respectively (Table 1 and

Figure 2). Speakman,6 using some of the same data as in

our analysis, observed a mean PAL of 3.3 (s.e., 0.2) for 74

small mammals weighing o2 kg.

Our assumptions and calculations support the view that

the AEE/REE level of primitive humans, before the develop-

ment of efficient tools and agriculture, approximates 1.8.

Modern humans engaged in vigorous physical activity such

as rural farmers in developing nations, athletes, and training

military personnel at or close to energy equilibrium can

reach an AEE/REE level near or above 1.8 with a range of

from 0.8 (PAL¼2.0) to 2.6 (PAL¼4.0) (Figure 2, Table 1, and

Appendix). In contrast, the AEE/REE ratio for representative

humans living in developed nations (B0.5 or PAL¼1.67) is

far lower than that observed in most free-ranging non-

human mammals and active humans.

Using time-energy budget modeling,7 we estimate that to

achieve an AEE/REE of B1.8, present-day humans with a

modern lifestyle (ie, with AEE/REE¼B0.5)7 would need to

spend, in addition to ‘usual’ activities, about 3.5 h per day

running at 14.5 km/h, 3.7 h per day swimming, or 5.7 h

walking over fields and small hills.

This AEE/REE value of B1.8 or PAL of 3.2 can be compared

to PAL levels of sedentary Western populations of approxi-

mately 1.4–1.6.5 The upper limit of sustainable human

metabolic rate, the highest level of energy expenditure at

which energy balance and body weight can be maintained

when food supplies are not limiting is a PAL of 2.2–2.5

for the general population and approximately twice as high,

4–5, in endurance-trained athletes.5

Early hominid evolution took place in woodlands and

grasslands with subsistence hard to obtain. Modern hunter–

gatherers who provide us with a model often travel 6–8

= 800xM

R2 = 0.96

= 253xM

R2 = 0.98

1

10

100

1000

10000

100000

En

erg

y E

xpen

dit

ure

(kJ

/d)

0.001 0.1 10 1000Body Mass (kg)

FEE

FEE

REE

REE

0.73

0.75

Figure 1 Free-ranging and resting energy expenditure (FEE, REE, in kJ/day), vs body mass (M, in kg) in mammals, both expressed as the logarithm. FEE

expenditure was evaluated by the doubly labeled water method and REE by conventional indirect calorimetry. FEE¼800�M0.73, R2¼0.96, Po0.001;

REE¼253�M0.75, R2¼ 0.98, Po0.001. The plotted data and associated references are provided in the Appendix.

Human activity levelsM Hayes et al

152

International Journal of Obesity

mi/day in search of food. Leonard and Robertson8 provided

minimal PAL values using time allocation data for two

groups, the low-activity hunting and gathering !Kung (PAL:

males/females, 1.71/1.51) and the high-activity Ache (2.15/

1.88). The semi-subsistence Upper Volta males and females

had high PAL values in the wet season (2.2/2.1) and lower

values in the less physically demanding dry season (1.5/

1.7).9 Leonard10 and Sorenson and Leonard,11 based upon

skeletal robusticity, estimated that Neandertals had extre-

mely high activity levels with PALs in the range of 2–3.

Maximal oxygen consumption (VO2 max) values provide

additional evidence in support of high physical activity

levels of young adult male modern hunter–gatherers.

Canadian Igloolik Eskimos with a PAL of 2.2 have a VO2

max of 56.4 ml/kg/min, Kalahari San (Bushmen) males

47.1 ml/kg/min, and industrialized Westerners 40.8.12 Ac-

cording to studies cited by Panter-Brick,13 the total energy

expended per kg body mass of typical Americans is 65% of

that of Paleolithic Stone Agers and the average 70 kg modern

human would need to add a 19 km/day (12 mi) walk to reach

a comparable physical activity level. Although time alloca-

tion studies tend to provide PAL estimates lower than those

by doubly labeled water, modern hunter–gatherers evaluated

by this approach thus appear to have physical activity levels

considerably higher than those of modern nonexercising

adult humans living in industrialized nations.

An important limitation of our analysis is the relatively

small literature on mammalian free-living energy expendi-

ture estimated by doubly labeled water. Data in large

mammals are particularly lacking, owing in part to the cost

and complexity of carrying out experiments in free-ranging

animals. Controlling for seasonal influences, age, and other

possible moderators of mammal energy expenditure were

not possible. We used all of the available data, given these

limitations, in our analyses. Our findings should therefore be

seen as an initial approach and analysis rather than as a

definitive examination of early human relative physical

activity levels.

Our findings suggest that a low PAL (ie, AEE/REE¼B0.5 or

PAL of 1.67), much less than that observed in free-ranging

non-human mammals or highly active humans, is present in

modern adult humans living within technologically ad-

vanced settings. These observations provide a quantitative

Table 1 Animal species (Figure 2) and human groups (Figure 3) and

associated references

# Animal Species (references) (Figure 2) # Humans (references) (Figure 3)

1 Pipistrellus pipistrellus (a) 1 Normal wt. women (g)

2 Plectous auritus (a) 2 Normal wt. women (g)

3 Myotis lucifugus (a) 3 Normal wt. men (g)

4 Tarsipes rostratus (b) 4 Normal wt. men (g)

5 Anoura caudifer (b) 5 Obese men (g)

6 Macrotus californicus (a) 6 Obese women (g)

7 Peromyscus crinitus (a) 7 Mexican pima farmers (h)

8 Mus domesticus (a) 8 Gambian female farmers (i)

9 Cleithrionomys rutilus (a) 9 Gambian male farmers (j)

10 Mus musculus (a) 10 Gambian male laborers (k)

11 Sminthropis crassicaudata (a) 11 Active service soldiers (l)

12 Peromyscus maniculatus (a) 12 Marine winter training (m)

13 Perognathus formosus (a) 13 Field training (n)

14 Peromyscus leucopus (a) 14 Arctic training (o)

15 Apodemus sylvaticus (a) 15 Jungle training (p)

16 Antechinus stuartii (b) 16 Female endurance runners (q)

17 Dipodomys merriami (a) 17 Male swimmers (r)

18 Acomys cahirinus (b) 18 Female swimmers (s)

19 Sekeetamys calurus (b) 19 Female mountaineers (t)

20 Microgale dobsoni (b) 20 Male mountaineers (t)

21 Microgale talazaci (b) 21 Females cross country skiers (u)

22 Acomys russatus (a) 22 Males cross country skiers (u)

23 Praomys natalensis (b)

24 Talpa europea (a)

25 Petaurus breviceps (a)

26 Gymnobelideus leadbeateri (a)

27 Psammomys obesus (a)

28 Pseuocheirus peregrinus (a)

29 Vulpes cana (a)

30 Petauroides volans (a)

31 Isoodon obesulus (b)

32 Settonix brachyurus (a)

0

1

2

3

4

5

6

7

1 2 3 4 5 6 7 8 9 10

11

12

13

14

15

16

17

18

19

20

21

22

0

1

2

3

4

5

6

7

1 2 3 4 5 6 7 8 9 10

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

Study Number

Study Number

AE

E /

RE

EA

EE

/ R

EE

Small Free-Ranging Mammals

Humans

Mean, all Free-Ranging Mammals

Non-Athletes

Active Rural

Military

Athletes

Mean

Figure 2 PALs (AEE) in humans (left) and small mammals (right) evaluated

with doubly labeled water, expressed as a multiple of REE. Data are presented

on human groups (nonathletes living in developed nations, military trainees,

individuals from rural areas engaged in high levels of physical activity, and

athletes in training) and small animal species for whom both FEE and REE data

were available as outlined in Table 1. The associated study number references

are summarized in the table and Appendix.


153


estimate of the degree to which modern adult humans may

differ from Paleolithic humans in physical activity and lends

support to the hypothesis that low activity-related energy

expenditure levels contribute to the rising prevalence of

obesity in developed nations.

Acknowledgements

We acknowledge the new primate doubly labeled water data

generously provided by Jeanne and Stuart Altmann of

Princeton University.

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Coward WA, Prentice AM. Effects of dietary supplementa-

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Golan R, Epstein Y. Energy expenditure variations in

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Energy expenditure of swimmers during high volume

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t Westerterp KR, Kayser B, Brouns F, Herry JP, Saris WHM.

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u Sjodin AM, Andersson AB, Hogberg JM, Westerterp KR.

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Appendix A1Free-ranging expenditure was evaluated using a composite

database developed from previously reported and new doubly

labeled water human and animal studies (see Table A1).

Table A1 Field and resting energy expenditure data in mammals

Common name Species Mass (kg) REE (kJ/day) FEE (kJ/day) Appendix ref.

Pipistrelle Pipistrellus pipistrellus 0.0076 5.2 29.3 a

Little brown bat Myotis lucifugus 0.0085 9.6 27.6 a

Brown-long eared bat Plectous auritus 0.0085 6.9 27 a

Northern pygmy gerbil Gerbillus henleyi 0.0093 26.5 b

Honey possum Tarsipes rostratus 0.0099 12.7 34.4 b

Flowering-visiting bat Anoura caudifer 0.0115 17 51.9 a

Big-eared bat Macrotus californicus 0.0129 6.7 22.8 a

Cactus mouse Peromyscus crinitus 0.0134 11.9 40.1 a

Wild house mouse Mus domesticus 0.014 12.1 45.1 a

Bank vole Cleithrionomys rutilus 0.0160 38.1 63.3 a

House mouse Mus musculus 0.0162 12.1 52.5 a

Fat-tailed dunnart Sminthropis crassicaudata 0.0166 9 68.7 a

Deer mouse Peromyscus maniculatus 0.0189 10.6 56.9 a

Long-tailed pocket mouse Perognathus formosus 0.0189 13.4 41.2 a

White-footed deer mouse Peromyscus leucopus 0.0194 16.3 52.5 a

Meadow mouse Microtus arvalis 0.02 90 b

Common wood mouse Apodemus sylvaticus 0.0205 16.9 56.4 a

Big brown bat Eptesicus fuscus 0.0208 45.6 a, ba

Allenby’s gerbil Gerbillus allenbyi 0.0228 35.6 b

Bank vole Cleithrionomys glareolus 0.0234 88 b

Field vole Microtus agrestis 0.0268 77.8 b

Brown antechinus Antechinus stuartii 0.0294 15.6 79.2 a, ba

Greater Egyptian gerbil Gerbillus pyramidum 0.0318 45.2 b

Australian native mouse Pseudomys albocinereus 0.0326 62.2 b

Wambenger Phascogale calura 0.0335 61.9 b

Merriam’s kangaroo rat Dipodomys merriami 0.0359 19.0 52.9 a

Meadow vole Microtus pennsylvanicus 0.0369 115 b

Common spiny mouse Acomys cahirinus 0.0383 22.3 51.8 b

Bushy-tailed jird Sekeetamys calurus 0.0412 22 44 b

Shrew-tenrec Microgale dobsoni 0.0426 20.1 77.1 b

Shrew-tenrec Microgale talazaci 0.0428 21 66.5 b

Golden spiny mouse Acomys russatus 0.045 19.7 47.6 a

Brown lemming Lemmus trimucronatus 0.0552 201 b

Multi-mammate mouse Praomys natalensis 0.0573 15.8 86.6 b

Chisel-tooth kangaroo rat Dipodomys microps 0.0579 101.0 a, ba

Broad-footed marsup. mouse Anetechinus swainsonii 0.0626 150 b

Jird Meriones crassus 0.0692 65 b

Spear-nosed bat Phyllostomus hastatus 0.0808 146 b

Water vole Arvicola terrestris 0.0858 119 b

Antelope ground squirrel Ammospermophilus leucurus 0.087 88 b

Mole Talpa europea 0.0877 29.9 173 a

Eastern chipmunk Tamias striatus 0.0963 143 b

Hamster Cricetus cricetus 0.103 26.1 c

Botta’s pocket gopher Thomomys bottae 0.104 130 b

Sugar glider Petaurus breviceps 0.124 42.8 172.5 a

leadbeater’s possum Gymnobelideus leadbeateri 0.129 49.7 226 a

Least weasel Mustela nivalis 0.15 80.1 c

Wood rat Neotoma albigula 0.15 45.35 c

Fat sand rat Psammomys obesus 0.171 43.6 165.4 a

Golden-mantled ground sqrl. Spermophilus saturatus 0.214 226 b

Golden bandicoot Isoodon auratus 0.333 285 b

Guinea Pig Cavia porcellus 0.41 146.9 d

European Squirrel Sciurus vulgaris 0.44 225.3 c

Arctic ground squirrel Spemophilus parryi 0.63 817 b

Ring-tailed possum Pseudocheirus peregrinus 0.717 192.7 556 a

Civet cat Bassariscus astutus 0.752 472 a

Long-nosed potoroo Potorous tridactylus 0.825 517 b


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Blanford’s fox Vulpes cana 0.945 163.5 604 a

Greater glider Petauroides volans 1.018 206.5 532 a

Short-nosed rat kangaroo Bettongia penicillata 1.1 593 b

Potto Perodicticus potto 1.2 202.6 c

Short-nosed brown bandicoot Isoodon obesulus 1.23 235 644 b

Kit fox Vulpes macrotis 1.48 1180 b

Black-tailed jackrabbit Lepus californicus 1.8 1296 a

Quokka Settonix brachyurus 1.9 310 548 a

Swift fox Vulpes velox 2.045 1634 a, ba

Rufous rat kangaroo Aepyrpimnus rufescens 2.86 1430 b

Echidna Tachyglosssus aculeatus 2.86 875 b

Yellow-bellied marmout Marmota flaviventris 3.19 2430 b

Wildcat Felis sylvestris 3.3 1112 e

Jack rabbit Lepus alleni 3.4 734.4 e

Anteater Tamandua tetradactyla 3.5 423.4 e

Armadillo Cabassous centralis 3.8 394.0 e

Agouti Dasyprocta azarae 3.8 886.5 e

Mangabey Cercocebus torquatus 4.1 814.8 e

Macaque Macaque sp. 4.2 866.1 d

Coypu Myocastor coypus 4.3 1449.0 e

Tammar wallaby Macropus eugenii 4.38 1150 b

Mantled howler monkey Allouatta palliata 4.67 980.7 f

Koala Phascolarctos cinereus 4.77 954 f

Tammar wallaby Macropus eugenii 4.8 672 f

Red-bellied marmout Thylogale billiardieri 5.98 1630 b

Mantled howler monkey Aloutta palliata 7.33 2580 b

Koala Phascolarctos cinereus 7.52 1710 b

Aardwolf Proteles cristatus 8.54 1850 b

Rock wallaby Petrogale xanthopus 8.9 2210 b

Yellow Baboons (female) Papio cynocephalus 1271.4 34007600 n¼ 6 in Kenya

Orangutan Pongo pygmaeus 16 1516.7 c

African wild dog Lyacon pictus 25.17 15300 b

Chimpanzee Pan troglodytes 33 4477.4 c, da

Antarctic fur seal Arctocephalus gazella 34.6 23000 b

Goat Oreamnos americanus 36 3347.2 d

Timber wolf Canis lupus 37.3 17700 b

Mule deer Odocoileus hemionus 39.1 18000 b

Springbok Antidorcas marsupialis 43.3 24100 b

Eastern grey kangaroo Macropus giganteus 44.5 8670 b

Sheep Ovis ammon musimon 46.6 5405.7 d

Northern fur seal Callorhinus ursinus 51.1 36100 b

California sea lion Zalophus californianus 78 38600 b

Australian sea lion Neophoca cinerea 83.5 39500 b

Common seal Phoca vitulina 99 52500 b

Cow Bos taurus 445 28261.5 d

Beef heifers Bos taurus 482 32442.7 d

aMean of combined values for data from more than one source.

Table A1 Continued

Common name Species Mass (kg) REE (kJ/day) FEE (kJ/day) Appendix ref.


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