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An Evolutionary Perspective on Sedentary Behavior

Article in BioEssays · December 2019

DOI: 10.1002/bies.201900156

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An Evolutionary Perspective on Sedentary Behavior

John R. Speakman

diseases attributable to being inactive sug-

Most people are aware of the health benefits of being physically active. The gest that physical inactivity (defined as not
question arises then why people so easily fall into sedentary habits. The idea attaining government guidelines for activ-
developed here is that sedentary behavior is part of a suite of behaviors to ity) is responsible for 6.8% of cardiovas-
reduce levels of physical activity that were strongly selected in the cular disease, 7% of type 2 diabetes, and
10% of breast and colon cancer.[6] Over-
evolutionary past, likely because high levels of physical activity had direct
all physical inactivity was judged to be re-
negative consequences for survival. However, hunter-gatherer populations sponsible for 5.3 million of the 57 million
could not reduce activity indefinitely because of the need to be active to hunt deaths that occurred worldwide in 2008.[6]
for, and gather food. Hence they never experienced low levels of activity that In comparison, smoking was estimated to
are damaging to health, and no corresponding mechanism avoiding low have caused 5 million deaths in 2000.[7] Re-
ducing the population levels of inactivity
activity evolved. Consequently, gene variants promoting efficiency of activity
by 10% would save around half a million
and increased sedentariness were never selected against. Modern society lives annually.[6] The lifespan gains for a per-
facilitates reduced activity by providing many options to become less active son who changes from being inactive to ac-
and divorcing food intake from the need to be active. Choosing the less active tive are estimated at 1.3 to 3.7 years in the
option is hard wired in the genes; this explains why being sedentary is so United States,[8,9] and 2.6 to 4.2 years among
common, and why reversing it is so difficult. Incentivizing activity may be East Asians.[10] There is considerable evi-
dence that the epidemiologically identified
enabled using modern technology, but ultimately may only end up replacing
links are causal.[11]
one set of health issues with others. Also see the video abstract here Although these benefits of being physi-
https://youtu.be/ekHbUwPw-v4. cally active are well known, large segments
of the world population are physically in-
active, not meeting the widely adopted
government guidelines to engage in 150 min of moderate to vig-
1. Introduction–The Problem of Physical Inactivity orous activity per week. The WHO estimated that in 2008, 31% of
the population of individuals aged >15 were insufficiently active.
Hippocrates was so enamored by the benefits of being active that Such data led to the WHO global strategy for physical activity.[4]
he stated “Walking is man’s best medicine.”[1] Being physically Despite such strategies, physical inactivity has continued to rise
active lowers the rates of cardiovascular disease, hypertension, and a recent estimate is that 77.1% of Americans in 2016 did not
stroke, type 2 diabetes, depression, and frailty.[2–4] Conversely, meet the recommended target levels.[12]
sedentary behavior (sitting and lying) increases the risk of coro-
nary heart disease, type 2 diabetes, and both breast and colon
cancer, and hence leads to shortening of lifespan.[5] Estimates of
2. If Physical Activity Is so Beneficial, Why Is
the fraction of the population levels of these non-communicable
Sedentary Behavior so Alluring?
Prof. J. R. Speakman Outside my apartment block there is a small paved path that runs
State Key Laboratory of Molecular Developmental Biology across an area of grass. At the end of the grass, it meets another
Institute of Genetics and Developmental Biology, Chinese Academy of path at an angle of about 90o . People have the choice at the end
Sciences of the first path to turn either left or right. At each side of the
Beijing 100100, China
E-mail: [email protected]
junction, the grass is worn away where people take a shortcut
Prof. J. R. Speakman
across the corner. You can see evidence of the same shortcuts
Institute of Biological and Environmental Sciences being used everywhere across the whole world. I walk past this
University of Aberdeen particular example twice every day and have often wondered why
Aberdeen Scotland AB24 2TZ, UK people do that. Walking across the corner, instead of staying on
Prof. J. R. Speakman the paved section, reduces the path of travel by at most 1 m. The
CAS Center of Excellence in Animal Evolution and Genetics time saving is about 1 s. However, corner cutting is so common
Kunming Institute of Zoology
Kunming Yunnan province, China that it must be hardwired into our biology. Individually cutting
corners results in trivial savings of time and energy, but by do-
The ORCID identification number(s) for the author(s) of this article ing it repeatedly, the savings can start to add up. Imagine I am a
can be found under https://doi.org/10.1002/bies.201900156 hunter-gatherer walking 13–21 000 steps per day.[13] There may
DOI: 10.1002/bies.201900156 be 200 or more occasions on my path where I can be a little more

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efficient by cutting a corner. So now I am walking 200 m less and over the same interval direct measures of physical activity lev-
saving 200 s every day. Over a whole year, that means I have saved els indicate it has decreased.[27] These observations have been ex-
walking 73 km and about 20 h and 15 min of walking time. tended more recently to hunter-gatherer populations, showing
The fact we still cut corners suggests that there was a strong that while HG populations may be more than twice as active
selective pressure for this type of behavior to evolve. This im- as their western counterparts, their total daily energy demands,
plies there is something negative about physical activity that is when corrected for body size, are not different.[26] It has been
selected against. I will develop the argument here that this nat- suggested that this may be because total energy expenditure is
ural selection pressure against physical activity is a major driv- constrained.[28] The fact there is a limit on total energy expendi-
ing factor that causes adoption of sedentary behavior in modern ture is well established.[29–31] Animals working at this limit can-
society. This view contrasts previous considerations of the evolu- not increase expenditure on a component of their energy budget
tion of physical activity which have generally suggested increas- without cutting back on some other aspect of their energy expen-
ing levels of activity to be advantageous and hence endurance was diture. When animals working at this limit become more phys-
strongly selected for.[14–17] The argument here is similar to that ically active, the expended energy on activity may increase, but
by Lieberman,[18] Cheval et al.,[19] and Lee et al.[20] but based on this must be compensated by savings in the budget elsewhere.
a different mechanism. These previous authors have suggested This would explain why levels of activity and total expenditure are
that there is a selection pressure to reduce the total energy that not closely linked. However, the levels of expenditure in hunter-
we expend on physical activity,[18–20] and thus favor behaviors that gatherer and other populations are well below the levels at which
minimize energy expenditure. It is suggested this minimization such limits are suggested to apply,[29–32] so the constraint explana-
was important during our evolutionary history because energy tion is unlikely to be correct. However, the fact that compensation
supplies were limited. In other words, because walking (or run- is observed is indisputable even if we do not fully understand the
ning) costs more energy than being sedentary, by cutting corners mechanism. The argument that we are driven to be inactive to
our activity overall becomes more energy efficient. We visit the save energy[18–20] therefore seems improbable.
same places over the course of a year on our walking travels, but Instead, I suggest there may have been other more direct ad-
it costs us less energy to do so. This would add to the numer- vantages of reducing activity time. The first is the risk of preda-
ous other adaptations we have when walking, such as modulating tion. I have already discussed elsewhere how risk of being pre-
step length,[21] and coordinating our arm movements[22] to mini- dated may have been extremely high in small early hominins
mize energy costs. Corner cutting is evidence that our ancestors like Australopithecus because African populations of predatory
experienced strong selective pressures to reduce activity time, po- cats were much higher and more diverse than presently.[33–35]
tentially to save energy. Taking opportunities to be sedentary as Australopithecine fossil bones often carry features indicative of
opposed to being active is an extension of this phenomenon, and predation. Once our ancestors discovered fire and started to use
a suggested selective pressure for us to be inactive whenever the weapons, the risk of predation probably declined enormously, but
opportunities arise.[18–20] by this time, there had been at least 3 million years of high pre-
How much actual energy would corner cutting save? Sitting dation risk and the die had been cast. This risk would select for
costs about 1.7 kcal min−1 .[23] In contrast, the cost of walk- behaviors minimizing the time spent active because it is active in-
ing is about 3 kcal min−1 ,[24] so each minute walking expends dividuals that are more likely to be predated.[36] Twenty hours less
an additional 1.3 kcal. Over the hypothetical year of corner activity per year from cutting corners is about 1% less total activ-
cutting, the energy saving would amount to 1580 kcal. With ity, but this would translate directly to a 1% reduction in mortality
most vegetables having energy densities in the range from 30– risk when foraging, much more important in terms of selection
50 kcal/100 g saving this amount of energy would reduce the than a 0.1–0.2% energy saving, that might in any case be compen-
annual amount of vegetables needed to be eaten by over 3 kg. sated elsewhere in the energy budget. Increases in sedentary time
For a diet comprising mostly of meat, with an energy density at the expense of activity would bring directly proportional ben-
around 200 kcal/100 g, the weight of saved intake would be lower, efits in survival, even if total energy expenditure was unrelated
but still about 0.8 kg. Although these seem large amounts of en- to such changes. Even if predation risk was dramatically reduced
ergy and food, hunter-gatherer and forager-horticulturalists en- 2 million years ago,[33–35] the avoidance of activity whenever possi-
ergy demands are around 2500–4000 kcal per day,[25,26] hence a ble might be sustained because there was no selective advantage
saving of 1500 kcal by cutting corners over the course of a year is to stop doing it (inertia), but also because there are other nega-
actually only 0.1–0.2% of the total annual energy budget. It seems tive aspects of activity sustaining selection on the trait. For exam-
unlikely that this would be a sufficient selective advantage to pos- ple, although snakes generally do not predate humans, the risk of
itively select for the behavior, particularly since this difference in fatal snake bite is massively increased among walking and run-
the level of available energy is only indirectly linked to the true ning individuals. Moreover, there is the increased risk of injury.
drivers of selection, which are reproduction and survival. Walking, and especially running, entail increased risks in terms
Nevertheless, taking opportunities to be sedentary would po- of ankle sprains and occasionally limb fractures, relative to be-
tentially provide much greater benefits in terms of saved energy ing sedentary. In earlier times, with no health care, such events
than cutting corners would, and so may be selected. However, would also likely be fatal. Hence, the direct mortality risks of ex-
I suggest based on recent evidence that this is unlikely to be the ercise likely provided a much stronger selection pressure against
case. It seems that there is little relationship between the levels of physical activity than their impacts on energy balance.
physical activity (and hence also inactivity) and total expenditure Gene variants favoring efficiency of physical activity (like cor-
of energy. For example, levels of total energy expenditure have ner cutting) and sedentariness were likely advantageous during
remained virtually constant since the 1980s despite evidence that our evolutionary history. This is probably why there is a high

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heritability of physical activity levels[37–39] and inactivity levels.[40] pressure to minimize activity, you could not sit around all day
Gene variants are known that cause reduced activity.[41] This ex- socializing. Long durations of physical inactivity were never ex-
plains why there is such a strong drive to reduce our physical perienced because of the more dominant urge to collect food to
activity whenever we can. This genetic imperative to minimize eat. It seems plausible that our ancestral populations were never
activity is hard wired into our biology. The idea developed above sufficiently inactive that they started to get the diseases we now
is that there are disadvantages of being active that select for genes associate with inactivity,[18–20] which would have selected against
pushing us to be sedentary, but is there also a fitness “pull” of be- this trait. Modern hunter-gatherers have virtually no cardiovas-
ing sedentary as well? One thing that primates and humans do cular disease, for example.[44] So consequently, there was never a
extensively when they are inactive is to socialize with each other. countervailing selection pressure against reducing activity levels
These social behaviors are among the most characteristic features whenever possible.[18–20]
of humans and non-human primates, and in theory, they may The main problem with modern life is that we have come to
provide a fitness pull toward being sedentary that magnifies the the point where we are able to indulge our drive to be sedentary,
fitness push from the mortality risks of activity. I suggest this is to a low level of physical activity not seen previously in our evo-
unlikely. This is because humans do not stop being social when lutionary history. Importantly, in addition, our food intake has
they are active. Sociality among groups of hunters and foragers become disconnected from physical activity levels. We can be in-
were likely extremely important allowing individuals to coopera- active without starving to death. Our ancestors could not. Our
tively hunt animals larger than themselves, butcher them in the genetic legacy pushes us toward inactivity at every opportunity,
field, and carry the pieces back to camp. This fitness benefits of and we succumb almost every time that opportunity arises. This
sociality probably therefore occur independently of whether indi- explains why large sectors of industrialized society have levels of
viduals are active or sedentary, and the difference in fitness due activity that are damaging to health (reviewed above). Our Aus-
to sociality between the states would be unlikely to provide a suf- tralopithecine genetics never prepared us for this eventuality, and
ficient fitness pull toward being sedentary. worse they help us to resist any initiative to reverse the trend.
How might these ideas be tested? Testing evolutionary argu-
ments is notoriously difficult because we cannot go back and re-
run evolution with a different set of selective pressures to see if 4. What Can be Done?
the outcome differs. The best we can hope then is to test the un-
derlying assumptions on which any given selection scenario is Because the desire to be sedentary is in our genes, it will be diffi-
based. A primary element of the argument I have developed is cult to reverse. Moreover, the history of public health science tells
that there is a fitness push toward being sedentary because of the us repeatedly that giving people information that their behavior
mortality risks encountered during periods of activity. A critical is damaging does little to motivate behavior change. The simplest
test of the idea then would be to measure directly such mortality way to reverse this trend would be to eliminate the possibilities
risks. If it was shown that being active did not bring significant to avoid activity. Recently, both elevators in my office building
risks of mortality above being sedentary that would be enough to were simultaneously broken. There was no option but to take the
disprove the hypothesis. stairs. If nothing else, this showed that people can do it if they are
forced to. Generally, initiatives to improve physical activity levels
will be most successful if they include removing the option to
3. Activity in the Modern World choose a more sedentary alternative. Grass only regrows at the
corners of intersecting pathways if the ability to walk across the
What implications does the fitness push to inactivity have for corner is restricted by a barrier. Lieberman makes similar sugges-
modern humans? I suggest it has a major impact. In the modern tions, but additionally suggests exercise might be encouraged if
world, we still cut corners and select the least expensive ways of we made it more fun.[18] This may work for sporting activity, but
getting around. We choose the moving walkway rather than walk- I am less sure that fun will be a sufficient motivator to overcome
ing. We order our shopping online rather than going to the super- our deep genetic history for more mundane activities. There is
market. We order takeaway instead of going to the restaurant or only so much fun that can be derived from walking up stairs.
cooking ourselves. The building where I work has six stories that Removing options to be sedentary will be difficult. Remov-
are accessed via two elevators and a staircase. Last week for a cou- ing elevators, thus forcing people to take the stairs, for example,
ple of days, I sat by the ground floor elevators and counted people would likely increase physical activity levels, but clearly also have
returning from lunch who used the stairs and who used the el- some undesired side effects. Elevators facilitate the movement
evators. One hundred and fifty seven people came by, of which of people who are disabled; moreover, they are also used to trans-
only two used the stairs. We make these choices even though we port things, other than people. Modern technology, however, may
know the options we do not choose are healthier. enable solutions to this problem. It would be relatively easy, for
Physical inactivity is bad for our health, so it is reasonable example, to issue people with disabilities who need to use ele-
to ask why there was not a counter-selection pressure against vators with swipe cards permitting them to do so, while blocking
being sedentary. The reason for this it seems is that to exist out individuals that do not have such needs. Many hotel elevators
as a hunter gatherer you need to do a fair amount of activity already have such technology to restrict individual access to guest
to collect food. There are few true hunger-gatherer populations room levels. This could be further enabled if the population us-
left, but those that have been studied walk a lot—about 6 h and ing the elevator was known, by using face recognition technology,
20 km per day.[12,42–44] It seems this level of activity is necessary thereby eliminating the need for physical swipe cards, and issues
to collect enough food.[18,45] So, while there was always a selective if people lost or forgot their cards. I recently visited a company in

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South China where using the elevators was simply not permitted Received: September 1, 2019
by staff with the stated goal of promoting their physical activity Revised: October 30, 2019
levels. I did not find out what the sanctions were for breaking the Published online: December 12, 2019
rules. As technology develops, one might envisage all sorts of sce-
narios where we could be incentivized to engage in activity. For
example, many people now carry smartphones that have integral [1] Hippocrates quotes, http://wwwbrainyquotecom/quotes/authors/
accelerometers that log physical activity. The advent of big data h/hippocrateshtml 2011 (accessed: August 2019).
means that conceivably such activity could be centrally accumu- [2] U.S. Department of Health & Human Services, Physical Activity
lated and people reaching certain activity levels could be given Guidelines Advisory Committee Report, U. S. Department of Health and
tax breaks because they would cost less money to the health ser- Human Services, Washington, DC 2008.
vice. One would need to have safeguards, for example, to prevent [3] D. E. Warburton, S. Charlesworth, A. Ivey, L. Nettlefold, S. S. Bredin,
people attaching their phones to their dogs to reach the tax break Int. J. Behav. Nutr. Phys. Activity 2010, 7, 39.
threshold, but that could be easily enabled by having a small im- [4] World Health Organization, Global Recommendations on Physical Ac-
tivity for Health, WHO, Geneva, Switzerland 2010.
planted ID chip linked to the external device.
[5] F. W. Booth, C. K. Roberts, M. J. Laye, Compr. Physiol. 2012, 2, 1143.
Sooner or later, however, if we force or incentivize large num- [6] I. M. Lee, E. J. Shiroma, F. Lobelo, P. Puska, S. N. Blair, P. T. Katz-
bers of individuals to take the stairs or walk the streets, some marzyk, Lancet physical activity working group, Lancet 2012, 380, 219.
of them will fall and get injured, or be hit by a car (predated), [7] M. Ezzati, A. D. Lopez, Lancet 2003, 362, 847.
and then we will be reminded why we are selected to be inac- [8] O. H. Franco, C. de Laet, A. Peeters, J. Jonker, J. Mackenbach, W.
tive. In the United States, for example, there are 2.3 deaths per Nusselder, Arch. Intern. Med. 2005, 165, 2355.
million residents per annum due to cars hitting cyclists.[46] If we [9] R. S. Paffenbarger Jr, R. T. Hyde, A. L. Wing, C. C. Hsieh, N. Engl. J.
increased the amount of cycling by a factor of 5 that would po- Med. 1986, 314, 605.
tentially lead to an extra 3600 activity-related deaths per year. The [10] C. P. Wen, J. P. M. Wai, M. K. Tsai, Y. C. Yang, T. Y. D. Cheng, M.-C.
question is will the frequency of such negative outcomes at the Lee, H. T. Chan, C. K. Tsao, S. P. Tsai, X. Wu, Lancet 2011, 378, 1244.
[11] F. W. Booth, C. K. Roberts, J. P. Thyfault, G. N. Ruegsegger, R. G.
population level offset the benefits of doing the extra physical
Toedebusch, Physiol. Rev. 2017, 97, 1351.
activity? [12] D. L. Blackwell, and T. C. Clarke, National Health Statistics Reports
Number 112 n June 28, 2018 State Variation in Meeting the 2008
Federal Guidelines for Both Aerobic and Muscle-strengthening Ac-
tivities Through Leisure-time Physical Activity Among Adults Aged
5. Conclusion 1864: United States, 2018, 2010–2015.
Reducing activity levels whenever possible has probably been un- [13] J. H. O’Keefe, R. Vogel, C. J. Lavie, L. Cordain, Am. J. Med. 2010, 123,
der strong selection during our evolutionary history. However, 1082.
[14] A. F. Bennett, J. A. Ruben, Science 1979, 206, 649.
the drive to eat which required us to collect food and hunt meant
[15] U. Wisloff, S. M. Najjar, O. Ellingsen, P. M. Haram, S. Swoap, Q. Al-
we never experienced low levels of activity. Hence there were
Share, M. Fernstrom, K. Rezaei, S. J. Lee, L. G. Koch, S. L. Britton,
never the conditions for a countervailing selection pressure to Science 2005, 307, 418.
avoid low activity levels that we now know are damaging to health. [16] S. B. Eaton, S. B. Eaton, Comp. Biochem. Physiol., Part A: Mol. Integr.
We consequently take whatever opportunities that present them- Physiol. 2003, 136, 153.
selves to reduce our physical activity, with critically no lower limit [17] J. H. O.Keefe, R. Vogel, C. J. Lavie, L. Cordain, Prog. Cardiovasc. Dis.
to this imperative. Modern industrialized society has largely fa- 2011, 53, 471.
cilitated this behavior change by continuously presenting us with [18] D. E. Lieberman, Curr. Sports Med. Rep. 2015, 14, 313.
options to reduce our activity and disconnecting the need to eat [19] H. H. Lee, J. A. Emerson, D. M. Williams, Front Psychol. 2016, 7,
from the need to be active. Attempts to encourage us to choose 1285.
[20] B. Cheval, R. Radel, J. L. Neva, L. A. Boyd, S. P. Swinnen, D. Sander,
the option requiring more activity are largely futile in the face of
M. P. Boisgontier, Sports Med. 2018, 48, 1389.
our evolutionary history and genetics. Probably, the only way to
[21] M. Y. Zarrugh, F. N. Todd, H. J. Ralston, Eur. J. Appl. Physiol Occup.
successfully increase activity will be to remove the low-activity al- Physiol. 1974, 33, 293.
ternatives, but this will often have other negative consequences. [22] S. H. Collins, P. G. Adamczyk, A. D. Kuo, Proc. Biol. Sci. 2009, 276,
Modern smart technology may enable solutions to these issues, 3679.
but in the end forcing people to be active may expose them to a [23] C. J. Fountaine, J. Johann, C. Skalko, Int J Exerc. Sci 2016, 9, 223.
different set of health issues. [24] D. P. Laroche, N. R. Marques, H. N. Shumila, C. R. Logan, R. S. Lau-
rent, M. Goncalves, Med. Sci. Sports Exerc. 2015, 47, 1017.
[25] L. Christopher, F. C. Madimenos, R. G. Bribiescas, S. S. Urlacher, J.
J. Snodgras, L. S. Sugiyama, H. Pontzer, Am. J. Hum. Biol. 2019, 31,
Conflict of Interest e23223.
[26] H. Pontzer, D. A. Raichlen, B. M. Wood, A. Z. P. Mabulla, S. B. Racette,
The author declares no conflict of interest.
F. W. Marlowe, PLoS One 2012, 7, e40503.
[27] K. R. Westerterp, J. R. Speakman, Int. J. Obes. 2008, 32, 1256.
[28] H. Pontzer, R. Durazo-Arvizu, L. R. Dugas, J. Plange-Rhule, P. Bovet,
Keywords T. E. Forrester, E. V. Lambert, R. S. Cooper, D. A. Schoeller, A. Luke,
Curr. Biol. 2016, 26, 410.
evolution, genetics, internet of things, physical inactivity, sedentary behav- [29] R. Drent, S. Daan, Ardea. 1980, 68, 225.
ior, smart technology [30] K. A. Hammond, J. Diamond, Nature 1997, 386, 457.

BioEssays 2020, 42, 1900156 1900156 (4 of 5) © 2019 WILEY Periodicals, Inc.

www.advancedsciencenews.com www.bioessays-journal.com

[31] J. R. Speakman, Adv. Ecol. Res. 2000, 30, 177. T. D. Spector, N. J. Wareham, R. J. Loos, Am. J. Clin. Nutr. 2013, 98,
[32] C. Thurber, L. R. Dugas, C. Ocobock, B. Carlson, J. R. Speakman, H. 1317.
Pontzer, Sci. Adv. 2019, 5, eaaw0341. [41] Z. Zhang, C. J. Hao, C. G. Li, J. Zhao, X. N. Li, A. H. Wei, Z. B. Wei, L.
[33] J. R. Speakman, Cell Metab. 2007, 6, 5. Yang, X. He, X. C. Zhen, X. Gao, J. R. Speakman, W. Li, PLoS Genet.
[34] J. R. Speakman, Int. J. Obes. 2008, 32, 1611. 2014, 10, e1004124.
[35] J. R. Speakman, J. Exp. Biol. 2017, 221, jeb167254. [42] F. W. Marlowe, The Hadza: Hunter-Gatherers of Tanzania, University
[36] A. I. Houston, J. McNamara, Ornis Scand. 1993, 24, 205. of California Press, Berkeley, CA 2010, p. 325
[37] J. H. Stubbe, D. I. Boomsma, J. M. Vink, B. K. Cornes, N. G. Martin, [43] H. Pontzer, B. M. Wood, D. A. Raichlen, Obes. Rev. 2018, 19, 24.
A. Skytthe, K. O. Kyvik, R. J. Rose, U. M. Kujala, J. Kaprio, J. R. Harris, [44] D. A. Raichlen, H. Pontzer, J. A. Harris, A. Z. P. Mabulla, F. W. Mar-
N. L. Pedersen, J. Hunkin, T. D. Spector, E. J. de Geus, PLoS One 2006, lowe, J. J. Snodgras, G. Eick, J. C. Berbesque, A. Sancillo, B. M. Wood,
1, e22. Am. J. Hum. Biol. 2017, 29, e22919.
[38] X. Y. Zhang, J. R. Speakman, Endocrinology 2019, 160, 840. [45] S. B. Eaton, M. Konner, M. Shostak, Am. J. Med. 1988, 84, 739.
[39] N. F. Butte, G. Cai, S. A. Cole, A. G. Comuzzie, Am. J. Clin. Nutr. 2006, [46] www.governing.com/gov-data/transportatio-infrastricture/most-
84, 646. bicycle-cyclist-deaths-per-capita-by-sate.html (accessed: August
[40] M. den Hoed, S. Brage, J. H. Zhao, K. Westgate, A. Nessa, U. Ekelund, 2019).

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