Sports

SPRINGER BRIEFS IN ANTHROPOLOGY
HUMAN BEHAVIOR, BIOLOGY AND E VOLUTION
Ann E. Caldwell
Human Physical
Fitness and Activity
An Evolutionary
and Life History
Perspective
SpringerBriefs in Anthropology
Human Behavior, Biology and Evolution
Series editor
Jane Lancaster, Department of Anthropology, University of New Mexico,
Albuquerque, New Mexico, USA
More information about this series at http://www.springer.com/series/15166
Ann E. Caldwell
Human Physical Fitness

and Activity
An Evolutionary and Life History Perspective
123
Ann E. Caldwell
Anschutz Health and Wellness Center
University of Colorado Denver, School
of Medicine
Aurora, CO
USA
ISSN 2195-0806 ISSN 2195-0814 (electronic)

SpringerBriefs in Anthropology
ISSN 2366-8792 ISSN 2366-8806 (electronic)
Human Behavior, Biology and Evolution
ISBN 978-3-319-30407-6 ISBN 978-3-319-30409-0 (eBook)
DOI 10.1007/978-3-319-30409-0
Library of Congress Control Number: 2016934017
© The Author(s) 2016

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To my Josephine
Series Preface
Just a few years after the 150 anniversary of the publication of “Origins of the
Species” and official Year of Darwin, the social and biological sciences are still
working to integrate their research into a more complete understanding of human
behavior and biology. This series hopes to close this gap as demonstrated by the
potential topics and their audiences: anthropologists, biologists, demographers,
economists, psychologists, geneticists, public health researchers, and others. This
series will closely model the aims and scope of the journal Human Nature. It will
focus on human evolutionary ecology, biology, psychology and behavior. The
Briefs are based on evolutionary and life history theory. The social, behavioral, and
biological sciences have traditionally pursued separate paths both in research and in
education. Yet it is increasingly apparent that the most pervasive and crucial issues
of our time are fundamentally behavioral in their nature and lie squarely at the
interface of these artificially separated disciplines.
To address the problems of modern society, we must first understand the very
substance of our species: how we evolved to be the way we are, why we behave the
way we do, and what are the social and ecological contexts that restrain, channel,
modify, and diversify our behavior. To this end, this SpringerBrief series will look
though a complex, multi-faceted lens: one that peers through time, across geo-
graphic space and into the diversity of human social and cultural experience. It will
emphasize the integration of biological, environmental, social, psychological and
behavioral factors that shape the expression of human behavior and the relevance of
a biosocial perspective to major scientific, social, and policy issues in the world
today.
The first book in the series, Human Physical Fitness and Activity: An
Evolutionary and Life History Perspective by Ann E Caldwell provides us with a
model for applying evolutionary and life history theory to understand the rela-
tionship of exercise with health and disease in modern, sedentary populations.
Natural selection has shaped human physiology and psychology to flexibly respond
to energetic demands and resource scarcity in order to optimize survival and
reproductive fitness. The one thing natural selection has not been able to do is to
vii
viii Series Preface
shape human biology to operate in an environment with an unlimited supply of

surplus energy with the only physiological tools at hand being a biology that readily
stores surplus energy in fat. Evolutionary theory tells us we should be thinking in
terms of how costs and benefits shaped by a world long gone operate in today’s
world. Life History Theory warns us to be cognizant of the fact that the costs and
benefits of energy expenditure and storage vary by sex, age and circumstance so
that approaches to increase physical activity can and must be tailored by altering the
built environment and the reward structure from a life history perspective.
Jane Lancaster
Acknowledgements
There are several people who contributed significantly this book. Namely, Jane
Lancaster, Angela Bryan, and Steve Gangestad, who served on my comprehensive
exam committee when I was forming many of the ideas presented here. The
application of life history theory to physical activity has also been shaped
immeasurably by Paul Hooper and Hilly Kaplan. Mel Konner, Herman Pontzer, and
Paul Hooper reviewed previous versions of the manuscript and provided essential
feedback and suggestions that significantly improved the research included, and
presentation of these ideas. In addition, my work has been critically shaped by my
experiences doing fieldwork among the Tsimane’. This book would not be possible
without the work of Hilly Kaplan, Mike Gurven, and the entire Tsimane’ Life
History Project, including Ben Trumble, Dan Cummings, Helen Davis, Adrian
Jaeggi, Chris von Rueden, and Matthew Schwartz, and the incredibly understanding
Tsimane’ people. I also want to thank all those who have studied and written about
this topic before me, cited within; without their insight and foundation, this book
would not have been possible. I would like to express my gratitude to all the people
who saw me through the writing of this book and provided support and encour-
agement, editing, proof reading, and/or listened to me talk about this topic end-
lessly. You know who you are.
ix
Contents
Part I Evolution and Life History Theory

1 Advantages of Evolutionary Theory for Understanding
Physical Activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Categories of Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Proximate and Ultimate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Phylogeny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2 Physical Activity and Life History Theory . . . . . . . . . . . . . . . . . . . 11

Energetic Costs of Physical Activity. . . . . . . . . . . . . . . . . . . . . . . . . 12
Benefits of Physical Activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Summary and Applications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3 Phylogeny and Life History Patterns . . . . . . . . . . . . . . . . . . . . . . . 19

Human Morphological, Physiological, and Metabolic Adaptations . . . . 19
Human Uniqueness in Life History Patterns . . . . . . . . . . . . . . . . . . . 22
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
4 Physical Activity and Energy Expenditure in Humans . . . . . . . . . . 27

Measurement. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
TEE in Humans and Other Primates. . . . . . . . . . . . . . . . . . . . . . . . . 30
Physical Activity in Humans over Time and Across Populations . . . . . 30
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
xi
xii Contents
Part II Physical Activity and Energetic Trade-Offs Through

the Lifespan
5 Energy Costs and Benefits During Fetal Development
and Infancy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
6 Energetic Trade-Offs and Physical Activity During

Childhood and Adolescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
7 Physical Activity and Reproductive Ecology in Adults . . . . . . . . . . 53

Women . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Men . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Post-reproductive Age Men and Women . . . . . . . . . . . . . . . . . . . . . 60
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Part III Summary of Proximate Mechanisms, an Integrated

Evolutionary Model and Applications
8 Proximate Mechanisms: Psychology, Neuroendocrine System,
and Central Nervous System. . . . . . . . . . . . . . . . . . . . . . . . . . ... 65
Interpreting Previous Findings: Why do Psychological
Variables Matter Sometimes but not Others? . . . . . . . . . . . . . . . . . . . 65
Neuroendocrine System as Potential Mechanism . . . . . . . . . . . . . . . . 66
Hypothalamo-Pituitary-Gonadal (HPG) Axis . . . . . . . . . . . . . . . . . 67
Hypothalamo-Pituitary-Adrenal (HPA) Axis . . . . . . . . . . . . . . . . . 71
Central Nervous System and Epigenetics as Potential Mechanisms. . . . 73
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
9 Summary, Conclusions, and Applications . . . . . . . . . . . . . . . . . . . 81

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
About the Author
Ann E. Caldwell received her Ph.D. in evolutionary psychology with distinction

from the University of New Mexico in 2013. Her areas of research also include
evolutionary anthropology and health psychology. She held a postdoctoral research
position in the Laboratory for Comparative Human Biology in the Department of
Anthropology at Emory University from 2014–2015. She is currently a
Postdoctoral Fellow at the Anschutz Health and Wellness Center at the University
of Colorado.
xiii
Introduction
The science of human physical fitness and activity is ripe for a novel theoretical
framework that can integrate the ecological, physiological and psychological factors
that influence physical activity.1 Sedentary behavior is the fourth leading cause of death
worldwide (Kohl et al. 2012; WHO 2008), contributing to an estimated 5.3 million
deaths per year (9 % of premature mortality; Lee et al. 2012). Physical activity is
protective against numerous avoidable, chronic, and deadly conditions: cardiovascular
and metabolic diseases (type 2 diabetes, stroke, heart disease), breast and prostate
cancer, osteoporosis, stress, anxiety and depression (F.W. Booth et al. 2012; Warburton
et al. 2006). Exercise also improves cognitive function and academic performance in
children and adults (Hillman et al. 2008). Despite the wide array of physical and mental
health benefits associated with physical activity, the vast majority of people in the US
(45 % of kids and <5 % of adults when measured objectively; Troiano et al. 2008) do
not get the recommended levels of activity for achieving these benefits.
Current approaches for understanding the psychology of physical activity among
largely sedentary, modern populations are only modestly effective in increasing
levels of physical activity (Spence and Lee 2003). These approaches rely primarily
on domain-general social-cognitive models of health behavior that, as of yet, have
not led to effective exercise interventions that initiate and maintain successful
behavior change. Despite efforts to explain exercise behavior, much remains
unclear about the determinants of exercise participation and adherence (Trost et al.
2001). Health and exercise psychologists generally adopt models of health behavior
such as the Health Belief Model (Rosenstock 1974), Social Cognitive Theory
(Bandura 1977), or the Theory of Planned Behavior (Ajzen 1991), that measure
1
In many cases it is important to distinguish between exercise and obligatory or habitual physical
activity, with exercise being a subset of physical activity explicitly designed to increase physical
fitness. For the purposes of this book, we will be comparing across human history and across
different groups across the world, and therefore will at times discuss both exercise and habitual
physical activity in the same context. For the purposes of this book, we will therefore use a
traditional definition of physical activity to mean, “any bodily movement produced by skeletal
muscles that results in energy expenditure” (Caspersen et al. 1995 p. 126).
xv
xvi Introduction
cognitive correlates of health behaviors. These models are limited in two crucial
ways: (1) the same models are applied across a wide range of behaviors—including
flossing teeth, wearing a seatbelt, participating in mammography screening, and
reducing alcohol consumption, smoking, and unhealthy eating—without taking
behavior-specific aspects of motivation and adherence into account (M.L. Booth
et al. 2000; Mirotznik et al. 1995); and (2) they focus almost exclusively on
rational, cognitive processes driving behavior. These drawbacks are particularly
problematic for behaviors like physical activity or exercise that are influenced by
the integration of multiple physiological systems (i.e., endocrinology, nervous
systems, and metabolism), in addition to explicit and implicit cognitive processes
and environmental factors. Several researchers have advocated more integrative
approaches that take into account evolutionary history (Astrand 1994; Biddle and
Mutrie 2008), or that consider the connections between ecological, genetic, and
physiological factors that influence physical activity (Bryan et al. 2007; Nigg et al.
2008; Spence and Lee 2003). However, a comprehensive evolutionary framework
has yet to be put forward.
Over the past two decades researchers across disciplines have begun to think
about the lifestyle factors that increasingly threaten health and well-being in light of
evolution. In terms of physical activity, this has meant understanding the ways
natural selection has favored high levels of physical activity in humans. First, we
can think about the mismatch or discordance between the energy expenditure
required to survive in current environments compared to those humans faced for the
vast majority of evolutionary history (Cordain et al. 1998; Eaton et al. 1988).
Nutrient dense foods are now abundant and energetically very easy to acquire.
Ancestrally, however, food was not always available, and was rarely easy to come
by. Furthermore, humans have several anatomical and physiological adaptations
that are unique among primates and suggest that endurance running was an
important part of human evolution (Bramble and Leiberman 2004). For example,
humans have springlike tendons and arches in the feet, collagen-rich tendons and
ligaments that effectively store and release elastic strain energy, long legs, short
toes, and other traits that help stabilize and correctly move our upright torso and
head, aid in thermoregulation, and reduce the stress of running. Not only is
endurance running possible in humans because of these adaptations, it also involves
neurobiological rewards (Raichlen et al. 2012). Recently, researchers have sug-
gested that endurance running (and an increased aerobic capacity) in humans also
played a role in the evolution of our uniquely large brains (Raichlen and Polk
2013).
Comparisons of estimated energy expenditure between humans living in primarily
sedentary Western settings with modern subsistence-level populations have been
drawn in the following way. On average, an office worker in the US expends
approximately 9 kilocalories per kilogram of body weight per day (kcal/kg/d). In
contrast, among modern subsistence populations, a !Kung man expends 20 kcal/kg/d,
and a !Kung woman 14.5 kcal/kg/d. Ache men expend 30 kcal/kg/d and women
Introduction xvii
15 kcal/kg/d (Cordain et al. 1998).2 Based on fossil evidence, early H. sapiens have
been estimated to have expended 22 kcal/kg/d, H. erectus, 23 kcal/kg/d, and
H. habilis, 21 kcal/kg/d (Leonard and Robertson 1997).
The view that human physiology has evolved to have high levels of physical
activity helps explain why low levels of physical activity lead to deleterious health
outcomes, and why physical activity has so many mental and physical health
benefits. A large body of evidence supports the claim that human physiology is not
designed to function properly under conditions of unlimited caloric intake and low
physical activity (Booth et al. 2012; Booth et al. 2002a, b; Chakravarthy and Booth
2004). But this view is sometimes taken to imply that natural selection has only
shaped human physiology and psychology to be driven to achieve high levels of
physical activity and exercise. It is as if there is some critical unknown factor that
made our ancestors so active, which we have somehow lost in recent history. If
scientists could simply rediscover this critical factor, it would make us all want to
be as active as the !Kung or Ache. It could also be interpreted as suggesting that
there is some ideal Paleo Exercise program that would prescribe the proper intensity
and duration of exercise, appeal to humans everywhere, and restore health and well-
being around the world. Unfortunately, these expectations are unlikely to be ful-
filled in such a simple manner. However, there is a silver lining. Applying evo-
lutionary theory does allow us to uncover many missing pieces of the puzzle to
understand physical activity, and to change current patterns for greater health and
well-being.
In this book, I summarize and extend the work that has been done thus far to
understand the ways natural selection has shaped physical activity in humans. I
propose that while humans are adapted to be physically active in many ways,
natural selection has shaped a flexible system that responds to the environmental
and individual costs and benefits to allocate a finite energetic budget across the
lifespan. This nuanced application of evolutionary theory leads to a more sophis-
ticated understanding of the circumstances under which natural selection favors
both sedentary and active behaviors in predictable ways.
Human patterns of physical activity have changed drastically over a relatively short
amount of evolutionary time. Over the past 10,000 years humans have created
innovative technological advances that have made resource extraction and storage
exponentially more efficient, resulting in a dramatic shift in the level of physical
2
It should be noted that these estimates of energy expenditure are obtained through the factorial
method (based on behavioral observations and estimated metabolic exertion of observed behav-
iors), which does not accurately estimate overall energy expenditure compared to more objective
measures (more on this in Chap. 4). More recently, researchers have compared objectively mea-
sured energy expenditure in the Hadza in Tanzania and in Western populations, and did not find
significant differences in total energy expenditure, despite significantly different lifestyles (Pontzer
et al. 2012). Much remains unclear about differences in objectively measured physical activity
across populations and ecologies. These estimates are also problematic because they are given
relative to body size, however there is a non-linear relationship between body mass and energy
expenditure, thus these estimates will be influenced by body size with smaller individuals having
higher ratios than larger individuals.
xviii Introduction
energy expenditure necessary to fulfill basic biological needs. This level of innovation
is unprecedented in the animal kingdom, but the impetus to increase efficiency is a
fundamental feature of life, universal to all organisms. This feature is a central pre-
mise of Life History Theory, a body of evolutionary thought on which this book
draws on heavily (see Chap. 2). The theory leverages the observation that all
organisms face finite energetic budgets, and that energy must therefore be strategically
allocated across the lifespan. Mechanisms that determine how energy is allocated
have been shaped by natural selection over millions of years in order to maximize the
probability of reproductive success. How energy is allocated varies across species,
across environments, and across individuals within species, and is fundamentally
linked to energy expenditure in the form of physical activity and exercise.
The first goal of this book is to present a novel theoretical perspective that
highlights domain-specific aspects of the evolved psychology and physiology of
energy expenditure, and that can lead to a more integrated and complete under-
standing of physical activity across the lifespan. This perspective is described in
detail in Part I. The second goal of this book is to synthesize and interpret cross-
disciplinary research that can illuminate original approaches to increase physical
activity in modern, primarily sedentary contexts. This includes a breakdown of the
human lifespan in Part II, and a discussion of the predicted costs and benefits of
physical activity at each stage of life. Knowing that many obstacles to physical
activity and exercise are functionally adaptive—or were in the environments that
they evolved—and identifying which factors are more modifiable than others can
help us to develop interventions and environments that are more conducive to
physical activity. An evolutionary framework also helps illuminate the salient
underlying motivations for physical activity that can be leveraged in new and
creative ways to tip the costs and benefits toward increasing physical activity.
It can be difficult to synthesize all the factors we know to be important for
influencing complex behaviors like physical activity. We know that genes, envi-
ronments, physiology and psychology all work together to influence physical
activity and exercise. How can we integrate across these different levels to
understand behavior? Part III presents a model that integrates the relevant predictors
of physical activity from the theoretical perspective introduced in Parts I and II.
This final Part also presents future directions for research and applications for
public health.
Throughout the book I will necessarily use the term fitness in two distinct ways.
When discussing fitness in terms of physical prowess, including cardiorespiratory
and muscular endurance, muscle strength, flexibility, balance, agility, and body
composition, I will use the term physical fitness. On the other hand, when dis-
cussing fitness in an evolutionary sense, it refers to the relative success of passing
genes on to a subsequent generation. In these instances, I will use the terms
reproductive or Darwinian fitness.
This book draws on data from contemporary small-scale societies, where
quantitative fieldwork has characterized multiple dimensions of behavior and
biology in populations living outside typical industrial economic systems. It draws
particularly heavily on data from the Tsimane’, an indigenous population in the
Introduction xix
Bolivian lowlands with a subsistence economy based on foraging and horticulture,

with whom I have had the honor of conducting fieldwork. Carrying out research
with members of small-scale societies is informative because, generally speaking,
they live in conditions much more similar to those faced by humans for the majority
of human history. In addition, data from different field settings provide stark
environmental contrasts that can highlight the ways environmental factors influence
behavior, which are often consistent with predictions derived from evolutionary
theory. When taken together with studies of physical activity in contemporary
Western populations, we can develop a more comprehensive understanding of the
environmental and evolved psychological and physiological factors that influence
physical activity and inactivity in humans. This comprehensive knowledge can be
applied in modern settings to increase physical activity and decrease the burden of
poor health outcomes that result from inactivity.
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Part I
Evolution and Life History Theory
Chapter 1
Advantages of Evolutionary Theory
for Understanding Physical Activity
Examining physical activity from an evolutionary perspective can lead to novel

hypotheses or ways of integrating evidence across different levels of analyses that
would not naturally be integrated from existing theoretical perspectives. This
approach is particularly informative for traits or behaviors such as physical activity
that are influenced by numerous environmental, psychological, and physiological
factors that have likely been shaped by natural selection over time. For a trait or
behavior to have been shaped by natural selection, it must satisfy three criteria. It
must have (1) variation1 (2) that is at least partly heritable and which also
(3) influences reproductive success. For physical activity, this would mean that
there must be variation in how active people are and that tendencies to be more or
less active are inherited and influence reproductive opportunities and outcomes. In
this chapter, I will discuss the four categories of analysis that an evolutionary
perspective provides and highlight the ways this differs from traditional approaches
from health psychology and public health.
Categories of Analysis
In 1963, evolutionary biologist Nikolaas Tinbergen specified four complementary

categories of analysis to explain a behavior: proximate (or mechanistic), ultimate
(or functional), phylogenetic, and developmental (Tinbergen 1963). Integrating
1
Traits can also not currently have variability, but have had variability at a certain point histori-
cally, until an optimal solution for all individuals arose and stabilized in the population. That there
is still variation in physical activity and underlying psychological motivation, and psychological
and physiological responses to physical activity suggests that there is not one solution that is
optimal for everyone and that selection is still shaping this behavior.
© The Author(s) 2016 3

A.E. Caldwell, Human Physical Fitness and Activity,
Human Behavior, Biology and Evolution, DOI 10.1007/978-3-319-30409-0_1
4 1 Advantages of Evolutionary Theory …
what we can learn from each category leads to a more thorough understanding of a
trait or behavior. Each category is summarized here and further detailed in subse-
quent chapters.
Proximate and Ultimate
Traditional approaches to understand physical activity in health psychology or

public health tend to only consider proximate mechanisms related to physical
activity. Hypotheses about what factors predict physical activity are often generated
by looking at patterns in empirical data, or from theoretical approaches that lump all
health behaviors together and assume that the cognitive process that underlies all
health behaviors is the same. An evolutionary approach contrasts with these tra-
ditional approaches in several important ways. First, evolutionary theory considers
that proximate predictors are directly or indirectly linked to ultimate/functional
predictors. An analysis that considers an ultimate explanation for a behavior seeks
to understand how the behavior or the underlying psychological and physiological
mechanisms that produce the behavior were and are shaped by natural selection.
Evolutionary theory can therefore provide a framework for generating hy-
potheses about proximate mechanisms that influence health behavior that may be
overlooked by traditional approaches, such as considering the costs and benefits of
performing physical activity as they relate to reproductive fitness (more on this in
Chap. 2). It can also lead to hypotheses about why or when certain proximate
predictors of physical activity are more or less likely to influence behavior at a
given time or in a given environment. This can provide a more comprehensive
understanding of predictors that have been shown to influence physical activity
inconsistently, or those that seem to influence behavior in one environment or
population, but not others. This framework can also guide the development of
interventions to focus on more malleable traits or psychological processes, or
encourage change in ways that are more consistent with evolved psychological and
physiological mechanisms. Including an ultimate or functional level of explanation
has advantages, but requires careful consideration of how natural selection operates,
as well as an understanding of how psychological adaptations are designed.
Natural selection operates through differential reproductive success. Those who
reproduce more have greater reproductive fitness, because their genes are passed on
to a subsequent generation at higher rate than those who reproduce less or not at all.
The external forces that determine which traits or behaviors result in greater
reproductive success are called selection pressures. Selection continues to act on
traits, behaviors, and their underlying psychological and physiological mechanisms
so long as they exhibit heritable variation that influences reproductive success. It
should be noted that adaptive behaviors or mechanisms need not increase the
probability of reproductive success in all environments, or across time if environ-
mental conditions change more rapidly than selection can favor adaptive changes.
Selection can favor certain traits or behaviors in one environment, but others in
Categories of Analysis 5
another environment. Selection can also favor mechanisms that respond differently
to different circumstances in ways that enhance the probability of reproductive
success. Therefore, traits and behaviors do not have to be universal in order to
suggest that they have been shaped by natural selection.
A discordance or mismatch can occur when environments are vastly different
than those under which selection was acting to shape a particular behavior or
underlying mechanism (Eaton et al. 1988, 2002; Lieberman 2013). In the novel
environment, the behaviors or mechanisms may no longer increase reproductive
success and can even be detrimental to reproduction or survival. In order to
understand the evolution of psychological and physiological factors that influence
physical activity behavior, we must consider the mechanisms and behaviors that
would have been adaptive in the environments faced by humans during the majority
of human history, when selection was acting to shape them, rather than behaviors
that are expected to be adaptive currently. We can hypothesize that over the course
of human evolutionary history, a willingness to selectively exert oneself physically
would have generated survival and reproductive benefits through the ability to
produce more resources, protect oneself and family from danger, compete with
rivals, defend resources, migrate over long distances, meet the energetic demands of
pregnancy, childbirth, and lactation, care and provide for children, ward off sexual
predators, and display one’s physical health and condition to gain mates and social
partners (Caldwell Hooper 2013; Hönekopp et al. 2004, 2007).
Since humans reproduce sexually, traits, behaviors, and their underlying
mechanisms are also shaped by sexual selection. Sexual selection is a specific type
of natural selection that favors characteristics that improve mating success by
appealing to the opposite sex, even when they are detrimental to health and sur-
vival. The peacock’s tail is a classic example of a trait that evolved through sexual
selection. Cumbersome and brightly colored tails are energetically costly and make
male peacocks much more susceptible to predation, but they have evolved because
they are attractive to female peacocks. A key difference between traditional and
evolutionary approaches to understanding health behaviors is an understanding of
the fact that environmental mismatches and sexual selection can lead to charac-
teristics that are detrimental to health and survival. It is also important to understand
how psychological adaptations are thought to operate.
Psychological adaptions are designed to take in information from the environ-
ment, process that information together with endogenous information about the
individual, and output behaviors that increased the probability of reproductive
success in the environments in which they evolved (for a comprehensive review see
Tooby and Cosmides 2005). These computations do not require conscious pro-
cessing in order to output behavior that is or was probabilistically adaptive based on
environmental and endogenous inputs. Additionally, selection is predicted to favor
domain-specific over domain-general processing, just as organs have evolved to
perform specialized functions that together influence the survival of an organism
rather than one master organ that performs all the functions necessary to survive
(Tooby and Cosmides 2005; Tybur et al. 2012). This is at odds with traditional
approaches from health psychology and public health that implicitly assume health
behaviors are predicated on generalized, rational, and cognitive processes that are
motivated to increase health overall.
Considering how psychological and physiological mechanisms that influence
physical activity and health have been shaped by natural selection is therefore
distinct from traditional approaches in health psychology in important ways. First,
natural selection can lead to adaptations that are detrimental to health through
environmental mismatch and sexual selection. Second, an evolutionary perspective
does not require that information processing is conscious. In addition, traditional
approaches in health psychology use a domain-general approach that assumes the
cognitive process underlying all health behaviors is the same. In contrast, evolu-
tionary psychology predicts that the psychological mechanisms that influence a
specific health-related behavior (i.e., physical activity or exercise) take in and
process endogenous and exogenous information that is relevant for that behavior
and output behavior that probabilistically led to increases in reproductive success in
the environment that they evolved.
Including an ultimate or functional understanding of proximate predictors can be
advantageous for understanding why current efforts to increase physical activity are
unsuccessful and providing novel approaches to achieve these ends. For example,
many interventions and public health campaigns aim to increase physical activity
by increasing conscious awareness of the many health risks of being inactive. In
modern, Western environments, the benefits of physical fitness and activity for
decreasing morbidity and mortality are striking, particularly with respect to chronic
cardiovascular and metabolic disorders. It may seem logical that natural selection
would favor a psychology that mitigated health risks because more healthy people
may survive longer and are more likely to reproduce successfully. However, evo-
lutionary psychology would not predict that a general psychological mechanism to
promote health would have evolved. As stated earlier, selection does not favor
generalized mechanisms (Tooby and Cosmides 2005; Tybur et al. 2012). Second,
natural and sexual selection can favor traits and behaviors that are harmful to health
and survival when they probabilistically led to greater reproductive success in the
environment they evolved. In addition, the most critical health risks related to
inactivity (e.g., cardiovascular disease, type 2 diabetes, cancer) typically do not
manifest until much later in life, and there is no paleontological or historical evi-
dence to suggest that these chronic diseases were primary sources of mortality until
the present day. They are what evolutionary medicine has termed mismatch con-
ditions. Mismatch conditions are diseases that are more common or severe currently
because they result from mechanisms or behaviors that evolved in environments
that are vastly different from current ones (Lieberman 2013). Ancestral environ-
ments are thought to be very different from current environments in both the
physical activity necessary to survive and the availability of food (Eaton and Eaton
2003; Eaton et al. 1988, 2002). Thus, it is unlikely that natural selection has shaped
human psychology in a way that would favor recreational physical activity
throughout life because of a conscious awareness that doing so may prevent health
risks in our modern, high-tech, largely sedentary environments. An evolutionary
perspective would instead predict that raising awareness of the more immediate
benefits or those that are salient for increasing the probability of reproductive
success at a given stage of life would be more effective at influencing behavior.
Indeed, one recent study has shown that people who report that they exercise for
health benefits exercise less than those who do so for improving current quality of
life (Segar et al. 2011).
Phylogeny
Another category of analysis that would not be considered by traditional health or

exercise research is a phylogenetic or comparative approach to understand the
factors that influence activity across species, particularly those closely related to
humans, such as chimpanzees, bonobos, and other great apes. This allows
researchers to compare the evolutionary trajectory between species to understand
how humans are expected to be similar or different based on other evolved dif-
ferences in bodies, brains, life span, muscle composition, social structure, and so
on. As an example of a phylogenetic approach, we can consider how other animals
allocate energy throughout their life span in foraging and reproduction that differ
from humans.
Chimpanzees are one of our closest primate relatives and are estimated to have
diverged from a common ancestor relatively recently (6–8 million years ago).
However, chimpanzees take a very different approach to childrearing. Chimpanzee
mothers care for one offspring at a time, without the help of fathers, for approxi-
mately five years (Sugiyama 2004). At this time, offspring can provision for
themselves entirely, and mothers can shift energy to her next offspring. In contrast,
human females have a much shorter interbirth interval—2–3 years in natural fer-
tility populations—and due to our unique, learning-intensive foraging niche, off-
spring cannot provide for themselves for decades (see Fig. 1.1; Gurven et al. 2006;
Hooper et al. 2015; Howell 2010; Marlowe 2010). Thus, human females care for
and help provision multiple dependent offspring simultaneously, and can be
pregnant, lactating, and raising older, dependent children at the same time.
All of these activities are calorically expensive and influence the energy that can
be allocated to food production by women, and the amount of food that needs to be
produced. It would not be possible for human females to meet these energetic
demands without the help of male partners and larger kin and social networks. The
differences in human patterns of reproduction and provisioning for offspring are
therefore likely related to male and female patterns of activity throughout the life
span, as well as body mass differences in fat and muscle distribution (we will return
to this in Part II, Chap. 3). These differences in parental investment and body
composition also lead to differential costs and benefits of physical activity over
human history and are predicted to continue to influence patterns of physical
activity in modern contexts.
Fig. 1.1 Net production

throughout the life span
among the Tsimane’. This
graph was reproduced from
(Hooper et al. 2015) and
demonstrates the unique
pattern of human life history
that is characterized by an
extended juvenile period of
dependency and large surplus
of production later in life
Development
Finally, an evolutionary perspective includes an understanding of the develop-

mental trajectory of a behavior throughout the life course and of developmental
plasticity. Developmental plasticity is a term to describe how an organism’s
genotype can give rise to a range of phenotypes depending on the environmental
conditions experienced throughout development (West-Eberhard 2003). A reaction
norm is a specific pattern of phenotypic expression in response to environment.
Some environmental factors that are predicted to influence reaction norms in
physical activity are resource availability, the energy required to extract resources,
exposure to pathogens and infections, and social and cultural factors. These changes
in phenotypic expression are not necessarily associated with variation in gene
frequency and thus are not necessarily heritable. Rather, they are outcomes of a
flexible system favored by selection that tended to be adaptive in the environments
in which it evolved.
For example, the human lineage began in Africa, but over time spread to a
variety of ecosystems, with unique abilities to physiologically, morphologically,
behaviorally, and culturally adapt to all of them, including high altitudes with
hypoxic stress, or a reduction in oxygen availability (Antón et al. 2002; Leonard
2015). Indigenous populations who live at high altitudes around the world exhibit
larger lung volumes in adulthood—which is achieved by faster growth of the
organs associated with oxygen transportation (heart and lungs) during childhood
and adolescence despite slow overall rates of growth (Leonard 2015). This physi-
ological adaptation undoubtedly influences the costs of physical activity, particu-
larly at high altitudes, whereas the benefits of investing more in heart and lung
growth only pay off when physical activity at high altitudes is a necessary part of
life. An evolutionary perspective would lead to predictions that those who live at
lower altitudes or potentially those who live at high altitudes but do not perform
physical activity would invest less in lung and heart growth during adolescence, and
would incur higher costs of activity at high altitudes.
Physiological and psychological factors that influence physical activity are
affected by environments and experiences faced during development, as well as
those that shaped these mechanisms ancestrally, and even those that shaped our
lineage before diverging from a common ancestor with chimpanzees, bonobos, and
other primates. An evolutionary perspective that incorporates each of these cate-
gories of analysis allows for a more complete understanding of physical activity and
underlying mechanisms that is not possible from other perspectives.
Summary
We have broadly explored the advantages of using evolutionary theory to help us

understand physical activity. Now, we will delve deeper into these advantages by
first exploring a body of theory from evolutionary biology, life history theory, that
is particularly relevant for understanding human physical activity behavior. Next,
we will examine evidence from a comparative, phylogenetic perspective and
summarize research on physical activity and energy expenditure in humans cur-
rently and across human evolution. In the following section, we will consider the
human life span and developmental trajectory and sex-specific constraints on
physical activity. In each chapter and section, our focus will be on how natural
selection is hypothesized to shape behavior, and the proximate predictors of
behavior, in ways that affect reproductive success and survival from relevant
mortality risks as the ultimate level of explanation.
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Chapter 2
Physical Activity and Life History Theory
…the fundamental theoretical issue faced by the social, behavioral, and medical sciences is
how to build models of an ancient, but flexible, response system in a very novel envi-
ronment.—Hillard Kaplan, 1997.
Life history theory (LHT) is a branch of evolutionary theory aimed at explaining

differences in the age schedules of growth, reproduction, and mortality (Charnov
1993; Stearns 1992). LHT proposes that humans, like other species, face trade-offs
in how to optimally allocate the limited resources of time and energy among growth,
reproduction, and maintenance (i.e., immune function, survival, longevity). Optimal
allocations between these demands are expected to depend on individual and eco-
logical parameters that influence resource production, mortality, and other factors
(Hill and Kaplan 1999). There are inherent trade-offs between somatic versus re-
productive effort, current versus future reproduction, mating versus parental effort,
and quantity versus quality of offspring. The theory posits that there have been
long-term evolutionary pressures to balance energetic effort, and channel resources
toward behaviors that maximize expected reproductive success across the life span.
Life history strategies of energy allocation interact with the environment to funda-
mentally determine rates of growth, age of reproductive maturation, the pace of
reproduction, reproductive senescence (particularly in humans), and life span.
Physical activity is particularly well suited for a life history perspective because
it both influences and is influenced by life history parameters (such as age of
reproductive maturation, survival, and longevity) and is central to both energy
expenditure and (at least traditionally) energy acquisition. According to LHT, a
trait, characteristic, or behavior is maintained over evolutionary time when its
adaptive benefits outweigh its costs of time, risk, and energy invested. In this book,
I contend that human physiology and psychology have been shaped by natural
selection to maintain physical fitness and selectively engage in physical activity
when the adaptive benefits outweigh the metabolic and opportunity costs.
Conversely, when the benefits do not outweigh the costs (in terms of reproductive
fitness, after all trade-offs), humans, like other animals, have evolved to conserve
energy through behavioral and physiological shifts in energy allocation. This drive

12 2 Physical Activity and Life History Theory
for optimization has been shaped throughout human evolution to flexibly respond to
environmental and individual conditions that influence the costs and benefits of
physical activity across the life span. Considering how the costs and benefits of
physical activity have been shaped by natural selection offers a prime example of
integrating proximate mechanisms within a functional framework.
Energetic Costs of Physical Activity
On the one hand, physical activity entails high energetic costs. In addition to the
direct metabolic costs of performing physical activity, costs are also incurred to
develop and maintain cardiovascular and muscular fitness in humans. Muscles
consume about 40 % of a body’s resting metabolic rate (Lieberman 2015). Muscle
is much more energetically costly than adipose tissue, and the high costs of car-
diovascular and muscular fitness in humans can be inferred from the significant
decreases in fitness and strength in response to detraining (Booth and Lees 2007;
Coyle et al. 1984; Houston et al. 1983). In a preliminary study, researchers saw
decreased fat-free mass and increased abdominal fat after one week of decreasing
the number of steps taken per day from *10,000 to *1400 steps by taking cabs
and elevators rather than walking, cycling, or taking the stairs (Olsen et al. 2008).
This result suggests that humans have a rapid shift in metabolism from devoting
calories to maintain muscles instead being partitioned into fat when muscles are not
being used, even for relatively short time periods. Exercise also leads to changes in
metabolism that are protective against risks of obesity and type 2 diabetes in our
modern environment (Bassuk and Manson 2005). However, the maintenance of this
protective metabolic state also appears to entail significant costs, because it is lost
remarkably quickly in the absence of physical activity. For example, improvements
in insulin action have been shown to be reversed back to untrained levels in as little
as 38 h without exercise in trained male athletes (Oshida et al. 1991), and insulin
action is significantly reduced after just one day of sitting among healthy adults
(Stephens et al. 2011).
A potential explanation for the rapid changes in metabolism in response to
physical activity is that our metabolism has evolved to facultatively respond to
changes in physical activity. Given that the majority of physical activity across
human history was “obligatory” (in that, it was directly necessary to meet everyday
needs), and the ability to consistently store food extra-somatically relatively recent,
decreases in physical activity were likely associated with decreases in the avail-
ability of food in the environment. Those who were more efficient at storing fat
during times of low activity would be better off during times when less food was
available (Brown and Konner 1987).
On top of the direct metabolic costs, and costs of developing and maintaining
strength and cardiovascular fitness, humans also face risks of injury in developing
fitness, and opportunity costs when performing physical activity. That is, time and
energy spent in physical activity or maintaining strength and physical fitness cannot
Energetic Costs of Physical Activity 13
be spent on other activities important for survival or reproductive success. Over the
course of human history, physical activity was obligatory for producing the food,
shelter, and protection necessary to survive and reproduce; this ensured that a
greater weight was placed on allocating energy to physical activities to achieve
these basic needs, relative to today. Opportunity costs also vary with age, sex, and
individual condition. Trade-offs in allocating energy to physical activity versus
reproduction are more acute at certain points in the life span. This is particularly
true for females more than males, as women bear the heaviest burden of energy and
time devoted to reproduction that is incompatible with physical activity. Differences
in opportunity costs of physical activity according to sex and age are illustrated in
an example from the Tsimane’.
The Tsimane’ are an indigenous population in the Bolivian lowlands whose
economy is primarily subsistence based. They engage in foraging and small-scale
horticulture. Like many South American indigenous groups, the Tsimane’ have
adopted soccer as a principal form of recreational physical activity. While soccer is
often played several times per week in any given village, there is large variation in
how much time individual men and women devote to playing. The current theo-
retical framework would suggest that this variation can be explained, in part, by
variation in the costs and benefits of participation. Among the Tsimane’, girls can
be found playing soccer often, but few women play. Among men, net caloric
transfers of food to offspring provide a clear quantitative indicator of reproductive
effort in the form of paternal investment. The peak net transfers from Tsimane’ men
to offspring occurs in their 30s, with an average net transfer of 1740 calories per day
(Hooper et al. 2015). This peak occurs when their children are likely to be living at
home, old enough to have substantial caloric needs, but young enough not to
contribute much to the household resources. When men are younger, however,
reproductive effort is concentrated much more on attracting mates and social
partners, which need not trade-off as clearly against allocating energy to playing
soccer. Many men do not marry until their mid-20s, and men recently married
appear to maintain some level of mating effort, because they are more likely to have
affairs, and many marriages break up in the first couple of years (Winking et al.
2007). Compared to younger men, older men may also be more prone to injury, or
have more to lose if they do get injured because their bodies take longer to heal and
they have more dependents relying on their food production.
Given these differential costs, I would predict that younger men (<30 years old)
play soccer more frequently than older men, who must spend more energy producing
food for their families and who may face higher risks of injury. A sample of 166 men
from six Tsimane’ villages supports this prediction (unpublished data from
Benjamin Trumble, UC Santa Barbara, and Daniel Cummings, University of New
Mexico). A t-test comparing men who are under 30, to those 30 and over showed
that younger men played soccer significantly more days per week than older men t
(87) = 5.42, p < 0.001. Younger men played an average of 3 days, with 75 %
who reported playing soccer at least 3 days a week, with two men playing everyday
of the week. Older men played just 2 days per week on average. Only 36 % of the
older men played 3–5 days per week, and none over 5 days. The most frequent
response, given by half of the older men, was only 1 day per week. While we know
that older men produce more calories (Hooper et al. 2015), this analysis does not
directly tell us whether there is a trade-off between time and energy spent playing
soccer versus producing food. It could be that older men tend to buffer against
unnecessary risks, or that their preferences for leisure time activities change over
time, or that their fitness decreases, making it more difficult and less fun to compete
with younger men. Each proximate explanation could serve the ultimate function of
prioritizing physical activity that is consistent with primary reproductive benefits that
change over time (i.e., mating versus parenting effort). Anecdotally, having seen the
enthusiasm and joy expressed by all the men I have seen playing soccer; even in the
face of losing to a rival community, heat, humidity, and exhaustion, I doubt that a
lack of interest or enjoyment drives men to play less soccer. However, there is
additional support for the hypothesis that time and energy spent producing food
trades off against leisure time soccer playing. In a subsample of these men (N = 50),
we also have data on how many hours per day they spend hunting (unpublished data
from Paul Hooper, Emory University). In a regression model, the number of hours
spent hunting per day negatively predicted the number of days per week spent
playing soccer (B = −2.24, p < 0.001; controlling for both age and age squared).
Therefore, holding the effect of age constant, averaging one more hour of hunting
per day is associated with 2.24 fewer days of playing soccer per week.
Benefits of Physical Activity
Physical activity also has many benefits. As discussed previously, the decreased
risk of many chronic and deadly diseases we see in our current environment are
very important when considering the benefits of physical activity today. We will
consider those in the final section on future directions, but the majority of this book
focuses on those benefits hypothesized to influence reproductive fitness in ancestral
environments. These hypothesized benefits include, but are not limited to: com-
peting with rivals; attracting mates and social partners through displays of physical
health and prowess; meeting the energetic demands of pregnancy, childbirth, and
lactation; caring for children; producing food through foraging; processing food;
protecting oneself and family from danger (including sexual predators); defending
resources; migrating over long distances; building shelters; and collecting fuel for
warmth and protection.
Summary and Applications
To summarize, there are substantial costs and risks associated with investing
energy and time in physical activity that must be balanced with investing time
and energy necessary to grow and maintain a healthy body, and reproduce
Summary and Applications 15
successfully. In humans, reproducing successfully involves not only mating effort,

but substantial parenting effort to increase the likelihood that offspring also survive
and reproduce.
Beginning with the advent of agriculture, humans have been very successful at
finding more efficient ways to extract resources. In even more recent human history,
technological advances and globalization have led to an especially rapid and dra-
matic increase in the efficiency of resource extraction, distribution, manipulation,
and consumption, such that little energetic expenditure is required to fulfill our
basic biological goals. It is no coincidence that modern, Western environments have
very few energetic demands in order to meet our survival and reproductive goals.
Rather, they reflect our innate, evolved drive to decrease the energy required to
meet these goals in order to conserve energy for growth, maintenance, and repro-
duction. The inborn conservatism of our evolved energy allocation systems dis-
courages us from maintaining energy expenditures similar to Pleistocene ancestors
in the absence of tangible evolutionary benefits. This innate drive to conserve
energy has only recently been coupled with massive abundances of calorically
dense, easy to acquire foods. Energy conserving behaviors and metabolic adapta-
tions that increased the likelihood of survival and reproduction in environments
with fewer and less predictable resources now lead to deleterious health problems
and can even hinder reproduction. This has been termed a discordance or mismatch
between adaptations and environments (Eaton et al. 1988, 2002; Lieberman 2013).
How can an understanding of the costs of physical activity as they relate to
reproductive success inform efforts to increase physical activity and exercise in
modern environments? One advantage of this approach is helping to differentiate
between costs and benefits that are malleable versus those that are not, to design
interventions to target those that are changeable, while working around those that are
not. For example, the metabolic costs of performing physical activity cannot be
changed by intervention techniques. They will change with experience performing
physical activity, but an intervention cannot make running or other physical exertion
somehow less costly, so people are more inclined to do it. Doing so would also make
some people less drawn to it, or defeat the purpose, as people in our current envi-
ronment seek exercises that help them expend as much energy as possible. Some
people go to great lengths to change their workouts regularly to keep their bodies
from acclimating to exercises and performing them more efficiently; our bodies want
to expend less energy, so we have to stay one step ahead if we want to exert more.
However, an evolutionary framework can explicitly model and make predictions
about factors that lead to individual differences in how people physiologically and
psychologically respond to physical activity, training, and fitness that change the
metabolic costs of activity at an individual level. For example, differences in
condition brought about by inherited genetic variation (and potentially epigenetic
changes), adaptations to developmental environments, prior physical fitness, burden
to the immune system, access to resources (socioeconomic status) are all predicted
to influence the individual costs of physical activity. Moreover, the metabolic and
opportunity costs of physical activity change throughout the life span and are
different in men and women (detailed in Part II). Considering the individual level
costs can help to know when it may be more or less difficult for individuals to begin
or continue exercise and may predict the extent to which conscious, cognitive
processing is likely have a strong impact on behavior at a given point.
In contrast to the relatively rigid energetic costs of physical activity, the benefits
are much more malleable because, while they all ultimately relate back to repro-
ductive fitness and survival, they are partly defined by individual, ecological, and
cultural factors. For example, there are traditional populations like the Tarahumara
of northwestern Mexico that continue cultural traditions that include high levels of
endurance running. Historically, the Tarahumara would hunt deer and antelope
using persistence hunting, a form of hunting that requires endurance running and
walking over long periods of time to exhaust and overheat prey (Lieberman 2014).
The Tarahumara no longer practice this hunting technique, but the importance of
endurance athleticism has remained a large part of their culture. They participate in
annual rarajipari (for men) and ariwete (for women) games that in which teams of
men kick and chase a small wooden ball for around 75 km (46.6 miles), and teams
of women chase a hoop over shorter, but still extensive distances (≤40 km,
*25 miles). In this culture, being able to show physical prowess likely confers
more benefits than in cultures where there is less of an emphasis on endurance
athletic ability.
It is important to stress that critics of an evolutionary perspective have concerns
that this approach is deterministic and presumes that behavior is determined by
genes that were shaped by natural selection and play out regardless of environments
or developmental experiences (e.g., Lickliter and Honeycutt 2003). In stark con-
trast, a LHT perspective transcends a false dichotomy of nature versus nurture.
Instead, a LHT perspective predicts that the optimization system of energy allo-
cation is what has evolved; it has been shaped to flexibly and adaptively respond to
environmental and developmental cues regarding resource availability, how those
resources are acquired, individual condition, and extrinsic mortality risks. This
distinction is crucial for those interested in increasing health and wellness in current
environments through increased physical activity, because it suggests that although
there is a mismatch between the environment and adaptive behavioral and physi-
ological responses, there is flexibility to shape the reaction norms, or trajectories of
energy allocation, by manipulating the costs and benefits, and changing environ-
mental conditions to require or at least favor higher levels of activity. A clear
understanding of life history parameters and evolved energy allocation systems that
influence physical activity in humans will also help us to find opportunities for
increasing physical activity by reducing the costs or increasing the benefits, so the
adaptive benefits outweigh the metabolic, maintenance, and opportunity costs
involved. Toward that better understanding, we will begin this quest by examining
the factors that are unique to human life history and the costs and benefits of
physical activity from a phylogenetic and comparative perspective.
References 17
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Chapter 3
Phylogeny and Life History Patterns
Phylogenic or comparative analysis allows us to examine how traits or behaviors

evolved in humans and other animals, and to discover factors that lead to simi-
larities and differences in traits and behaviors over evolutionary time. This chapter
focuses on unique morphological, physiological, and metabolic adaptations that
influence energy expenditure and physical activity in humans compared to primates
and other mammals. Next, it examines the evolution of human life history patterns
and how the human pattern compares to other primates and mammals. Finally, it
compares physical activity patterns and energy expenditure in primates and
humans. Throughout, I highlight the ways the unique aspects of human anatomy,
energetics, and life history are hypothesized to influence costs and benefits of
physical activity, and developing and maintaining physical fitness and strength.
Human Morphological, Physiological, and Metabolic

Adaptations
The hominin line diverged from the last common ancestor with chimpanzees
between 6 and 8 million years ago. Ecological changes led to two major shifts in
behavior after this divergence: a descent from the trees and a shift to bipedal
locomotion. At the end of the Miocene, forested habitats were becoming less
common, and vast open landscapes with a cooler, drier climate set the scene for
hominin ancestors who increased roaming distances to find more distantly spaced
food patches (Cerling et al. 1997, 1998). Many of the details of hominin evolution,
including the evolution of bipedalism, are not conclusive, but current research
suggests that hominin ancestors have morphological adaptations such as
sideways-facing pelvic bones that stabilize the torso, long hind limbs, and feet with
arches and toe joints that allow for propulsion that made locomotion, especially
bipedal locomotion more efficient by the mid-Pliocene (Pontzer et al. 2009, 2014;

20 3 Phylogeny and Life History Patterns
Sockol et al. 2007). Researchers have estimated that the earliest evidence for effi-
cient bipedalism is in Australopithicus afarensis, about 3.4 million years ago
(Pontzer et al. 2014). Modern subsistence humans exhibit much larger ranging
behavior (9–15 km/day among the Hadza; Marlowe 2010) compared to chim-
panzees (2–5 km/day Pontzer and Wrangham 2004, 2006).
Around two million years ago, Homo erectus emerged, with a much larger brain,
long legs, and generally humanlike body (Lieberman 2013, 2015). Humans have
several anatomical and physiological adaptations that are unique among primates
and suggest that not only bipedal walking, but also endurance running was an
important part of human evolution (Bramble and Leiberman 2004). For example,
humans have springlike tendons and arches in the feet, collagen-rich tendons, and
ligaments that effectively store and release elastic strain energy, long legs, short
toes, and other traits that help stabilize and correctly move our upright torso and
head, aid in thermoregulation, and reduce the stress of running. Not only is
endurance running possible in humans because of these adaptations, but it also
involves neurobiological rewards in the form of increased endocannabinoids.
Increases in endocannabinoids in response to running are also exhibited by dogs
(another mammal that is considered to have evolved to habitually engage in
endurance running), but not in ferrets (mammals not considered to have evolved for
endurance running; Raichlen et al. 2012). The neurobiological reward may help to
encourage intense aerobic activity despite the high energetic costs and risks asso-
ciated with it. Recently, researchers have suggested that endurance running (and an
increased aerobic capacity) in humans also played a role in the evolution of our
uniquely large brains (Raichlen and Polk 2013). Since the divergence from the last
common ancestor with chimpanzees and bonobos, it seems that human ancestors
became morphologically adapted to cover long distances by walking and running.
On the other hand, humans have not simply evolved in ways that favor car-
diovascular and muscular fitness. Instead, humans seem to require habitual activity
to maintain physical fitness and favor uniquely high levels of energy storage.
Paleontological research suggests that human ancestors were substantially more
muscular than humans living in modern, sedentary contexts (Booth et al. 2002). But
when we compare humans to other primates, humans actually have much less
muscle tissue and more adipose tissue than other primates, which some researchers
have suggested has helped humans balance the increased metabolic costs of a large
brain (Leonard and Robertson 1997; Leonard et al. 2007; Zihlman and Bolter
2015). As discussed in Chap. 2, humans rapidly shift energy from maintaining
muscular fitness to storing fat. One study found decreases in fat-free mass and
increased abdominal fat after only one week of reduced physical activity (Olsen
et al. 2008). These rapid declines in allocation of effort to muscle tissue when not
being used may be unique to humans. Captive bonobos have high muscle mass,
despite having lower levels of activity than in the wild, which suggests that high
levels of activity are not necessary for maintaining muscle mass among one of our
closest primate relatives (Zihlman and Bolter 2015).
Human Morphological, Physiological, and Metabolic Adaptations 21
One recent and exciting advance for examining differences between species in
metabolism across tissues is to measure metabolite distribution across species. One
can then determine whether changes in the concentrations of metabolites in these
tissues can be explained by genetic divergence expected based on phylogenetic
divergence, or when the species diverged from a common ancestor. Bozek et al.
(2014) examined the metabolite distributions in brain, kidney, and skeletal tissues
across humans, chimpanzees, macaques, and mice. Among humans, they found that
while divergence in metabolite distributions in one brain region and in kidneys was
similar to what would be expected based on phylogenetic divergence, the metabolic
divergence in the prefrontal cortex and in skeletal muscle tissues was 4 and 8 times
higher (respectively) than what was expected based on phylogenetic divergence.
These results suggest that the metabolism of the prefrontal cortex and skeletal
muscle tissue has gone through rapid and parallel evolution since humans diverged
from the last common ancestor with chimpanzees. The authors interpret this finding
as supporting the prediction that the cost of our big brains—and specifically in the
prefrontal cortex—trades off against investment in skeletal muscle and strength. It
should be noted that this area of research is intriguing, but is still nascent, and does
not clearly demonstrate a trade-off in energy allocation between brain and muscular
development. These researchers also examined differences between humans,
chimpanzees, and macaques in pulling strength. Despite comparing a sample of
humans comprised of athletes and professional rock climbers to samples of captive
chimpanzees and macaques, the chimps and macaques averaged twice the pulling
strength of humans over a series of tasks. Taken together, along with the evidence
that humans rapidly lose muscle tissue in the absence of activity, it appears that
human physiology is driven to avoid the metabolic costs to build and maintain
skeletal muscle when not necessary, particularly in the upper body, in favor of
greater energy storage.
Humans appear to be somewhat unique among primates in their propensity for
storing energy. Humans have larger fat deposits (between 14 and 26 % among
healthy adults, even among modern foragers) than wild and captive chimpanzees
and bonobos (3–10 %; Zihlman and Bolter 2015; Zihlman 1984) and wild baboons
(Altmann et al. 1993). However, the body fat of other captive primates (lemurs,
baboons, and Western lowland gorillas) tends to fall within the human range.
Female primates also tend to have more fat than males. Humans are clearly unique
in the ability to store energy extra-somatically, with evidence of food storage in
granaries occurring before the domestication of plants during the Pre-Pottery
Neolithic A (≈11,300–11,175 years before present) to buffer against seasonal or
annual shortages in resources (Kuijt and Finlayson 2009). On an evolutionary
timescale, this is still a relatively recent development. Over longer time spans,
humans have had complex networks for food sharing which are unique in the
animal kingdom and have allowed us to buffer fluctuations in resource availability
and competing demands of energy allocation that interfere with resource production
(Kaplan and Gurven 2005).
Human Uniqueness in Life History Patterns
Recall that the ways energy is allocated across the life span determine pre- and
postnatal growth rate, gestation length, size at birth, age of reproductive maturation,
length of reproductive age, rate of reproduction, and life span. Across mammals,
these biological parameters are closely related to body size in predictable ways.
Typically, small organisms mature quickly and produce a large number of off-
spring. In contrast, large mammals have long life spans, slow growth rates, late age
of reproduction, slower reproductive rate, and fewer offspring, but greater invest-
ment in each offspring. Holding body size constant, life history parameters also
vary in patterned ways, with correlations between long developmental periods, slow
reproductive rates, and long life span in mammals and birds (Jeschke and Kokko
2009; Sibly and Brown 2007). Understanding the differences in these parameters,
and the environment-, species-, and individual-specific factors that give rise to
them, is central to a life history approach.
What does the life history pattern of non-human primates look like? Non-human
primates typically grow slower, reach reproductive age later, and reproduce more
slowly than would be predicted given their body size (Charnov 1993). These dif-
ferences are commonly attributed to an increase in somatic or bodily investment,
particularly in large, metabolically costly bigger brains. Wild chimpanzees typically
reach menarche at age 10 and have their first offspring at around thirteen years of
age. Captive chimpanzees, on the other hand, reach menarche earlier, at around
8 years, and have their first offspring between 10 and 11 (Sugiyama 2004). Earlier
age of menarche and first birth among captive chimpanzees demonstrates that while
life history parameters have been shaped at a species level, they are still sensitive to
changes in environment. In captivity, physical activity is lower, and diet is con-
sistent, which would all lead to earlier age of menarche and a shorter interbirth
interval, which is indeed the pattern that is observed. Wild orangutans reproduce at
15 and show a mean interbirth interval of 9.3 years. In captivity, however, age of
first reproduction is cut in half, and they can have interbirth intervals of less than
one year (Shumaker et al. 2008). These comparisons demonstrate how life history
parameters are sensitive to energetic condition and work to maximize reproductive
success.
In humans, a shift to nutrient-dense, difficult-to-acquire resources is theorized to
have strongly influenced the coevolution of increased brain size and life span, an
extraordinarily long dependent juvenile period that allows for complex development
and learning, complementary between the sexes in provisioning for multiple
dependent offspring, and an extended post-reproductive life span (Kaplan 1997;
Kaplan et al. 2000; Lancaster and Kaplan 2009). During childhood, energy is
focused on growth (particularly of brains) and maintenance (including immune
function) with zero effort allocated to current reproduction. Human brains are hugely
metabolically costly and take 6–7 years to grow, compared to the chimpanzee brain,
Human Uniqueness in Life History Patterns 23
which takes three years (Lieberman 2013). It has been suggested that the slow rate of
human growth is due to the high energy costs of our brains. These costs have been
estimated to peak in childhood, utilizing 66 % of resting metabolism, and 43 % of the
overall energy requirement around age four (Kuzawa et al. 2014). This study also
found that growth is slower during times when the energetic demands of the brain
(measured by glucose uptake) are higher, suggesting a trade-off between allocating
energy between brain and body growth.
Throughout development, children learn about their environment and develop
the skills they need to live and produce resources in a given ecology. Older children
spend a substantial amount of time and energy caring for younger siblings, which
allows mothers to put more energy into other activities (Kramer 2005), but do not
typically contribute significantly in the production of resources until older ages.
Children’s productivity varies between populations and depends in part on the type
of food production and the safety of performing this production (Jones et al. 1996).
In less safe settings, children produce fewer resources and tend to stay close to
home. During puberty, energy must be balanced between the costs of growth,
maintenance, reproductive effort, and the beginning stages of mating effort.
Following puberty, there are trade-offs in energy allocation toward reproductive
effort between mating and parenting effort, and between the quantity versus quality
of offspring.
The energetic costs of parental investment are much greater for humans com-
pared to other primates, because of the huge investment in neurological develop-
ment, altriciality at birth, and caloric dependency into the second decade of life
(Aiello and Key 2002; Hooper et al. 2015; Kaplan et al. 2000). Humans have babies
that are bigger than predicted by maternal body size and have significantly more
body fat (15 %) than other primates (Kuzawa 1998; Zihlman 1997). The energy
costs of fat deposition in human babies increase the energy demands on human
mothers compared to other primates (Dufour and Sauther 2002). At birth, the
human brain utilizes a large proportion of resting metabolic rate and total energy
budget, which has historically been produced completely through lactation
(Lieberman 2013). Women pay a much higher price than men for the initial
metabolic investment through pregnancy, particularly in the last trimester when
fetuses are storing extra fat, increasing energy expenditure by 465 kcal/day over
prepregnancy levels. Lactation is estimated to necessitate an extra 625–700 calories
per day for nursing mothers (Butte and King 2005; Ellison 2001). The high energy
costs for women, and the returns to sex- and age-specific division of labor, combine
to favor increased investment from fathers and other close kin, while pregnant and
lactating women theoretically and empirically decrease energy devoted to resource
production when possible (Hooper et al. 2015; Hurtado et al. 1992; Madimenos
et al. 2011; Marlowe 2010). These unique characteristics of human life history
patterns impact the energy available to allocate to physical activity (and related
maintenance) as well as the costs and benefits of physical activity throughout the
life course.
Summary
One way to utilize a life history approach to understanding physical activity is to

consider each stage independently to examine the energetic trade-offs theoretically
expected to influence the optimization of energy allocation. This optimization has
been shaped by natural selection to increase predicted reproductive success.
Considering how energy allocation across the life span has been shaped to maxi-
mize reproductive success can lead to hypotheses about what factors influence the
costs and benefits of energy expenditure in the form of physical activity. One can
also take a cumulative approach, because LHT also predicts that experiences and
environments at earlier stages of life, particularly during development, will influ-
ence later energy optimization strategies. It is therefore important to look at each
stage of life to know what the salient costs and benefits are within that stage, but
also how it may influence behavior in later stages, and how earlier experiences and
environments may have shaped responses to current environmental cues. The goal
of Section II is to do just that, tracing the stages of the human life course through
fetal development, childhood and adolescence, reproductive age, and
post-reproductive age in men and women. In each stage, predictions from a life
history framework will be presented regarding the energetic trade-offs that are likely
to influence the costs and benefits of allocating energy to physical activity. In
addition, each chapter will synthesize cross-disciplinary research that support the
use of this framework, or can be meaningfully interpreted in light of theoretical
predictions on energetic trade-offs that favor optimizing predicted reproductive
success. First, Chap. 4 will summarize some measurement issues in physical
activity research and the state of research on physical activity patterns in humans.
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Academy of Sciences, 111(4), 1433–1437. doi:10.1073/pnas.1316940111.
Pontzer, H., Raichlen, D. A., & Sockol, M. D. (2009). The metabolic cost of walking in humans,
chimpanzees, and early hominins. Journal of Human Evolution, 56(1), 43–54. doi:10.1016/j.
jhevol.2008.09.001.
Pontzer, H., & Wrangham, R. W. (2004). Climbing and the daily energy cost of locomotion in
wild chimpanzees: Implications for hominoid locomotor evolution. Journal of human
evolution, 46(3), 317–335. doi:10.1016/j.jhevol.2003.12.006.
Pontzer, H., & Wrangham, R. W. (2006). Ontogeny of ranging in wild Chimpanzees. International
Journal of Primatology, 27(1), 295–309. doi:10.1007/s10764-005-9011-2.
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104(45), 17707–17712. doi:10.1073/pnas.0707725104.
Sockol, M. D., Raichlen, D. A., & Pontzer, H. (2007). Chimpanzee locomotor energetics and the
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States of America, 104(30), 12265–12269. doi:10.1073/pnas.0703267104.
Sugiyama, Y. (2004). Demographic parameters and life history of chimpanzees at Bossou Guinea.
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evolving female: A life-history perspective 78, pp 86–106.
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sapiens has implications for changes during human evolution. Proceedings of the National
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1505071112.
Chapter 4
Physical Activity and Energy Expenditure
in Humans
Measurement
Before I discuss physical activity and energy expenditure in humans and other
primates, I must briefly discuss the different ways energy expenditure and physical
activity are measured. Doubly labeled water (DLW) is considered the gold standard
measure of energy expenditure. This technique uses stable isotopes of hydrogen and
oxygen to trace the flow of water and carbon dioxide (CO2) through the body, and
compute the rate of oxygen consumption (Butler et al. 2004). The primary
advantage of DLW is that it is a precise and relatively accurate way to measure total
energy expenditure (TEE). Total energy expenditure encompasses all energy
expended, including basal metabolic rate (BMR), the thermogenic effect of food
(the energy it takes to digest food), non-exercise activity thermogenesis, exercise,
and post-exercise oxygen consumption. From TEE, physical activity level (PAL) is
often estimated by first using a predicted value of BMR based on sex, age, height,
and weight, and then using one of several equations to estimate PAL.
The DLW technique has several limitations for understanding physical activity
patterns. First, DLW is typically considered to be a precise method, meaning that
when measured repeatedly, it consistently gives similar results; however, few
studies have actually tested its precision (Butler et al. 2004). In addition, its
accuracy may be less robust than is often assumed. Accuracy is the closeness of a
measurement to its true value. This can be evaluated with validation studies that
compare estimates of CO2 production and oxygen consumption derived from
simultaneous measurements from DLW with those from another type of mea-
surement, such as indirect calorimetry. In these validation studies, it is common for
some individual’s measurements to vary by more than 20 %. This suggests that
while in aggregate the two techniques yield relatively similar results, there is
potentially meaningful variation in accuracy between individuals that remains
poorly understood (Butler et al. 2004). Further, the way PAL is typically estimated

28 4 Physical Activity and Energy Expenditure in Humans
introduces non-negligible error by first estimating BMR and then having multiple
methods for calculating PAL, which make it difficult to compare across studies and
detract from the primary advantage of this method as being precise and accurate.
Further, the primary component of energy expenditure, BMR or resting metabolic
rate (RMR), changes due to several factors. For example, RMR is evidenced to be
influenced by an immune response to even minor respiratory infections
(Muehlenbein et al. 2010), reduced caloric intake (Martin et al. 2007), pregnancy
and lactation (Melzer et al. 2009), and acute and long-term bouts of physical
activity (Speakman and Selman 2003). In addition, RMR is often estimated using
weight, but is affected by body composition, with fat free mass explaining over
81 % of the variance (Sparti et al. 1997). When examining the proportion of energy
allocated across competing demands, the DLW method alone may not provide the
right level of detail necessary to understand differences across populations or
between individuals. It can be difficult, costly, and increases the subject burden to
directly measure BMR or RMR, but it may be worthwhile to more clearly and
accurately tease apart where energy is being allocated in concert with markers of
immune function and reproductive hormones. Even when reducing the costs by
estimating BMR, using the DLW technique very costly, and researchers wishing to
implement this technique in small-scale populations who live in remote parts of the
world without electricity must also have methods to freeze, store and transport urine
samples.
These difficulties aside, the DLW technique does not provide information about
the frequency, intensity, or duration of physical activity, which are critical for
understanding the numerous health benefits of physical activity and exercise.
Recent studies have examined energy expenditure measured by DLW over time and
across populations and have concluded that physical activity and exercise are not
related to recent increases in obesity. These studies do not find significant differ-
ences in total daily energy expenditure since the 1980s, and have shown that it is
not different from that in less developed, and third world populations (Dugas et al.
2011; Westerterp and Speakman 2008). Another study compared total energy
expenditure (TEE) between the Hadza in Africa and Westerners, and found no
significant differences between these populations, despite vastly different activity
patterns (Pontzer et al. 2012). Paradoxically, while there were significant differ-
ences between the Hadza and Westerners in PAL (2.26 and 1.81, respectively),
overall energy expenditure was not different. The authors interpret their findings to
suggest that because higher levels of physical activity do not increase overall
energy expenditure, diet is solely responsible for weight gain and/or recent
increases in overweight and obesity.1 However, another take-home message from
1
Decreases in physical activity are clearly not the only factor leading to increased rates of over-
weight and obesity. An entire book could (and should) be dedicated to understanding energy
balance, and the effects of diet and physical activity together for influencing human health and
well-being, weight gain, and obesity. It is premature to conclude that the proportion of energy
expenditure spent in active rather than sedentary activities does not contribute to weight gain, fat
storage, and obesity.
Measurement 29
these studies is that estimates of physical activity based on the DLW method may
not reflect true physical activity patterns and do not help us to understand the health
benefits of physical activity. While the Hadza are much more active than Western
populations, this higher activity level did not translate into higher overall energy
expenditure. In a follow-up study, these researchers demonstrated that measures of
physical activity such as daily walking distances or walking speed did not predict
TEE (Pontzer et al. 2015). Thus, for researchers aiming to understand the physical
activity energy expenditure and the many health outcomes associated with it,
estimates of overall energy expenditure, and potentially PAL estimates based on
TEE, are not likely to be the sole variables of interest. Teasing apart the proportion
of the energy budget that is spent on physical activity, or patterns of physical
activity and sedentary behavior may be more important for understanding the health
benefits of physical activity. In addition, much more research is needed to under-
stand the role of physical activity and energy balance in overweight and obesity.
It is counterintuitive that physical activity level is not directly related to overall
energy expenditure in the way we would expect, with higher levels translating into
higher overall expenditure. This is perplexing for researchers wishing to understand
energy expenditure from an evolutionary perspective. Given that total energy
expenditure is not different when physical activity is different, populations with high
levels of obligatory physical activity or those who regularly voluntarily exercise may
adaptively downregulate the energy necessary to perform other functions (Pontzer
et al. 2015). They may also perform somatic maintenance more efficiently, resulting
in similar TEE across individuals or populations with very different physical activity
patterns. It remains unclear whether the proportion of energy allocated to physical
activity in relation to other metabolic functions, such as tissue growth and mainte-
nance, and immune function influences weight gain and the recent and sudden
increases in obesity in the last 30 years. Further, physical activity, and likely the
proportion of the energy budget spent on physical activity, is associated with a
myriad of health benefits outside of weight loss. Thus, it is premature to conclude
that because TEE may not be different across populations, physical activity is
unimportant for health and/or weight loss. While TEE may be informative for telling
us the total energy budget across species or across individuals, which is important
for understanding the life history (Pontzer et al. 2014), it cannot tell us everything
about how that energy budget is being allocated across physiological systems, or
how that proportional allocation influences health and well-being.
Physical activity can also be measured by a factorial method that is calculated
from time allocation research methods. Researchers observe people and the activ-
ities they engage in over a period of time, or conduct recall interviews. Physical
activity is then estimated by computing the time spent in activities and the estimated
costs of those activities as a multiplier of estimated basal metabolic rate. Further
calculations can then translate these values into daily PAL levels. The factorial
method may be limited because the number of estimations that have to be made to
calculate PAL introduces measurement error. It has also been criticized because
when recall interviews are used, this can introduce even more error. It can also
underestimate total energy expenditure (Leonard et al. 1997). Given what we now
know about physical activity levels in the Hadza, measures of PAL or estimates of
TEE from the factorial method are unlikely reliable or accurate with current
methods of estimation as much remains unclear about how physical activity and
other metabolic functions combine to produce total energy expenditure.
A more objective method to measure physical activity is to use accelerometers or
heart rate monitors, or a combination of accelerometers and heart rate monitors.
Accelerometers measure movement in one or three axes, but do not capture
resistance-based movements, so can lead to underestimates of PAL if used exclu-
sively (Lyden et al. 2011). A combination of heart rate monitors and accelerometers
can be used to account for this underestimations. Where the accelerometer counts
are low but heart rate is at an active level, it is assumed that resistance activity is
taking place. Estimates for the intensity of those activities based on heart rate
measurements are thus added to the overall estimates of energy expenditure mea-
sured by accelerometer. As with the other measures of physical activity, many
estimations have to be made that can introduce measurement error in estimating
PAL and energy expenditure from accelerometers and heart rate monitors (in-
cluding estimates of BMR). Further, there are many different equations for esti-
mating energy expenditure from accelerometer counts, and equations are
necessarily different across the life span, which makes comparison across studies
difficult.
TEE in Humans and Other Primates
Researchers have examined the level of energy expenditure expected given body
size among placental mammals and found that primates have surprisingly low
energy expenditure, about 50 % lower energy expenditure than expected based on
body size (Pontzer et al. 2014). Primates also devote a greater proportion of their
energy budget to basal metabolic rate (BMR), which may be due to the metabolic
costs of big brains and longer life spans. This pattern is consistent with primate life
histories and suggests that TEE measured by DLW is an alternative to BMR for
examining the evolution of life history patterns across primates (Pontzer et al. 2014;
Sibly and Brown 2007).
Physical Activity in Humans over Time and Across

Populations
Human evolutionary history has largely taken place during the preagricultural
period, lasting until roughly 10,000 years ago (Biddle and Mutrie 2008; Blair
1988). During this time, humans subsisted through foraging and hunting nutrient
Physical Activity in Humans Over Time and Across Populations 31
rich but difficult to extract resources entailing moderate to high levels of obligatory
physical activity (estimated from both paleontological records and comparisons
with contemporary foragers). Starting in the Holocene, a transition from foraging to
agriculture took place for many human groups, which overall led to increased
physical activity (Panter-Brick and Pollard 1999) until approximately 200 years ago
with the beginning of the Industrial Revolution. The Industrial Revolution resulted
in a marked shift to machine-operated food production and decreased importance of
human physical energy expenditure for resource acquisition. This shift from a
reliance on human physical energy expenditure for food production and survival
has increased through the present. While technological advances have improved
health overall through sanitation, public health measures, and medical advances, a
lack of physical activity is a now considered a key risk factor for two of the leading
causes of disease burden worldwide, cardiovascular disease and depression (WHO
2008). Sitting, according to many obesity researchers, is considered to be the new
smoking, in terms of describing the extent of the health risks associated with a
sedentary lifestyle.
The transition to a primarily sedentary lifestyle and abundant caloric intakes has
taken place fairly recently in evolutionary terms, which has resulted in a mismatch
between human physiology, psychology, and the genes that code for that physi-
ology, and current environments. This mismatch or discordance has led to poor
mental and physical health (Booth et al. 2002; Chakravarthy and Booth 2004;
Cordain et al. 1998; Eaton et al. 2002; Eaton and Eaton 2003). However, our
behavior and physiology are not acting in maladaptive ways with respect to the
environments we evolved in; instead, natural selection has shaped our behavior and
physiology to conserve energy whenever possible in order to buffer against difficult
times and allocate more energy to increase reproductive success.
Does this mean we are doomed? No. But it does mean that it will be critical to
consider the selection pressures that have shaped our bodies and minds over mil-
lions of years, and work with our evolved energy allocation systems if we are
interested in increasing physical activity in modern contexts. Toward that aim, the
rest of the book will examine what we know about physical activity in traditional
and modern contexts, in light of life history theory to interpret results and derive a
testable framework for integrating the complex factors that influence physical
activity.
What can this perspective and available research tell us about the variables that
we can predict to be relevant for physical activity in humans? Measures of physical
activity vary largely across populations, as shown in Figs. 4.1, 4.2, and 4.3.
These figures include measures of physical activity level (PAL) in populations
from around the world and highlight the great variation in PAL between men and
women, and across a range of ecologies and modes of food production (Dufour and
Piperata 2008; Gurven et al. 2013; Katzmarzyk et al. 1994; Leonard 2008; Leonard
et al. 1995; Norgan et al. 1974; Panter-Brick 1996; Pearson 1990; Pontzer et al.
2012; Yamauchi et al. 2001; Yamauchi 2000). A clear, predictable pattern does not
Fig. 4.1 Variation in physical activity level (PAL) by sex among hunter-gather/forager
populations. PAL was estimated using a variety of data collection modalities: factorial method,
heart rate (HR), accelerometer, heart rate and accelerometer together, and doubly labeled water
(DLW)
emerge when comparing PAL across these dimensions and populations, even
within similar modes of subsistence. Men are sometimes, but not always more
active than women, and differences in physical activity are even observed within the
same mode of food production.2
Figure 4.4 depicts PAL levels across samples from which energy expenditure
was measured by the DLW method in both men and women in a meta-analysis
(Dugas et al. 2011). The populations sampled were categorized as low/middle or
high human development index (HDI). In meta-analytic analyses, physical activity
levels were significantly higher among men in low/middle HDI populations com-
pared to those in high HDI populations (1.88 and 1.79, respectively). Women in
high HDI countries had slightly, but not significantly higher PALs than women in
low-middle HDI (1.72 and 1.70, respectively).
2
It should be noted that PAL estimates were made using different methods of data collection,
which can also lead to differences.
Fig. 4.2 Variation in physical activity level (PAL) by sex among pastoralist populations. PAL
was estimated using a variety of data collection modalities: factorial method, heart rate (HR),
accelerometer, heart rate and accelerometer together, and doubly labeled water (DLW). Asterisk
denotes samples for which seasonal variation was averaged to obtain one estimate
Dufour and Piperata (2008) highlighted the importance of understanding the

factors that lead to variation in PAL among farming populations, by closely
comparing social and ecological conditions that influence PAL among women in
Yapu (a village in the Vaupes region of Columbia) and women living around the
Caxiuanã Forest Reserve in Brazil. Many indices of ecological condition among
these women were very similar: Both were subsistence agriculturists whose diet
consisted primarily of cassava as the staple and fish as the primary source of protein
in rural, rainforest environments that did not have electricity or running water.
However, they differed significantly in body fat and physical activity measured by
time allocation (PAL = 1.77 among the Yapu; PAL = 1.55 among the Caxiuanã).
A closer look at the differences in energy required by women to grow, harvest, and
process cassava partially explains the differences in PAL. Among the Yapu, women
were responsible for all agricultural work and processing. However, among the
Caxiuanã, men shared agricultural responsibilities as it was not entirely compatible
with childcare (women reported concerns about taking their children to the fields). The
Caxiuanã also processed cassava using mechanical graters, which were primarily
operated by the men. Carxiuanã women’s tasks varied widely in difficulty, and many
could be done with the help of older children. In both groups, the primary source of
Fig. 4.3 Variation in physical activity level (PAL) by sex among farming populations. PAL was
estimated using a variety of data collection modalities: factorial method, heart rate (HR),
accelerometer, heart rate and accelerometer together, and doubly labeled water (DLW). Asterisk
denotes samples for which seasonal variation was averaged to obtain one estimate
physical activity was derived from walking to and from the agricultural fields; thus, a
reduction in work in the fields among the Carxiuanã was reflected in lower levels of
physical activity. This comparison exemplifies the ways human physical activity is
reduced when possible, as well as the importance of considering environmental, social,
and cultural factors that together shape physical activity patterns.
This detailed comparison and the variation in physical activity that is highlighted
in Figs. 4.1, 4.2, 4.3, and 4.4 exemplify that there is not simply one level of physical
activity that humans have evolved to perform. Instead, the variation suggests that
humans flexibly respond to competing energetic demands across environments,
across seasons, and across the life span to selectively exert energy when the
Fig. 4.4 Variation in physical activity level (PAL) by sex for populations where PAL has been
estimated from doubly labeled water (DLW). Populations were defined as having low/medium
human development index (HDI) or a high HDI. In a meta-analysis (Dugas et al. 2011), physical
activity levels were significantly higher among men in low/middle HDI than those in high HDI
populations
adaptive benefits to doing so outweigh the costs, and otherwise conserve and/or
redirect energy. The following section serves to flesh out life history and epi-
demiological research to help understand variation in the costs and benefits of
physical activity across the life span.
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Part II
Physical Activity and Energetic
Trade-Offs Through the Lifespan
Chapter 5
Energy Costs and Benefits During Fetal
Development and Infancy
Despite the fact that there are no sizeable physical activity patterns to examine for
fetuses, it is important to consider early-life environments for understanding energy
allocation and physical activity. During fetal development and in infancy, all energy
is focused on somatic development of brains and bodies, growing, accumulating fat,
and maintaining tissue. One fundamental thing to consider at this stage is the huge
energy costs of human brains (infant brains require 50–60 % of resting metabolic
rate; Kuzawa et al. 2014). This stage is also important to consider because several
lines of research suggest that poor nutrition or energetic stress during prenatal and
early-life environments has a profound impact on health, development, and even
physical activity. Epidemiological evidence has consistently shown that early-life
undernutrition, indexed by low birth weight (alone and when paired with postnatal
overnutrition), is related to high blood pressure, type 2 diabetes, cardiovascular
disease, and stroke in adulthood (Barker 1989, 1994, 2004; Curhan et al. 1996).
After observing these patterns, researchers termed this phenomenon the fetal or
developmental origins of adult disease (reviewed in Barker 2004). Hales and Barker
(1992) hypothesized poor fetal and postnatal nutrition can lead to a “thrifty phe-
notype” which results in impaired development of the pancreas and endocrine
disruptions that lead to increased risk of type 2 diabetes.
Some evolutionary researchers have proposed that when faced with energetic
stress prenatally and in infancy, developmentally plastic energy allocation systems
make predictive adaptive responses (PARs) and shape more thrifty energy alloca-
tion systems later in life (PAR; Ellison and Jasienska 2007; Ellison 1990;
Gluckman et al. 2005; Kuzawa 2005; Kuzawa et al. 2007; Lipson 2001). It is
hypothesized that maternal cues of constrained resource availability were likely to
signify future environments with unstable resource availability, and therefore, it
was adaptive to shift reaction norms to develop a suite of metabolic characteristics
that help buffer against hard times with increases in fat accumulation, lower muscle
mass, insulin resistance, lower vascular density, compact body shape, and earlier
reproductive age. Proponents of the PAR hypothesis contend that over the course of

42 5 Energy Costs and Benefits During Fetal Development and Infancy
hominin evolution, later environments predictably did resemble prenatal environ-

ments, and selection would favor energy-saving mechanisms. Ancestrally, errors
were unlikely to produce deleterious health outcomes because all environments
required high levels of obligatory physical activity and had limited food availability
even in the best of times (Gluckman et al. 2005; Kuzawa 2007). They argue that the
changes in metabolic physiology that buffer against hard times would only lead to
harmful health outcomes when there is a mismatch between energetic stress during
fetal and infant development, paired with later-life environments with abundant
resources and much lower levels of obligatory physical activity.
This is particularly relevant for populations undergoing transition from a more
traditional lifestyle to one that is integrated with a market economy, as well as
examining health disparities based on socioeconomic status. Market integration is
typically accompanied by increased access to high-calorie, fatty, processed foods
including refined carbohydrates that were not previously available, which our
bodies are not physiologically adapted to process. Some researchers have argued
that rather than being an adaptive response, correlations between later-life health or
life history traits and early-life nutritional stressors merely reflect individuals
“making the best of a bad start” (e.g., Bogin et al. 2007). PAR theorists agree that
some environmental factors merely disrupt development, rather than serving an
adaptive function. It remains an open question whether or not this energetic pattern
reflects an adaptive shift in reaction norms to a more conservative energy allocation
strategy, or merely a disruption during development that leads to poorer health
outcomes. A considerable body of epidemiological and experimental evidence
supports the relationship between intrauterine environment and changes in devel-
opmental trajectory and metabolism (for a review, see Gluckman et al. 2005).
A similar, relevant, and related pathway for understanding the relationship
between early-life environments and later health is that of the impact of early-life
environments on the development of the immune system. From a life history per-
spective, investment in the immune system and immune responses is a costly but
necessary form of somatic investment that appears to flexibly respond to envi-
ronmental conditions, namely resource availability and exposure to parasites,
pathogens, and infectious diseases (Blackwell et al. 2010; McDade et al. 2005,
2008). In resource-scarce environments, it may be adaptive to reduce immune
investments overall, or to shape the immune system in energy-saving ways—ways
that increase less costly immune responses, or targeted responses to more likely
threats, and reducing efforts in other immune responses (McDade 2003, 2005).
Early-life undernutrition has been shown to moderate life history trade-offs between
growth and immune response in a preliminary analysis among Tsimane’ toddlers
(McDade et al. 2005). Among toddlers with poorer nutritional status, those with
larger investments in immune response, measured by high C-reactive protein (CRP;
a biomarker of the first, pro-inflammatory line of defense against pathogens) at
baseline, had less growth over a three-month period. Among toddlers in better
nutritional status, CRP was not related to growth, which indicates an energetic
trade-off between growth and the immune system. Those with more energetic
reserves face less severe trade-offs among life history traits, because there is more
5 Energy Costs and Benefits During Fetal Development and Infancy 43
energy available to invest in immune system and growth simultaneously. Energetic

trade-offs are more evident among those who are under energetic stress.
How is this relevant for understanding physical activity? As outlined in Part I,
the energy costs of performing physical activity and developing and maintaining
physical fitness and strength are high. Part of a thrifty phenotype may be a con-
servative energy allocation strategy that further reduces the likelihood of partici-
pation in non-obligatory and even obligatory forms of physical activity. A small but
growing number of studies on humans and rats are consistent with this perspective.
A meta-analysis of 13 Scandinavian birth cohorts demonstrated that lower-than-
normal birth weight was related to less leisure time physical activity later in life
(Andersen et al. 2009). Another study found low birth weight was related to higher
energy costs of running (Baraldi et al. 1991). In a recent study, birth weight pre-
dicted muscle efficiency when exercising, such that low birth weight was associated
with lower muscle efficiency (Workman et al. 2015). Lower muscle efficiency and
higher energy costs of running among those with low birth weight may support the
hypothesis that energetic stress early in life leads to a system that is not fully or
optimally developed, rather than an adaptive, energy conserving shift. It may also
be that these shifts are necessary to make other activities that are more necessary to
survival and reproduction more efficient. Either way, having lower muscle effi-
ciency or higher energy costs during activity are predicted to decrease the likeli-
hood of physical activity among those with lower birth weight, contributing to the
poorer health outcomes associated with early-life energetic stress.
In an experimental study of Wistar rats, mothers’ undernourishment predicted
offspring physical activity irrespective of postnatal feeding (Vickers et al. 2003).
Offspring of malnourished mothers were significantly smaller and less active than
offspring of ad libitum-fed mothers. Surprisingly, but consistent with the mismatch
hypothesis, the least active rats were offspring of malnourished mothers who were
fed a hypercaloric diet after birth. Taken together in light of life history theory,
these results suggest that reaction norms for energy allocation strategies are sen-
sitive to cues of resource availability early on in development. This may lead to
adaptive shifts in the optimization of energy allocated to later physical activity
similarly to the shifts observed in metabolism and immune function. In our modern
environments, we can test the hypothesis that individuals who experience nutri-
tional stress in utero or in infancy may have developed a more conservative allo-
cation optimum that buffers against energetic stress and prioritizes behaviors and
physiological systems more closely linked to reproductive success. This may lead
to higher costs for performing physical activity (e.g., if it is relatively less efficient,
or more difficult to develop muscle mass), or more psychological barriers (e.g.,
more displeasure in response to activity, lower motivation). For these individuals,
non-obligatory physical activity may be less likely to occur naturally, and more
difficult to initiate and/or maintain without substantial increases in the benefits, or
fewer trade-offs against investment in reproductive success.
44 5 Energy Costs and Benefits During Fetal Development and Infancy
References
Andersen, L. G., Angquist, L., Gamborg, M., Byberg, L., Bengtsson, C., Canoy, D., et al. (2009).
Birth weight in relation to leisure time physical activity in adolescence and adulthood:
Meta-analysis of results from 13 nordic cohorts. PLoS ONE, 4(12), e8192. doi:10.1371/journal.
pone.0008192.
Baraldi, E., Zanconato, S., Zorzi, C., Santuz, P., Benini, F., & Zacchello, F. (1991). Exercise
performance in very low birth weight children at the age of 7–12 years. European Journal of
Pediatrics, 150(10), 713–716. doi:10.1007/BF01958761.
Barker, D. J. P. (1989). Growth in utero, blood pressure in childhood and adult life, and mortality
from cardiovascular disease. British Medicine Journal, 298, 564–567.
Barker, D. J. P. (1994). Mothers, babies, and disease in later life. London: BMJ.
Barker, D. J. P. (2004). The developmental origins of adult disease. Journal of the American
College of Nutrition, 23(sup6), 588S–595S. doi:10.1080/07315724.2004.10719428.
Blackwell, A. D., Snodgrass, J. J., Madimenos, F. C., & Sugiyama, L. S. (2010). Life history,
immune function, and intestinal helminths: Trade-offs among immunoglobulin E, C-reactive
protein, and growth in an Amazonian population. American Journal of Human Biology, 22(6),
836–848. doi:10.1002/ajhb.21092.
Bogin, B., Varela Silva, M. I., & Rios, L. (2007). Life history trade-offs in human growth:
Adaptation or pathology? American Journal of Human Biology, 19, 631–642.
Curhan, G. C., Chertow, G. M., Willett, W. C., Spiegelman, D., Colditz, G. A., Manson, J. E.,
et al. (1996). Birth weight and adult hypertension and obesity in women. Circulation, 94,
1310–1315. doi:10.1161/01.CIR.94.6.1310.
Ellison, P. T. (1990). Human ovarian function and reproductive ecology. American
Anthropologist, 92, 933–952.
Ellison, P. T., & Jasienska, G. (2007). Constraint, pathology, and adaptation: How can we tell
them apart? American Journal of Human Biology, 19, 622–630.
Gluckman, P. D., Hanson, M. A., & Spencer, H. G. (2005). Predictive adaptive responses and
human evolution. TRENDS in Ecology and Evolution, 20, 527–533.
Hales, C. N., & Barker, D. J. P. (1992). Type 2 (non-insulin-dependent) diabetes mellitus: The
thrifty phenotype hypothesis. International Journal of Epidemiology, 35, 595–601. doi:10.
1093/ije/dyt133.
Kuzawa, C. W. (2005). Fetal origins of developmental plasticity: Are fetal cues reliable predictors
of future nutritional environments? American Journal of Human Biology, 17, 5–21.
Kuzawa, C. W. (2007). Developmental origins of life history: Growth, productivity, and
reproduction. American Journal of Human Biology, 19, 654–661.
Kuzawa, C. W., Gluckman, P. D., Hanson, M. A., & Beedle, A. S. (2007). Evolution,
developmental plasticity, and metabolic disease. In S. C. Stearns & J. C. Koella (Eds.),
Evolution in health and disease (2nd ed.). Oxford: Oxford University Press.
Lipson, S. F. (2001). Metabolism, maturation, and ovarian function. In P. T. Ellison (Ed.),
Reproductive ecology and human evolution. New york: Aldine de gruyter.
McDade, T. W. (2003). Life history theory and the immune system: Steps toward a human
ecological immunology. American Journal of Physical Anthropology, 37, 100–125. doi:10.
1002/ajpa.10398.
McDade, T. W. (2005). Life history, maintenance, and the early origins of immune function.
American Journal of Human Biology, 17(1), 81–94. doi:10.1002/ajhb.20095.
McDade, T. W., Leonard, W. R., Burhop, J., Reyes-García, V., Vadez, V., Huanca, T., & Godoy,
R. A. (2005). Predictors of C-reactive protein in Tsimane’ 2 to 15 year-olds in lowland Bolivia.
References 45
McDade, T. W., Reyes-García, V., Tanner, S., Huanca, T., & Leonard, W. R. (2008). Maintenance
versus growth: Investigating the costs of immune activation among children in lowland
Bolivia. Journal of Physical Anthropology, 136(4), 478–484. doi:10.1002/ajpa.20831.
Vickers, M. H., Breier, B. H., McCarthy, D., & Gluckman, P. D. (2003). Sedentary behavior
during postnatal life is determined by the prenatal environment and exacerbated by postnatal
hypercaloric nutrition. American Journal of Physiology Regulatory, Integrative and
Comparative Physiology, 285, R271–R273.
Workman, M., Baker, J., Lancaster, J. B., Mermier, C., & Alcock, J. (2015). Birth weight
predicted baseline muscular efficiency, but not response of energy expenditure to calorie
restriction: An empirical test of the predictive adaptive response hypothesis. American Journal
of Human Biology. doi:10.1002/ajhb.22818
Chapter 6
Energetic Trade-Offs and Physical Activity
During Childhood and Adolescence
One of the unique aspects of the human life span is an extended childhood. In contrast
with other primates, human offspring remain primarily dependent on their parents to
produce food for well over a decade after weaning. All energy during childhood is
focused on somatic growth and maintenance, including substantial continued
investments in brain growth, learning, and immune system development. Compared
to other primates, humans grow more slowly, and grow longer during this time, which
appears to reflect increased effort in building and maintaining our brains (summarized
in Chap. 3). For humans, childhood presents a unique opportunity for physical
activity free from energetic trade-offs with direct investment in reproductive effort, or
pressing demands to produce food for oneself or family. The benefits of activity are
also inherently high because physical activity can help to build the coordination,
strength, and skills necessary to survive in a given ecology, learn the energetic
demands of the environment, test individual ability to meet those demands, and foster
play. Play provides opportunities for children to interact with others, developing
critical skills necessary for living in our extraordinarily social species (Gray 2009).
Overall, a life history framework would predict that physical activity should be
high among children, particularly those who are healthy, well fed, and safe. This
perspective would predict that physical activity be reduced during times when
somatic and brain growth are higher. In addition, physical activity may involve
increases in safety risks and injury, and the relative safety of ranging among children
is likely to influence children’s physical activity and cultural norms to mitigate risk
or reduce dangerous forms of physical play. Recall from Chap. 3 that growth is
slower at times when brains uptake the most energy during development (at age
4, Kuzawa et al. 2014). These researchers also noted that physical activity is lower
during times when brains use more energy (among children aged 3–5 years com-
pared to later in childhood). The authors interpret this as consistent with the
hypotheses presented in this book, that physical activity trades off against other
forms of somatic investment and growth. A recent study of macaques in Thailand
demonstrated that locomotor play helped with skill acquisition, but was also related

48 6 Energetic Trade-Offs and Physical Activity …
to less growth, demonstrating the life history trade-off between investment in growth
and investment in developing physical skills through play (Berghänel et al. 2015).
Chapters 2 and 5 highlighted some predictions and interpretation of research
from a life history framework for understanding how resource availability in
developmental environments may influence health later in life through adaptive,
energy conserving shifts in metabolism, immune function, and physical activity. In
addition to environmental cues of resource availability, childhood environments
also vary in exposure to infectious diseases and parasites, which both impact
mortality risks, and thus are likely to have lasting influences on optimal energy
allocation strategies. The energetic budget necessary to fight parasites or infections
impacts how much energy can be put toward other endeavors: immune functions,
growth, reproduction, and physical activity. In addition, where mortality risks are
higher, it is predicted that energy allocation will favor reproducing earlier over
investing in longevity-related somatic investment and longer growth.
Archaeological studies have shown that exposure to viral infections and nutritional
deficiencies during development has a lasting influence on somatic investment and can
even be detected in a diagnostic capacity by the traces left in the bones and teeth
(Goodman and Armelagos 2012). The trade-off between growth and the immune
system was observed among Tsimane’ children between the ages of 2–10 (McDade
et al. 2008). Immune activation was measured by C-reactive protein (CRP), which was
predicted to negatively influence growth over a 3-month period. Children in poorer
energetic condition (with low body fat) and high CRP grew significantly less than
those with low CRP. Among children in better energetic condition, CRP did not
influence growth rates. Exposure to intestinal parasites or helminths has also been
shown to have a lasting effect on the pattern of investment of the immune system that
may favor fighting parasites over other pathogens (e.g., bacteria, viruses, see
Blackwell et al. 2010). Exposure to helminthes is associated with shifts in the T-cell
populations of the immune system toward a TH2-based phenotype as opposed to a
TH1-based phenotype. The TH2-based phenotype is characterized by increases in
immunoglobulin E (IgE) production, and decreases in TH1 and the pro-inflammatory
immune response (the first line of defense of the immune system).
Blackwell et al. (2010) tested whether IgE (a biomarker of the proinflammatory
response indicative of exposure to intestinal parasites or helminths) was related to
CRP and growth among the Shuar forager-horticulturalists in Ecuador. In line with
predictions, they found that IgE was inversely related to CRP. Among children
(particularly those aged 8–15 years), those with higher IgE (and thus greater hel-
minths exposure) were shorter for their age. These studies show the trade-offs
between investment in growth and somatic investment in immune function, and
how these trade-offs are sensitive to resource availability and exposure to parasites
and other pathogens in the environment. Investment in growth and immune func-
tion are important trade-offs to consider when examining energetic trade-offs that
influence physical activity during childhood.
Once children reach puberty, the opportunity costs of physical activity increase
because adolescents are dealt a double whammy of increased energetic trade-offs.
Energy must be allocated to support reproductive maturation, and growth velocity
6 Energetic Trade-Offs and Physical Activity … 49
actually increases around the same time (Baxter-jones et al. 2005; Walker et al.
2006). Given the tremendous increase in trade-offs that emerge as individuals reach
puberty, a life history perspective would predict a decrease in physical activity at
this stage of life in any environment, but particularly those in which physical
activity does not substantially increase the likelihood of reproductive success.
More likely than not, if you are reading this book, you have experienced this
phenomenon first hand. At a certain age, sleeping until noon replaced your desire to
rise with the sun and start playing. It may surprise you that this pattern of decreases
in physical activity and increases in sedentary behavior during pubertal maturation
is not unique to the developed world, but is common among traditional populations
as well. This is exemplified beautifully in a classic ethnographic documentary that
details the life of a !Kung woman named N!ai, when she describes her son by
saying, “…my son was almost a man. He was of the age, the lazy, difficult age
which we call ‘owners of the shade’” (Miesmer and Marshall 1980).
My own preliminary data among the Tsimane’ also demonstrate this pattern.
Physical activity was measured with accelerometers among a cross section of
Tsimane’ children, adolescents, and young adults (aged 8–22 years). Minutes of the
day spent being sedentary significantly increased with Tanner stage of pubertal
maturation (B = −28.66, t = 3.47, p < 0.001) even when taking into account the
effects of age (ns) and sex. Overall, knowing an individual’s Tanner stage, sex, and
age accounted for over half of all the variation in sedentary time (51 %).
Meanwhile, there were significant decreases in minutes of the day spent in moderate
to vigorous physical activity, again controlling for the effects of age and sex
(β = −9.00, t = −2.00, p < 0.05). Together, Tanner stage, age, and sex accounted for
38 % of the variation in moderate to vigorous physical activity (unpublished
results).
This pattern is also consistently seen in large-scale epidemiological studies in
Western populations. In the USA, when physical activity is estimated objectively
with accelerometers and strict cutoffs are used to define moderate physical activity,
less than half (42 %) of children get the recommended amount of activity, but only
8 % of adolescents get the recommended amount (Troiano et al. 2008). In a
European sample, 97 % of children at age 9 get the recommended amount of
physical activity, whereas only 82 % of boys and 62 % of girls get the recom-
mended amount at age 15 (Riddoch et al. 2004). In a longitudinal study in the USA,
most kids at age 9 most met the recommended levels of physical activity, but by 15,
only 31 % met the recommended levels on weekdays and even fewer (17 %) met
the recommended levels on the weekends (Nader et al. 2008).
Studies have consistently shown age-related decreases in physical activity that
occurs earlier for girls than boys (Lopes et al. 2007; Nader et al. 2008; Troiano et al.
2008). Recently, some researchers have shifted the focus more toward examining
biological age or pubertal maturation instead of chronological age to explain the sex
differences in physical activity decreases, since males mature later than females.
Indeed, when pubertal maturation is accounted for, the sex difference in decreases
in activity is attenuated (Cumming et al. 2008; Sherar et al. 2007; Thompson et al.
2003). This pattern is often interpreted as a by-product of the psychosocial stress
that transitioning to young adulthood typically involves, particularly among girls.

Researchers hypothesize that maturing early or late causes more stress and therefore
results in risky and unhealthy behaviors, like not exercising, smoking, drinking
alcohol, and initiating sexual behaviors earlier. However, when studies focus on the
impact of maturing early or late on physical activity, the results are inconsistent and
typically do not support the prediction that maturing early or late consistently leads
to decreases in physical activity (Sherar et al. 2010).
A life history framework offers an alternative, simpler interpretation: Given a
finite energetic budget, increased investment in rate of growth and reproductive
maturation trades off against other allocations, including physical activity. This
perspective requires a paradigm shift from thinking about being lazy or less willing
to be physically active as a pathology or risky health behavior, but rather as a
conservative strategy that conferred reproductive fitness benefits over evolutionary
time. There comes a point when it is more advantageous to invest in reproductive
capacity than running around playing.
Second, while the psychosocial stress that adolescents face is remarkable and
often influences behavior in negative ways, it may not be maladaptive and could
potentially serve as a mechanism to decrease non-essential physical activity.
Depression, for example, which is heightened in early maturing girls specifically
(Sherar et al. 2010), may proximately increase malaise and fatigue, and in turn,
sedentary behavior. But one can hypothesize that the ultimate mechanism is that
depression during this stage actually partially functions to help girls conserve
energy, and decrease investment in non-essential forms of physical activity in order
to redirect energy to growth (particularly of fat stores) that are more likely to have
Darwinian fitness, but not physical fitness benefits.
Does this perspective tell us that no matter what we do, physical activity is
programmed to decrease at this age and there is nothing we can do about it? My
answer is, yes and no. I think decreases in physical activity are somewhat inevitable
at this age; however, this perspective also has the potential to uncover individual,
developmental, and ecological factors that influence the decrease. These factors can
then be leveraged to minimize the impact of this energetic shift so that physical
activity levels are not driven down to levels resulting in deleterious mental or
physical health outcomes. Kwon et al. (2011) recently found that even light levels
of physical activity inversely predicted body fat among older children (>11 year
old) but not younger children. The authors suggest that encouraging light, rather
than moderate to vigorous physical activity at this age, may be sufficient to help
reduce overweight and obesity. Accepting that pubertal adolescents face increasing
opportunity costs to perform physical activity may help shape our expectations and
approaches in ways that increase health and well-being overall. We can also work
to increase the benefits side of the equation by diversifying activities to allow
adolescents to tailor their physical activities to be consistent with social goals.
Activities that help in forming supportive social bonds or those that help to increase
physical self-worth such as yoga, dancing, or taking long walks with friends may
increase the likelihood of participation. Some adolescents will enjoy and thrive in
6 Energetic Trade-Offs and Physical Activity … 51
competitive and intense sports, but opportunities for other types of physical activity
should be readily available and highly encouraged for those who do not.
Given that high levels of physical activity in childhood are related to high levels
of physical activity in adulthood (Friedman et al. 2008), one can hypothesize that
humans learn, through maternal cues and environmental demands during prenatal
and postnatal development: (1) the likely level of resource availability; (2) the
extent to which physical exertion and somatic investment are necessary for survival
and reproduction; (3) risks of parasites and other infections; and (4) the degree to
which their individual physiology is able to carry out and withstand physical
exertion. Individual and environmental variation during development might thus
lead to variation in equilibrium investments in physical activity and somatic
maintenance throughout life. I propose that puberty is a critical point in the life
span for targeting physical activity, because energetic trade-offs between growth,
reproductive maturation, immune function, and maintenance are intensified. This
further highlights the importance of physical activity during prepubertal develop-
ment. The next chapter will examine sex differences in the costs and benefits of
physical activity during the reproductive stage of life.
References
Baxter-jones, A., Eisenmann, J., Baxter-jones, A. D. G., Eisenmann, J. C., Sherar, L., & Sherar, L.
B. (2005). Controlling for maturation in pediatric exercise science. Pediatric Exercise Science
17, 18–30.
Berghänel, A., Schülke, O., & Ostner, J. (2015). Locomotor play drives motor skill acquisition at
the expense of growth : A life history trade-off
Blackwell, A. D., Snodgrass, J. J., Madimenos, F. C., & Sugiyama, L. S. (2010). Life history,
immune function, and intestinal helminths: Trade-offs among immunoglobulin E, C-reactive
protein, and growth in an Amazonian population. American Journal of Human Biology, 22(6),
836–848. doi:10.1002/ajhb.21092.
Cumming, S. P., Standage, M., Gillison, F., & Malina, R. M. (2008). Sex differences in exercise
behavior during adolescence: Is biological maturation a confounding factor. Journal of
Adolescent Health, 42(5), 480–485.
Friedman, H. S., Martin, L. R., Tucker, J. S., Criqui, M. H., Kern, M. L., & Reynolds, C. A.
(2008). Stability of physical activity across the lifespan. Journal of Health Psychology, 13(8),
1092–1104.
Goodman, A. H., & Armelagos, G. J. (2012). In risks mortality populations archaeological, 21(2),
225–243
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Play, 1, 476–522. doi:10.1300/J082v41n02_07.
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levels in childhood and adolescence assessed with accelerometry. Journal of Sports Medicine
and Physical Fitness, 47(2), 217–222.
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B., et al. (2004). Physical activity levels and patterns of 9- and 15-yr-old European children.
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Chapter 7
Physical Activity and Reproductive
Ecology in Adults
In adulthood, energy is no longer required for growth, but the energetic demands of
reproduction increase and vary between the sexes. In this chapter, the research that
helps characterize the sex-specific costs of physical activity is summarized to help
understand the factors that are likely to influence physical activity through adult-
hood. Human reproductive ecology is a subfield of biological anthropology that is
grounded in a life history framework and is particularly focused on understanding
the evolutionary physiology of human reproduction. Peter Ellison and colleagues,
in particular, have focused on the influence of energetics on reproductive function
in women and men (Ellison 1990, 2001, 2003, 2008; Jasienska 2001, 2003). This
research is essential for understanding the costs and benefits of physical activity in
humans of reproductive age. Because the costs and benefits are sex-specific due to
differential investment in reproduction in humans, I will discuss women and men
separately.
Women
As detailed in Chap. 3, women incur substantial energetic costs in order to

reproduce. Natural selection tends to err on the side of caution, favoring mecha-
nisms that ensure reproduction can take place, even under changing circumstances.
For human females, however, a growing body of theoretical and empirical research
in reproductive ecology suggests that women have evolved mechanisms that
decrease the likelihood of conception when facing short-term cues of energetic
stress, in order to make conception more probable in conditions that are more
favorable for meeting the energetic demands of pregnancy and lactation. Studies
have consistently shown that both negative energy balance alone, and high levels of
energy expenditure coupled with neutral or negative energy balance can harm
reproductive function among women. Ovarian function is a key component of
female fecundity, or the biological capacity to reproduce. While ovarian function
54 7 Physical Activity and Reproductive Ecology in Adults
was once considered an on/off phenomenon measured by the presence or absence of

menses, it is now well accepted that the presence of menses does not indicate that
ovulation has occurred, and ovarian function varies along a continuum of fecundity
that can be indirectly measured by levels of ovarian steroids. Low levels of estradiol
during the follicular phase or progesterone during the luteal phase have been shown
to reflect decreased fecundity (Lipson and Ellison 1996). Menstrual cycle irregu-
larities measured by a short luteal phase (<10 days), or by oligomenorrhea (cycle
length between 30 and 90 days) also signal potential disturbances in reproductive
function, or fecundity.
There is strong evidence that female fecundity is sensitive to transient shifts in
energy availability in both Western and non-Western populations, and even in
chimpanzees (Thompson and Wrangham 2008). Low ovarian steroid levels, for
example, have been demonstrated among women in Western populations who are
losing weight, are restraining their intake below appetite, have less stored fat and
lower lipid profiles, and have high energy expenditure through exercise or work-
loads with neutral or negative energy balance (reviewed in Ellison 2008; Vitzthum
2009). The combination of high energy expenditure and low energy intake has a
more pronounced negative affect on fecundity (Ellison 2001). It is interesting to
note, however, that increased workload or physical activity/exercise has been
shown to decrease steroid profiles even when food availability is not limited (e.g.,
Jasienska and Ellison 2004; Jasienska et al. 2006; Williams et al. 2015).
Recently, researchers attempted to tease apart the effects of energy expenditure
alone compared to negative energy balance attained through energy expenditure
and reduced caloric intake on menstrual cycle irregularities in a randomized con-
trolled trial among sedentary women in the US (Williams et al. 2015). In one
condition, participants increased exercise, but had compensatory increases in
energy intake to attain a neutral energy balance. In the three other conditions,
participants exercised and reduced caloric intake resulting in deficits of 15, 30, or
60 % depending on condition. Over three cycles, just 13 % of the women in the
exercise only condition experienced menstrual cycle irregularities, resulting in short
(<10 days) or inadequate progesterone (<5.0 ug/ml) during the luteal phase. In
contrast, nearly all (88 %) of those in the most extreme energy-deficient condition
exhibited irregularities. In addition to demonstrating the dose–response nature of
energetics on fecundity, this study importantly shows the individual differences in
how responsive reproductive systems can be to energetic stress. While some
women had very sensitive reproductive responses, showing menstrual irregularities
even when increased energy expenditure was compensated with increases in diet,
others had rather robust reproductive systems and did not show signs of menstrual
irregularities even in the face of increased exercise and a 60 % caloric deficit.
What are the factors that lead to these differences? It can be difficult to know
where to begin testing proximate predictors to examine these differences. Is it
socioeconomic status? Ethnicity? Education? Social stress? Armed with a life
history theoretical framework, I would predict that reproductive responsiveness to
energetic stress is influenced by a woman’s age, parity, overall health, and immune
system, the energetics of her prenatal and developmental environment, and previous
Women 55
experiences with physical activity and fitness. I would further predict that women
whose reproductive systems are particularly sensitive to energetic stress face higher
costs of performing physical activity, particularly non-obligatory physical activity
than those whose system is more robust and are therefore less likely to engage in
physical activity in the absence of greater benefits relevant for reproductive success
(i.e., mating, current or future parental effort).
In a more traditional setting, the Tamang in Nepal provide a natural experiment
for examining the impact of energetics on reproductive function because they
subsist on small-scale agriculture and pastoralism and have seasonal variation in
physical workloads—from moderately heavy in the winter to very heavy during the
monsoon season. Overall, Tamang women exhibit weight loss and decreased
ovulation between seasons (from 71 % of women ovulating in winter to 38 %
during the monsoon). Notably, however, young women aged 17–23 exhibited
negligible seasonal changes in ovarian function and actually gained weight during
the heavy work season (Panter-Brick et al. 1993). Vitzthum (2009) has hypothe-
sized that acclimation and high consumption levels allow some Tamang women to
buffer against ovarian suppression in the face of high energetic output. It could also
be that younger, non-lactating, nulliparous women, have a larger energy budget to
put toward increasing the likelihood of becoming pregnant, and are therefore less
affected by acute shifts in energy expenditure. Since the data are cross-sectional,
and only look at one year, it may also be an epigenetic cohort effect, where there
was something about the environment during a crucial stage of development for this
group of women that made them less vulnerable to shifts in energetic condition.
A study by Jasienska and colleagues (2006) also demonstrated that energetic
stress during fetal development may affect the sensitivity of women’s reproductive
systems to increases in physical activity or workload. Among a sample of rural
Polish women, size at birth, measured by ponderal index (PI; kg/m3), moderated the
relationship between physical activity and ovarian suppression (measured by
mid-follicular salivary estradiol) such that those who had high PI at birth were
buffered from ovarian suppression at moderate levels of physical activity. On the
other hand, women who experienced a constrained fetal environment, reflected by
low or moderate PI at birth, showed a heightened sensitivity of fecundity to
increased energetic demands. These results support the predictive adaptive response
theory that fetal environments with constrained energetic resources may lead to
adaptive shifts in metabolism or energy allocation that favor energy conservation.
The energy allocation system potentially ‘anticipates’ that increases in physical
activity or workload are likely to result in negative energy balance and would
therefore benefit from temporarily delaying reproduction.
It is not well understood why a high level of physical activity would adaptively
reduce ovarian function in the absence of direct energetic stress; Jasienska and
Ellison have suggested two possible hypotheses to explain why this may be: the
“preemptive ovarian suppression” hypothesis and “constrained downregulation”
hypothesis (Jasienska and Ellison 2004; Jasienska 2001, 2003). The former
assumes that historically, increased energy expenditure would have been reliably
coupled with negative energy balance because of limited food availability.
Therefore, human physiology preemptively decreases reproductive effort in antic-

ipation of negative energy balance. However, it is not clear why a system antici-
pating energetic stress would have been selected rather than a system that responds
to the energetic stress directly. An anticipatory system could reduce fecundity
unnecessarily, resulting in non-negligible reproductive fitness costs. The latter
hypothesis is based on evidence that women who are chronically energetically
stressed reduce their basal metabolic rates in order to allocate more energy to
reproduction (Jasienska 2003). Physical activity, on the other hand, increases basal
metabolism and may therefore prohibit or constrain the redirection of energy to
reproduction. It should be noted, however, that there is no evidence that reducing
basal metabolism is a necessary condition for conception in women who are not in
poor nutritional condition; therefore, reduced fecundity in response to increased
physical activity remains poorly understood. What is theoretically clear, and sup-
ported empirically, is that temporary reproductive suppression in the face of en-
ergetic stress (in the form of increased physical activity and/or decreased dietary
intake) is an adaptive response to hold off on reproducing until conditions are more
favorable for meeting the heavy metabolic costs of reproduction.
Energy balance also influences female reproduction in terms of offspring birth
weight and length of gestation, with smaller birth weight and shorter length of
gestation among women facing undernutrition (Ellison 2003; Kline et al. 1989).
Energetics also modulate the length of time that breastfeeding women do not do not
ovulate, termed lactational amenorrhea. Lactation is extremely energetically costly
for women, estimated to increase metabolic load by 625–700 kcal/day (Butte and
King 2005; Ellison 2003) and typically leads to ovarian suppression. The duration of
lactational amenorrhea varies greatly between and within populations and is thought
to be directly affected by energetic demands and fat stores available for producing
breast milk. The direct role of physical activity in the duration of lactational ovarian
suppression is not well understood and is difficult to tease apart from other factors
that influence lactation, such as frequency and duration of breastfeeding. Women
who have heavy workloads often breastfeed less frequently and begin to supplement
children’s diet with alternative food sources earlier, decreasing maternal levels of
prolactin and reducing the drive for milk production (Panter-Brick and Pollard
1999). Given the high metabolic costs of gestation and lactation for human females,
researchers have hypothesized that this heightened responsiveness to energetic
condition and energetic stress would be adaptive in order to decrease the likelihood
of conception when it is questionable whether the energetic demands of pregnancy
and/or lactation can be met (e.g., Ellison 2001; Jasienska 2003; Vitzthum 2009).
Women are also at a disadvantage for transporting oxygen to working muscles
during exercise because they have a smaller heart and lower filling volume, maximal
stroke volume, cardiac output, and lower blood hemoglobin concentration. Men also
have more lean body mass and a higher proportion of slow-twitch muscle fibers,
which contribute to greater overall muscular strength. Notably, however, when
expressed as relative to lean body mass, women do not differ in strength gains or
increased muscle hypertrophy in response to training (Robergs and Roberts 1997).
Women 57
Evidence suggests that women catabolize more fat during physical activity than
men, in spite of having higher returns to stored adipose tissues, since they are
utilized during pregnancy and lactation (reviewed in Lancaster and Kaplan 2009).
Furthermore, in a study of previously sedentary men and women, exercise changed
metabolism-regulating hormones1 in a direction associated with increased appetite
(i.e., increased ghrelin and decreased insulin), but only in women (Hagobian et al.
2009). Physical activity may therefore increase required food intake, or perceptions
of required food in take, specifically among women. In an environment of limited
food resources, as was experienced for humans over the majority of human history,
this would have been a substantial cost to all physical activity, but particularly any
inessential physical activity. In modern environments, this may also make it more
difficult for women who are using physical activity or exercise as a means to lose
weight, as they are also likely to experience increases in appetite and have a more
difficult or unpleasant experience when reducing caloric intake.
Thus, numerous lines of evidence suggest that both empirically and theoretically,
it is advantageous for human females to be more conservative with physical activity
whenever possible during reproductive years to divert more energy to pregnancy
and lactation. This coevolved with humans’ unique pattern of sociality involving
pair-bonding, sex-specific divisions of labor, extended family food sharing, and
cooperative breeding social networks (discussed in Chap. 3).
Not surprisingly, this pattern of physical activity is observed in modern contexts
as well. Physical activity decreases as women get married, become pregnant, or are
postpartum (Brown and Trost 2003; Fell et al. 2009). A life history perspective
helps us to understand why this is the case and provides novel ways to help women
combat becoming sedentary during this critical period with higher costs of physical
activity. One study noted that while moderate and vigorous activity decreased
during pregnancy and postpartum, walking did not (Pereira et al. 2007). Walking
and other activities of a lower intensity may be preferable for women at this time
when competing energetic demands and opportunity costs for physical activity are
so high. Consistent with life history theory, the women in this study who reported
having other children in the home, having long work hours, and lacking childcare
during physical activity, became less active during and after pregnancy. Activities
that are compatible with childcare or more childcare opportunities that women are
comfortable with can help reduce the costs of physical activity for reproducing
women. Another study held focus groups to talk to working mothers and fathers
about the barriers and facilitators of physical activity. The barrier listed most fre-
quently was family responsibilities, while what facilitated exercise the most was
participating in an activity that was consistent with childcare responsibilities. One
mother described how she anticipated the trade-off in advance: “I work full-time…I
1
The thyrotrophic axis and endocrine pathways associated with metabolic regulation and energy
intake specifically are also likely to be essential for a complete characterization of the hormonal
mediation of energy allocation decisions related to physical activity. They were not included in this
book because much less research has focused on these hormones relative to life history parameters,
and they are primarily examined in terms of maladaptive responses to current energy conditions.
knew I was gonna have to give something up…and exercise was it.” (Mailey et al.
2014, p. 660).
These studies show that, while the ultimate motivation that drives this trade-off is
not necessarily processed on a conscious level, women are consciously and clearly
perceiving relevant trade-offs, and unfortunately for public health, appropriately
weighing the costs and benefits to ‘give up’ activities that are less likely to increase
reproductive success. This in no way implies that women are consciously motivated
to increase reproductive success, but rather that priorities have been shaped by
natural selection to value investing in offspring over expending energy on
non-obligatory forms of physical activity, particularly when offspring are so young
and need heavy investment from their mothers. This perspective can inform even
very inexpensive approaches for helping make physical activity more likely among
reproductive aged women by changing the way messages are framed. Instead of
sending the message to exercise with current recommendations, the message could
be tailored to allow women to scale back expectations and instead encouraging them
to keep up taking a walk every day, or finding creative ways to make physical
activity obligatory, more fun, or consistent with pregnancy and childcare. The
message can express that it is normal for physical activity to feel more difficult
during this time, or to weight barriers (particularly those related to reproduction,
provisioning, and childcare) more heavily because it was adaptive to have physi-
ological and psychological mechanisms that favor energy conservation in women of
reproductive age, particularly pregnant women and new or overly burdened mothers.
Framing benefits in terms of how physical activity can help women invest more in
offspring or fostering a social network at the same time as increasing physical
activity may also be more likely to resonate with pregnant women and mothers.
Men
Men’s reproductive physiology, in contrast to women’s, is generally less energet-

ically costly and more robust in the face of reductions in energy availability or
increases in energy expenditure. The quantity and quality of sperm production
appear to be insensitive to short-term changes in energy availability (Bribiescas
2006). Moreover, physical activity is largely compatible with men’s reproductive
effort, increasing their ability to compete with other males for access to mates, and
in environments with strenuous demands for resource production, provisioning
wives and children.
In addition, research in modern contexts shows that investment in fitness and
muscle mass actually increases men’s access to mates (Frederick and Haselton
2007; Gallup et al. 2007; Hönekopp et al. 2007; Lassek and Gaulin 2009). While
men do not appear to face nearly the same trade-offs between physical activity and
reproductive investment as women, variation in physical activity may still be
affected by resource availability and individual condition. Kuzawa and colleagues
(2010) observed that the rate of weight gain within 6 months after birth (a measure
Men 59
of early developomental resource availability) predicted metabolically costly

somatic characteristics such as reproductive hormones, adult size and strength, as
well as sexual activity among Filipino males. Those with rapid early growth
reached puberty earlier, were taller and more muscular, had higher T levels, greater
grip strength, had sex at earlier ages, and had more lifetime sexual partners than
those who did not grow rapidly during infancy. Lassek and Gaulin (2009) explicitly
examined the costs of musculature in terms of increased energy intake and
trade-offs with immune function in the USA. They found that free fat mass and limb
muscle volume were positively related to food intake and were inversely related to
two proxies of immune function: CRP and white blood cell counts.
Testosterone (T)—which modulates muscle mass, as well as allocations to
mating effort (Burnham et al. 2003; Ellison and Gray 2009; Gray et al. 2007;
Kuzawa et al. 2009)—appears to be unresponsive to short-term changes in energy
availability, but sensitive to longer-term energetic deficit (Ellison 2003). T has also
been shown to be negatively related to immune function (Muehlenbein and
Bribiescas 2005; e.g., Muehlenbein et al. 2010). Some researchers have argued that
investment in muscle mass can be thought of primarily as somatic effort aimed at
reproduction by increasing mating opportunities (Bribiescas 2001, 2006). More
recently, others have pointed out that somatic investment in musculature is also
required for paternal investment through provisioning and protecting partner and
offspring from external threats (Lancaster and Kaplan 2009). Long-term energetic
deficits and environments with constrained resources, heavy pathogen or infection
risk, or a burdened immune system are all predicted to result in lower levels of T,
which may therefore hinder men’s reproductive effort in both mating effort and
provisioning for offspring. However, Alvarado and colleagues (2015) recently
demonstrated that T was not correlated with strength or musculature among a
sample of rural Polish subsistence agriculturalists who regularly engage in stren-
uous manual labor. The authors propose the Paternal Provisioning Hypothesis
which predicts that in humans, muscle is more sensitive to physical workloads than
androgens (i.e., T, as in other primates), allowing for lower T levels to foster
pair-bonding and paternal investment while simultaneously maintaining muscle
mass by regularly performing physically demanding resource production. In this
case, energetic stress would not necessarily influence men’s mating or paternal
investment significantly. This may also influence the ability for humans to maintain
muscle in the absence of activity. Recall from Chap. 3 that compared to other
primates, humans lose muscle mass rapidly in response to decreases in use.
The opportunity costs of physical activity for men increase substantially, how-
ever, in environments where provisioning for offspring requires sedentary rather
than strenuous physical labor. Physical activity is therefore predicted to be lower
among committed, mated men with children in these environments, but men
focused on mating rather than parenting effort are still expected to favor somatic
investment in muscle mass in order to attract mates. As with females, it may be
possible to increase physical activity among mated fathers in these more sedentary
economies by reframing it to be more compatible with providing for children. This
point is exemplified by the following quote from a regularly exercising father in the
focus group of working parents discussed earlier, “Now that I have a family I’ve
gone away from trying to be as big and strong as possible to, I want to be as healthy
as possible. I want to be able to be there, and be able to pick them up and go play
with them and stuff like that. To have that energy, so that is my motivation” (Mailey
et al. 2014, p. 661).
Post-reproductive Age Men and Women
Another unique characteristic of the human life span is that humans have a long
period of life after reproduction ceases among women (i.e., after menopause). In
contrast, most other animals continue reproducing until close to the end of their
lives. Some men have children with other partners after their partners reach
menopause, but it is far more common for men to have their last child at the same
time their partners do, and thus similarly have an extended post-reproductive life.
This is thought to have evolved because humans exhibit returns to production late
in life, an extended period of juvenile dependency, and high maternal physiological
demands of reproduction. Help from partners, parents, and in-laws allows human
females to decrease production during pregnancy and lactation when the demands
for resources go up (Hooper et al. 2015; Marlowe 2010). The costs and benefits of
physical activity in post-reproductive individuals are theoretically predicted to be
influenced by (1) how much grandparental investment in offspring and
grand-offspring is dependent on physical activity, as well as (2) how much
grandparental investment significantly contributes to offspring and grand-offspring
survival and reproduction. Messages about the long-term health effects of physical
activity are likely to be much more salient and effective at increasing physical
activity during this stage of life. At earlier stages, these messages are unlikely to
have much of an effect, particularly among those with a more energetically con-
servative life history strategy that does not favor investments in longevity.
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Part III
Summary of Proximate Mechanisms,
an Integrated Evolutionary Model
and Applications
Chapter 8
Proximate Mechanisms: Psychology,
Neuroendocrine System, and Central
Nervous System
An evolutionary, life history framework can provide useful interpretation of pre-

vious findings in psychological and epidemiological research on the proximate
mechanisms that promote physical activity. It can also allow for innovative and
unique hypotheses to be generated that can be tested in future research. Basic
science studies can test hypotheses about the psychological and physiological
mechanisms that influence physical activity. Hypotheses can be tested to determine
whether physical activity behavior reflects predicted adaptive trade-offs between
competing energetic demands that influence the costs and benefits of performing
obligatory and non-obligatory physical activity. It can also lead to predictions about
factors that moderate energy allocation strategies such as individual condition, life
stage, developmental and current environments, and epigenetic changes. Finally,
intervention and public health researchers can test whether manipulating the rele-
vant and malleable costs and benefits in favor of physical activity is possible, and
truly leads to appreciable and sustained increases in physical activity. The following
sections will flesh out some avenues for research that are possible from an evolu-
tionary and life history framework.
Interpreting Previous Findings: Why do Psychological

Variables Matter Sometimes but not Others?
It is beyond the scope of this book to include all the relevant examples of psy-
chological predictors of physical activity. Instead, I will highlight two relevant
examples of how an evolutionary and life history framework can lead to a mean-
ingful interpretation of previous findings.
Researchers have examined whether physiological factors may mediate or
moderate the relationships between psychological predictors of physical activity
and exercise. In one study, researchers tested the relationship between self-efficacy

66 8 Proximate Mechanisms: Psychology, Neuroendocrine …
(or confidence in performing physical activity) and physical activity among normal
weight and overweight/obese individuals. The relationship between self-efficacy
and physical activity was moderated by weight status, such that self-efficacy pos-
itively predicted physical activity among normal weight participants, but not among
overweight or obese individuals (Blanchard et al. 2005). A life history perspective
can shed light on why physiological factors, such as weight status, may moderate
the extent to which self-efficacy was related to physical activity. I would expect that
psychological variables will predict physical activity, but only when the net costs
and benefits of physical activity are equal.
Increases in weight inherently increase the metabolic costs associated with
physical activity. In addition, heavier individuals have a more difficult time
responding to heat stress, which at the very least increase the perceived costs of
physical activity. The ability to dissipate heat declines with greater body mass, and
heavier subjects are less able to handle increases in metabolic heat production
(Leonard 2015). For the net costs and benefits of physical activity to be equal
between normal weight and overweight/obese individuals, there would need to be
compensatory decreases in competing costs (e.g., compatibility with investing in
children), or relevant adaptive benefits. When net costs and benefits are unequal,
psychological variables are expected to have unequal influences on behavior. This
example also demonstrates how a negative feedback loop can begin once people
gain weight. Physical inactivity can lead to weight gain, and the inherent metabolic
costs of physical activity get higher with additional weight, making it that much less
likely to occur.
McCormack and colleagues (2010) also showed that some ecological variables
influenced behavior through cognitive processing, while others influenced behavior
directly. In their study, perceived behavioral control (i.e., self-efficacy) mediated the
relationship between access to facilities and physical activity, while crime rate
influenced behavior directly. Considering the adaptive function of strategic energy
allocation offers a simple explanation for why this pattern was observed. Concerns
about safety are particularly detrimental to survival and reproduction, and thus,
where safety is a concern, this will supersede cognitive appraisals of how capable
an individual is to be active.
Neuroendocrine System as Potential Mechanism
In the human evolutionary sciences, life history theory has been widely applied for
understanding differences in the timing of crucial events across the life span (e.g.,
mortality, reproduction, senescence). Hormones play a central role as physiological
mechanisms that coordinate the allocation of energy across competing demands and
can influence behavior. Research that aims to apply a life history perspective is
increasingly including the measure of hormones as objective measures of physio-
logical investment in growth, immune function, reproduction, mating, pair-bonds,
cooperation, parenting (Finch and Rose 1995; Stearns 1989; Trumble et al. 2015;
Neuroendocrine System as Potential Mechanism 67
Worthman and Barrett-Connor 2002; Zera and Harshman 2001). As Worthman and
Barrett-Connor eloquently state:
Hormones establish the short-and long-term balance of resource allocation between growth,
reproduction, and maintenance. Hormones juggle net energy availability by modulating
metabolism and setting internal regulatory parameters, they regulate the rate of growth and
the timing of developmental transitions such as puberty, and they dynamically manage the
interface between the individual and environment by orchestrating responses to everything
from stress to workload. (p. 198)
Rather than provide a summary of all potentially important neuroendocrine

mechanisms for understanding physical activity, this section focuses on hormonal
pathways associated with the HPG and HPA axes that are likely to play funda-
mental roles in regulating life history allocations relevant to physical activity in
both men and women.
Hypothalamo-Pituitary-Gonadal (HPG) Axis
The hypothalamo-pituitary-gonadal (HPG) axis controls the onset of reproductive

maturation, maintains and regulates reproductive function in adults, and regulates
reproductive senescence (menopause) in women (Worthman and Barrett-Connor
2002). These functions are carried out through the hypothalamic pulsatile secretion
of gonadotropin-releasing hormone (GnRH), which stimulates the anterior pituitary
to release follicle-stimulating hormone (FSH) and luteinizing hormone (LH). FSH
and LH then stimulate gonadal steroid production, principally estradiol and pro-
gesterone in women, and testosterone in men. This cycle is regulated by the neg-
ative feedback of circulating ovarian steroids on GnRH production. The following
two sections will summarize the sex-specific aspects of the HPG axis in women and
men that are relevant for measuring energy allocation to reproduction across the life
span and are also influenced by physical activity.
Women: In women, the HPG axis is briefly active after birth, followed by a long
period of inactivity until puberty. After several years of non-fecund or sub-fecund
cycling, the axis attains full maturity and performs cyclical ovulation, except during
pregnancy and lactational amenorrhea (Worthman and Barrett-Connor 2002).
A typical menstrual cycle is characterized by rising levels of FSH, peaking during
the mid-follicular phase, a gradual decrease prior to ovulation, a small hump right at
ovulation, and a slight, gradual increase throughout the luteal phase. LH, on the
other hand, is low during the follicular phase, surges and peaks right around
ovulation, and gradually decreases through the luteal phase.
Estradiol (the main human form of estrogen) is produced in the ovaries in
response to both FSH and LH. In general, estradiol increases steadily during the
follicular phase, with a surge and peak just prior to ovulation, and then decreases
and remains at a low but non-negligible level during the luteal phase until the onset
of menses. Progesterone is similarly produced in the ovaries in response to LH, is

low during the follicular phase, and exibits a post-ovulatory surge and peak, fol-
lowed by a gradual decline until the onset of menses. Progesterone levels remain
high if conception has occurred at ovulation, and remain elevated during pregnancy
due to the embryonic, fetal, and placental production of human chorionic gona-
dotrophin (hCG). Reproductive senescence in women occurs over several years
during the fifth or sixth decade upon follicular depletion, which marks the begin-
ning of the uniquely human, post-reproductive period in life.
Compared to our closest primate relatives, human females have much more
discrete ovulation and lack obvious estrual swellings or dramatic shifts in sexual
behavior. While it was once thought that human ovulation was completely con-
cealed (Daniels 1983), this assumption has recently been called into question by
research exploring less overt, but still measurable changes in women’s and men’s
mating preferences and behavior across the cycle (Thornhill and Gangestad 2008).
This growing body of research supports two hypotheses: First, that women have
shifts in mating behavior and preferences across the cycle (Gangestad et al. 2005;
Garver-Apgar et al. 2006; Haselton and Gangestad 2006; Pillsworth et al. 2004);
and second, that there are cues of increased fecundity that are perceptible by males
(Miller et al. 2007; Miller and Maner 2010). These changes are theoretically linked
to changes steroid hormones; however, due to the difficulty in collecting estradiol
and progesterone across the cycle, hormones are often merely estimated based on
actuarial data and self-reported cycle day. The exact hormonal role in these changes
remains unclear.
Physical activity, in conjunction with overall energy balance, affects the func-
tions of the HPG axis in women. As summarized in Chap. 7, female age of
reproduction, ovarian function and fecundity, and lactational amenorrhea are all
sensitive to energetic constraints and demands. Therefore, female activity patterns
are intricately linked to life histories. The mechanism through which energetic
condition influences ovarian function is not well understood, but is theoretically
viable through either a central or peripheral pathway (Ellison 2001). The central
pathway model would involve communication between the energetic condition and
the HPG axis, which affects pulsatile GnRH production, and in turn, the hourly
pulses of LH. A peripheral endocrine pathway—where energetic condition is
communicated through changes in adrenal and metabolism-regulating hormones
such as cortisol, insulin, growth hormone, or leptin, and/or their receptors (de-
scribed more fully in detail below)—may be more likely. Further, a combination of
both peripheral and central pathways may also influence the relationship among
energy expenditure, balance, and ovarian function (Ellison 2001).
Ovarian steroids are associated with acute and chronic physical activity in
various ways. Estradiol and progesterone increase acutely in response to exercise,
absent from increases in their trophic hormones FSH and LH (Bonen et al. 1979).
Estradiol, in turn, increases the mobilization of free fatty acids from adipose tissue
and inhibits glucose uptake by peripheral tissues, indirectly favoring catabolic
metabolism. Progesterone is known to increase ventilation during exercise (Robergs
and Roberts 1997). On the other hand, chronically higher levels of physical activity
in the forms of habitual physical activity and exercise are inversely related to
estradiol (Jasienska et al. 2006; Robergs and Roberts 1997).
In a sample of Polish women, estradiol levels were also quadratically related to
body fat, with both very low and very high levels of body fat associated with
significantly lower levels of estradiol than women with a moderate amount of body
fat (Ziomkiewicz et al. 2008). If having high levels of body fat prohibits repro-
duction in women and/or men, people with moderate amounts of fat will have
greater reproductive success, and phenotypes with very high fat will be selected out
over time; but natural selection is a slow process, and this probably the first time in
human history that natural selection will have the chance to act on extreme adi-
posity that hinders reproduction.
As discussed in Chap. 7, high levels of physical activity are associated with
lower ovarian steroid profiles, though this is not universally true. Some women are
more sensitive to increases in energy expenditure, exhibiting menstrual cycle
irregularities even when energy expenditure is paired with compensatory increases
in caloric intake. Other women do not have exhibit menstrual cycle irregularities
even in the face of increased energy expenditure and decreased caloric intake
(Panter-Brick et al. 1993; Williams et al. 2015). Future research can examine factors
that are predicted to moderate this relationship, such as individual condition, age,
and parity (whether she already has other children), and examine whether women
who’s reproductive function is more robust to energy demands of physical activity
are more likely to do it. This would support the hypothesis that there is a trade-off
between allocating energy to physical activity and reproductive effort.
Men: In human males, the HPG axis follows a similar life course pattern as in
females until puberty. Once reproductive maturity is reached, men’s HPG axis
steadily functions throughout adulthood, without cyclic variation, and decreases
during senescence, but does not fully cease functioning. FSH secretion leads to the
production of testosterone (T) in the testes in men.
The regulation of male traits by T levels has pronounced implications for the
evolution of not just human, but all vertebrate life histories (Hau 2007). As outlined
in the previous section, T levels have long been understood in terms of investment
in future reproductive opportunities. A growing body of research has documented a
link between testosterone (T) levels and monogamous relationship and/or paternal
status among men in various populations. On average, men with children and those
in monogamous relationships have lower testosterone than single men (Alvergne
et al. 2009; e.g., Burnham et al. 2003; Gray et al. 2002, 2007; Kuzawa et al. 2009).
Though most studies have been cross-sectional, longitudinal research (e.g., on
divorced men or new fathers) generally suggests that T levels change with rela-
tionship status or paternal circumstances (Berg and Wynne-Edwards 2001; Mazur
and Michalek 1998; Storey et al. 2000). These findings are consistent with data
from several other species with high paternal care and/or extensive pair-bonding
(e.g., Clar and Galef 1999; Nunes et al. 2001).
Exceptions to the general pattern have been documented, but even they fit a
conceptually meaningful pattern. For instance, Muller et al. (2009) compared T
levels of fathers and non-fathers in two neighboring groups in Tanzania, Hadza

foragers and Datoga pastoralists. They found the typical pattern of lower T levels in
fathers than non-fathers in the Hadza, but not the Datoga. Datoga fathers, however,
are minimally involved in direct child-care, and hence, there may be less oppor-
tunity for paternal status to affect T levels in that group.
Sociosexual orientation (measured by the sociosexual orientation index (SOI), is
the quantification of interest in uncommitted, and/or extrapair sex) has been shown
to moderate the relationship between men’s T and relationship status in Western
samples. Specifically, partnered men who are relatively open to and interested in
mating opportunities outside their current relationship (those having unrestricted
sociosexual orientation; Simpson and Gangestad 1991) have higher T levels than
partnered men with more restricted SOI scores (McIntyre et al. 2006). In fact, these
men show no evidence of reductions in T level as a function of relationship status.
In this study, only men specifically committed to remaining faithful to their partners
experienced lower T levels.
Similarly, Gray (2003) found that while Swahili men had slightly lower T levels
when they had young offspring, married men did not have lower T levels than
unmarried men. This is likely due to the high degree of polygyny in this culture,
such that marriage did not necessarily reduce men’s mating effort, a finding sup-
ported by the greater T levels in polygynously married men. Alvergne et al. (2009)
also found higher T levels in polygynously mated versus monogamously paired
men, with T levels relating to relative investment in parental care versus mating
effort. Gray et al. (2007) did not find this result among Ariaal pastoralists in
Northern Kenya. They noted, however, that in this population, men with multiple
wives were older, and thus likely to have lower T levels, as they had to accumulate
enough wealth to acquire multiple wives.
Differences across individuals and populations in commitment to monogamous
pair-bonds thus may, as with paternal investment, modulate the association between
relationship status and T levels in a meaningful way. It is worth noting that T levels
may be necessary for men devoted entirely to parental investment for the devel-
opment of physical strength in subsistence activities, handling dangerous domestic
animals, and protecting women and children from outside threat, for example
(Lancaster and Kaplan 2009). As discussed earlier, however, Alvarado et al. (2015)
found that in a rural sample of Polish farmers who regularly performed physically
demanding agricultural labor, strength was not related to T levels, which were
lower than typically observed in Western samples. The authors hypothesize that
Polish men’s musculature is maintained through regular use, and similar to other
primates, is not dependent on T levels.
Testosterone, just as progesterone and estradiol, is known to increase acutely in
response to physical activity, and this increase is unrelated to increases in LH.
Further, T levels are lower in response to longer bouts of endurance exercise and in
highly active men (Robergs and Roberts 1997). T levels aid in protein synthesis in
muscle tissue and therefore lead to increased musculature, strength, and lean body
mass. As noted in Chap. 7, while men are stronger than women, due to their higher
proportion of lean body mass, there are no sex differences in strength and power
when expressed as relative to lean body mass (Robergs and Roberts 1997). T levels
are often thought of as somatic investment related to increasing mating effort,
mediating life history allocation to future reproduction (Bribiescas 2001; Ellison
2003). A recent study found evidence partially supporting this idea among a sample
of Filipino men. Interestingly, T levels predicted lean body mass and strength, but
only among men who were active, suggesting that T levels mediate investment in
costly somatic traits, specifically in conjunction with environmental interaction
demanding regular muscular use (Gettler et al. 2010).
Hypothalamo-Pituitary-Adrenal (HPA) Axis
In response to both physiological and psychological challenges, the hypothalamo-

pituitary-adrenal (HPA) axis is activated, and adrenal hormones adaptively shift
energy allocation by mobilizing resources to respond both physiologically and
behaviorally to the challenge presented. The fast acting, sympatho-adrenal-
medullary axis organizes the immediate endocrine response, releasing adrenal
neurotransmitters or catecholamines (epinephrine and norepinephrine) within mil-
liseconds, which leads to a rapid increase in heart rate and heightened arousal,
facilitating vigilant behavioral reactions (Worthman and Barrett-Connor 2002).
This response is costly, and continued or prolonged activation can result in
hypertension and functional impairment. A slower hormonal cascade begins with
corticotropin releasing hormone (CRH) produced in the hypothalamus. CRH pro-
duction stimulates the anterior pituitary to release adrenocorticotropic hormone
(ACTH), which in turn results in adrenal production of cortisol. Cortisol is often
thought of specifically as a stress hormone; however, when applying a life history
approach it is more useful to appreciate the broader function of cortisol as mediating
physiological energy allocation from long-term (growth, reproduction) to
short-term (vigorous activity, alertness) functions (Worthman and Barrett-Connor
2002; Worthman and Kuzara 2005). CRH is produced in the hypothalamus in
response to stimuli and in circadian rhythms. Cortisol is a glucocorticoid that
promotes immediate survival needs at the expense of long-term processes: It pri-
oritizes increases in blood glucose at the expense of protein and fat stores and
vigilant focus, while reducing food intake, HPG activity, gut motility, growth,
memory formation, and alters immune activity (increasing leukocytes in the short
term and decreasing immune function if elevated over a prolonged period (Robergs
and Roberts 1997; Sergstrom 2010; Sherwood 2001).1
1
It should be noted that HPA also regulates dehydroepiandrosterone (DHEA) and its sulfite
DHEAS, the predominant steroid hormone. While potentially important for life history and
physical activity, much remains unclear about its actions and responses to physical activity and
thus was excluded from this book. For more information, see Worthman and Barrett-Connor
(2002).
Through the life span, the HPA axis has regular diurnal periodicity by 6 months
of age, with little age-related change until activity gradually increases from third
decade on. As reviewed in the previous section, stress exposure and high levels of
helminths can be detrimental to growth and reproductive function in both men and
women. Women appear to have adaptive mechanisms to allow reproduction to
resume if stress exposure is chronic, as well as buffer a growing fetus from ener-
getic stress. However, stressful environments or experiences may alter stress
reactivity that change the extent to which individuals can handle the stress of
physical activity.
The HPA axis responds acutely to physical activity, with the catecholamines and
cortisol increasing as intensity increases (Robergs and Roberts 1997; Worthman
and Barrett-Connor 2002). These increases stimulate the mobilization of free fatty
acids that can be used by the liver for gluconeogensis. Given that liver gluconeo-
genesis is not needed during short-term exercise, it is thought that acute increases in
cortisol are particularly helpful for recovering during short-term exercise. In
addition, circulating amino acids are increased through reducing the uptake of
amino acids by peripheral tissues. Thus, exercise and physical activities are phys-
iologically experienced as a stressor. While people are often told that physical
activity will feel very good, there is also a lot about it that feels really bad. There are
individual differences in stress reactivity that may make it even more aversive for
some people, thereby increasing the costs. Future research can examine the factors
that lead to differences in stress reactivity (post-traumatic stress disorder, epigenetic
change, etc.) to test if those who are more reactive to stress are less active, find it
more aversive, perceive higher costs, or have a more negative psychological
response. If individuals who have a hyperactive HPA or sympathetic nervous
response to stress indeed have a more negative psychological response to physical
activity, they may be more successful at initiating exercise if they begin with
regular exercise at lower intensities, or with approaches that help stress reactivity
like yoga, or mindfulness meditation (Ross and Thomas 2010; Tang et al. 2009). In
addition, many sport and exercise researchers examine the conditions under which
people focus their attention inwardly toward body sensations (associative) or out-
wardly and away from body sensations (dissociative) in order to cope with energy
expenditure (reviewed in Masters and Ogles 1998). Individuals with a hyperactive
HPA may be more likely to initiate or increase physical activity or exercise if they
can use a dissociative coping strategy with their minds otherwise occupied, such as
watching television or listening to music. The primary goal would be to help
individuals find the activity or coping strategy that decreases the perceived stress (or
costs) of energy expenditure given individual differences in stress response to
energy expenditure. Training can also diminish this HPA hyperactivity (Luger et al.
1987), but getting these individuals to train enough to reach this point may prove
difficult. A heightened stress response to exercise may also contribute to the dif-
ficulties in beginning exercise in adults who did not exercise earlier in their lives.
The accumulation of experiences with physical activity as a stressor throughout
development, termed habituation, may dampen the physiological stress response to
exercise, which would in turn reduce energetic costs by not mounting an alarmed
HPA response in the face of common or unavoidable stressors in a given ecology.

This is similar to the California ground squirrel’s stress response to snakes, which
has been shown to depend on environmental exposure to the predators. Snakes are
long-term, formidable predators to ground squirrels, who have thus evolved rather
gutsy antisnake behaviors including biting, pouncing, and other forms of
“close-range harassment” (Coss and Biardi 1997, p. 295). In a quasi-experiment,
researchers measured squirrels’ stress and behavioral response to rattle and gopher
snakes. Squirrels from environments where the snakes are abundant and encounters
frequently have a lower physiological stress response than squirrels from envi-
ronments without rattle or gopher snakes. A decreased stress response likely helps
squirrels in tactical decision making for the necessary, yet dangerous antisnake
behaviors. For humans, this underscores the importance of consistently engaging in
physical activity throughout development, because not doing so likely results in a
heightened HPA response to an unfamiliar stressful stimulus, thereby increasing the
energetic costs of physical activity and decreasing the likelihood of the behavior.
Relative to the HPG axis, the HPA axis is largely similar in men and women. Sex
differences in adrenal response to ACTH emerge at puberty, however, with more
cortisol produced in women in response to the same level of ACTH. The mecha-
nism driving this difference remains unclear, though it is likely to occur through
different ACTH delivery to the adrenal gland or a heightened sensitivity or
responsiveness by the adrenal gland to ACTH stimulation.
The HPG and HPA axes demonstrate the ways in which hormones and endo-
crine pathways that mediate life history traits are affected by, and exert influence on
physical activity. Furthermore, there are hormonal and other physiological sex
differences that modulate the costs and benefits of physical activity for men and
women. Elucidating the bidirectional relationships between the ecological and the
individual factors that influence life history energy allocation, the hormonal milieu
necessary for carrying out the energy allocation, and physical activity can help lead
to a better understanding of the ways in which environmental, physiological, and
psychological factors influence individual physical activity behavior. A more clear
understanding of each of these factors can lead to the creation of more successful
ways of increasing physical activity in modern, primarily sedentary environments.
Central Nervous System and Epigenetics as Potential

Mechanisms
The central nervous system is another potential proximate mechanism that functions
to weigh the costs and benefits of physical activity at an individual level. This is
partially possible through how the stress and exhaustion of physical activity are
perceived. There is not a clear consensus among exercise physiologists about what
causes fatigue during exercise. It was initially thought that fatigue happened
peripherally in the muscles, but central pathways have recently received much more
attention. Central hypotheses predict that changes in the brain reduce neural drive or
have a protective “governor” that functions to prevent absolute fatigue and damage
to organs (St Clair Gibson and Noakes 2004). The central nervous system may also
play a critical role in strength gains by controlling how many muscle fibers are
recruited in order to perform an action, making effort more or less efficient (Folland
and Williams 2007). Future work can examine if these two processes are influenced
by life history stage, environments, and individual condition, health and/or fitness
and therefore make physical activity more or less costly. Those who are in poorer
condition, have higher parasite burden, have less previous experience with activity,
or are in harsher environments would be predicted to have more conservative
central pathways (whatever they may be) than those who were in better condition
currently, or who did not face energetic or immune stress during development.
Another nascent but exciting area to consider is ways epigenetic changes may
influence individual differences in the costs or perceived costs of physical activity.
Epigenetic changes are potentially heritable changes in gene function that do not
change an individuals genetic makeup, but whether or not certain genes are “turned
on.” They are sensitive to environments and behavior. It is an empirical question for
future research whether environmental exposures, particularly in resource avail-
ability, may actually lead to inherited epigenetic alterations that affect energy
allocation optimization in future generations.
Summary
An evolutionary and life history perspective provides a larger framework to con-

textualize the ways physical activity is influenced by trade-offs in energy allocation
that have been shaped by natural selection to maximize predicted reproductive
success. In light of this perspective, and the research presented in this book, I
propose a model (depicted in Figs. 8.1 and 8.2) to characterize the interrelationships
between the complex factors that determine the costs and benefits of physical
activity across the life span in humans and influence behavior.
At a population level, natural selection has shaped phylogenetic and species
level differences in behavior and life histories over evolutionary time. This process
has shaped humans’ genetic makeup, which in turn influences individual condition
and developmental trajectories, as well as the proximate mechanisms that make
energy allocation “decisions,” and result in behavior was adaptive in a wide array of
environments. Natural selection has not shaped behaviors that are adaptive in
environments with an overabundance of calorically dense, easy to acquire resour-
ces, because such environments did not exist until recently. Reaction norms have
been shaped by individual and environmental conditions, particularly the
Summary 75
Fig. 8.1 Integrative evolutionary model that assimilates the ultimate (including phylogenetic),
proximate, developmental, and environmental factors that collectively influence physical activity
(represented as “Behavior” in the center of the model). Temporally, the model begins on the
bottom left. Ancestral environments provided the selection pressures that shaped the evolution of
factors that influence physical activity within a species and a population. On an individual level,
genes together with epigenetic and environmental factors shape the developmental trajectory of
energy allocation throughout the life span. This trajectory, individual condition, and the life stage
of an individual at a given moment in time influence the costs and benefits of physical activity. The
proximate mechanisms are hypothesized to reflect the costs and benefits of physical activity and
are also influenced by current environmental conditions. Proximate mechanisms directly influence
behavior. The model is intentionally circular, because behavior can also shape the evolution of a
species when it influences reproductive success. It can also lead to epigenetic changes and changes
to individual condition and the proximate mechanisms that influence the behavior, making it more
or less likely to happen subsequently
availability of resources and risks of pathogens and other infections, to establish

energy allocation strategies that influence the rate of life history progression and the
costs of physical activity. Current environments also influence the costs and ben-
efits of physical activity through determining the amount of energy required to
extract resources and provide for oneself and offspring. In modern contexts, this
includes the opportunities or necessity for physical activity in the built environment.
Cultural variation can also influence the benefits of physical activity, where social
and cultural values encourage more physical activity.
This model is not exhaustive, but it can serve as a heuristic to help synthesize the
complex web of ultimate and proximate factors that influence physical activity. It
provides causal paths that can be tested empirically in the service of a more inte-
grated and comprehensive understanding of the factors that influence physical
activity throughout life.
Fig. 8.2 Provides a non-exhaustive list of possible variables to include in order to measure
individual and environmental factors that influence the proximate psychological and physiological
mechanisms that directly affect physical activity
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Chapter 9
Summary, Conclusions, and Applications
This book advocates the basic thesis that the physiological and psychological
mechanisms that influence physical activity have been shaped by natural selection
to flexibly respond to energetic demands and resource availability, in order to
optimize survival and reproduction within a given ecology. This system was, for the
vast majority of human evolution, consistently constrained by resource scarcity, and
in response, mechanisms have evolved to decrease energy expenditure adaptively in
response to decreased intake, early cues of nutritional constraints, or fewer adaptive
benefits of expenditure (Shetty 1999; Vickers et al. 2003). However, selection has
not yet had the opportunity to shape mechanisms to increase physical activity in
order to match the resource abundance that has arisen in recent history. Moreover,
increased weight inherently increases the metabolic costs of physical activity,
making it less likely for those who are overweight. In the absence of compensatory
increases in energetic demands or adaptive benefits, this can trigger an obesogenic
spiral, where gaining weight makes activity less and less likely.
The lack of evolved mechanisms to balance energy intake with expenditure,
together with a system that was shaped to readily store surplus energy as fat and
lose muscle mass quickly when not used, encourages the development of
pathologies such as obesity, type 2 diabetes, cardiovascular disease, metabolic
syndrome, and other chronic and often deadly disorders. The silver lining is that the
system that evolved was shaped to flexibly respond to the costs and benefits of
physical activity that varies across individuals, environments (including social and
cultural factors), and throughout the life span. The costs and benefits can be
influenced by malleable environmental, sociocultural, psychological, and physio-
logical factors to increase or decrease the likelihood of physical activity.
This perspective builds on evolutionary approaches that have been used thus far
that highlight the ways humans are physically and mentally adapted for high levels
of physical activity. When interpreted simplistically, this research can suggest that
high levels of physical activity are part of human nature. A more comprehensive
evolutionary approach however—represented by the literature in human life history

82 9 Summary, Conclusions, and Applications
theory and reproductive ecology—suggests that the evolutionarily relevant benefits

and costs of engaging in physical activity and investing in physical fitness are
vitally important for the natural history of human physical activity. Efforts to
increase physical activity and/or exercise will benefit from an understanding of
human nature as an energy conserving physiological and psychological system that
makes “decisions” about the costs and benefits in terms of maximizing potential
reproductive success throughout the life span.
A life history perspective can help us to understand typical patterns of energy
allocation across the life span. It can predict circumstances that lead to deviations
from that pattern that may be adaptive. It can also predict the stages of the life course
where allocating energy to physical activity will be more likely to occur (e.g., play
among children, hunting, gathering, or competing for access to mates), as well as
stages where it is less likely to occur (e.g., during puberty, pregnancy, lactation, and
when offspring care is incompatible with physical activity). From there, approaches
to increase voluntary physical activity can be tailored according to sex, life stage,
and specific to individual condition and developmental experiences.
This book has indicated the important role of ecology in producing variation in
the benefits and costs of physical activity. Among horticulturalists with seasonal
resource availability, energy expenditure is reduced when not required for har-
vesting resources, particularly among women. Human females also demonstrate
reproductive function that is acutely responsive to increased energetic demands
(Ellison and Jasienska 2007), and female foragers show greater variation than males
in production according to the intensity and compatibility of childcare demands
(Dufour and Piperata 2008; Hurtado et al. 1992; Lancaster and Kaplan 2009).
Changes to the built environment that increase opportunities for obligatory physical
activity as transportation, etc., as well as economic incentives or reduced trade-offs
between provisioning, childcare, and physical activity are potential avenues for
public and private policy initiatives that may help increase physical activity and
overall health.
Genetic, epigenetic, nervous, and neuroendocrine systems likely work in concert
to influence physical activity within a given environment across the life span.
Though much of the optimizing “decision making” about whether or not the
benefits of physical activity outweigh the costs is subconscious, conscious psy-
chological processes are also involved and may be leveraged to increase physical
activity. The extent to which psychological factors effectively influence behavior, or
conditions that make rational, cognitive processes more or less likely to influence
behavior can all be examined in light of the ultimate function of behavior to
increase survival and reproductive success.
Thus, an evolutionary perspective provides a novel approach to increasing
physical activity on a large scale by changing environments, social norms, and
socioeconomic policies in ways that tip the cost/benefit ratio to favor greater
activity despite the mechanisms that have evolved to help humans conserve energy.
This approach may actually lead to net economic gains because of the over-
whelming and rising costs of health care, the chronic nature of diseases associated
with physical inactivity, and the mental health benefits associated with physical
9 Summary, Conclusions, and Applications 83
activity. To conclude, I will offer a list of take-home messages and suggestions for
changes in private and public policy, as well as basic science and intervention-
oriented research aimed at understanding and increasing physical activity based on
this evolutionary framework.
Take-home messages to guide future research and public policy
• Energy allocation systems have been shaped through natural selection to
increase predicted reproductive success. Therefore, a flexible system has evolved
to selectively engage in physical activity in domains that increase reproductive
success and survival, and in other circumstances reduce physical activity.
• There is not one dose or physical activity/exercise regime that is the best,
healthiest, or least harmful used by humans ancestrally and that we can or
should replicate (Lieberman 2015). But inactivity is harmful, and we are not
physiologically adapted to deal with environments that do not require energy
expenditure to acquire resources or survive.
• Resource availability through maturation is predicted to have an important and
lasting effect on energy allocation strategies that can make physical activity
more or less likely in adulthood. Understanding the level of resource scarcity
individuals experience throughout development may inform how thrifty their
energy allocation strategy is, which in turn affects how they physiologically and
psychologically respond to non-obligatory energy expenditure.
• Immune stress can have an important and lasting effect on energy allocation that
influences physical fitness and activity. Investment in the immune system is
predicted to trade-off against somatic investment in physical fitness and activity.
• Childhood is an important time for incorporating physical activity because
(1) there are not competing demands for energy to be put toward reproductive
effort; (2) developmental environments are important for shaping energy allo-
cation strategies; and (3) childhood activities may communicate the energetic
demands of the environment, influencing adult levels of physical activity.
– We should be increasing recess and physical education rather than
decreasing them.
– We should encourage children to discover activities from a diverse range of
intensities and modes that they will want to stick with throughout
adolescence.
• Once humans reach pubertal maturation, physical activity decreases are highly
likely due to simultaneous increases in competing demands of growth and
reproductive maturation. It may be appropriate to come up with innovative
approaches that incorporate social goals, improve self-worth, or have lower
expectations about the intensity of activities to expect at this critical stage of
development.
• Physical activity will be more likely to be adopted and maintained when the
adaptive benefits (i.e., benefits that increase reproductive success) outweigh the
costs. These costs and benefits vary throughout the life span, between the sexes,
and between individuals.
84 9 Summary, Conclusions, and Applications
– Approaches to increase activity should be sensitive to factors that make the

metabolic or opportunity costs of physical activity higher, for example, those
who:
• are overweight or obese;
• have more thrifty energy allocation strategies due to energetic stress
through development;
• face higher costs due to competing energetic demands specific to a given
life stage (i.e., during puberty, pregnancy, and lactation);
• face or faced during development higher demands for immune function.
• Public and private policies, as well as intervention efforts, can increase physical
activity by tailoring the built environment and benefit structure so that activity
and exercise present fewer trade-offs and create more opportunities for activity
that are compatible with mating effort, parenting, and producing resources.
– Built environments should be changed in ways that encourage or require
more obligatory activity.
– We should encourage playful physical activity (particularly in kids) and a
diverse range of activities that vary in intensity and compatibility with
competing energetic demands.
– Employers can incentivize physical activity in the workplace, decreasing the
trade-offs between work (the modern, usually sedentary form of parental
investment) and physical activity, and create more benefits that are consistent
and relevant for investing in children. Progressive businesses that adopt a
“benefit corporation” model that prioritizes the needs of society and the
environment in addition to profit are growing (Hiller 2013). This is often
paired with caring more about employees than corporations who adopt a
shareholder-driven model. Improving employees’ health and well-being
would fit well into the “benefit” model, as well as lead to savings in
healthcare costs and increased productivity that can offset the costs of such
incentive structures. Some ways progressive businesses can encourage
employee health and well-being and increase productivity are as follows:
• Employers can offer exercise facilities where hourly employees can be
paid to exercise for an hour each day, to decrease the production/physical
activity trade-off sedentary jobs involve.
• Exercising regularly could be incentivized monetarily through employer
contributions to health savings accounts, insurance premiums, childcare
flexible spending accounts, or college funds.
• Employers can incentivize or make technological advances widely
available that allow electricity for computers to be powered by cardio-
vascular exercise.
References 85
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infants, children and adults. European Journal of Clinical Nutrition, 53, S14–S33.
Vickers, M. H., Breier, B. H., McCarthy, D., & Gluckman, P. D. (2003). Sedentary behavior
during postnatal life is determined by the prenatal environment and exacerbated by postnatal
hypercaloric nutrition. American Journal of Physiology Regulatory, Integrative and
Comparative Physiology, 285, R271–R273.
Index
A Cortisol, 71, 72
Accelerometer, 30 Costs of physical activity, 12, 15, 81, 82
Accuracy in activity measurement, 27
Ache, xii, xiii D
Adaptation, physiological, 8 Developmental origins, 41
Adaptations, metabolic, 15 Developmental plasticity, 8
Adaptions, psychological, 5 Discordance, 5, 15, 31
Adaptive benefits, 11, 66, 81, 83 Domain-specific processing, 5
Adolescence, 8, 9, 24, 47, 83 Doubly labeled water (DLW), 27, 28, 30, 32
Adrenocorticotropic hormone (ACTH), 71
Agriculture, 15, 31, 55 E
Applications, 81 Ecology/Ecological, 19, 23, 33, 47, 53, 66, 81
Endurance running, 16, 20
B Energetic costs of physical activity, 12
Basal metabolic rate (BMR), 27, 30 Energetic stress, 53–56, 59
Benefits of physical activity, 12, 14, 82 Energetics, 53–56
Birth weight, 41, 43, 56 Energy allocation, 11, 16, 48
Body composition, 7, 28 Energy balance, 53–56
Body size, 22, 23, 30 Energy expenditure, 19, 23, 24, 27–32, 54, 58,
Bonobos, 7, 20, 21 69, 72, 81, 82
Brain Epigenetics, 73
evolution of, 20 Estradiol, 54, 67, 68, 70
metabolite distributions in, 21 Estrogen, 67
Brain growth, 47 Evolutionary psychology, 6
Evolutionary theory
C analysis, categories, 3
C-reactive protein (CRP), 48 phylogeny analysis category, 7
Catecholamines, 72 proximate analysis category, 4
Caxiuanã, 33 traditional versus, 4
Central nervous system, 73 ultimate analysis category, 4
Chimpanzees, 7, 19–22, 23, 54 Exercise, 82, 84
Cognitive processing, 4, 6, 16, 66, 82
Comparative analysis” “See also Phylogeny, F
19 Factorial method of physical activity
‘Constrained down regulation’ hypothesis, 55 measurement, 29
Corticotropin releasing hormone (CRH), 71 Fat storage, 21, 23

Human Behavior, Biology and Evolution, DOI 10.1007/978-3-319-30409-0
88 Index
Fecundity, 53–56 metabolic adaptations, 19

Female fecundity, 54 morphological adaptations, 19
Flexible response, 12, 42, 81 of non-human primates, 22
Foraging, 7, 13, 30 Life history theory, 47, 49, 50, 81, 82
Framework, life history, 4, 12, 15, 24, 47, 48, Lifespan, 7, 11–13, 15, 22, 24, 30, 67, 72, 74,
65, 67, 83 81–83
Framework, 13 Lifestyle, 31, 42
Longevity, 11, 48, 60
G Luteal phase (LH), 67
Generating hypotheses, 4
Gorillas, 21 M
Great apes, 7 Macaques, 21, 47
Growth Measurement, physical activity, 27
immune system and, 48 Men
HPG in, 69
H physical activity in, 58
Hadza, 28–30, 70 reproductive ecology in, 58
Health behavior, 4–6, 50 Menopause, 60, 67
Health psychology Menstrual cycle, energy expenditure during, 54
physical activity in, 4 Metabolic adaptations, 19
Heart rate monitors, 30 Metabolic costs in physical activity, 66
Heritable variation, 4 Mismatch, environmental, 5, 6, 16, 31
Hormones, 66 Morphological adaptations, 19
Horticulture, 13 Mortality, 6, 11, 48
HPA axis, 71– 73 Muscle mass, 20
HPG axis, 67–69
Human chorionic gonadotrophin (hCG), 68 N
Human development index (HDI), 32 Natural selection, 4
Human uniqueness in␣life history patterns, 22 traditional approach versus, 6
Hypothalamo-pituitary-adrenal (HPA)␣axis, Neuroendocrine, 65–67, 82
71 Neuroendocrine system, 65
Hypothalamo-pituitary-gonadal (HPG)␣axis, as potential mechanism, 66
67 Non-human primates, 22
in men, 69 Non-obligatory activity, 43, 55, 58, 65, 83
in women, 67
O
I Obligatory, 29
Immune system/immune function, 42, 47, 48 Obligatory activity, 12, 13, 31, 42, 58, 65, 84
In utero stress, 41, 43 Oligomenorrhea, 54
Inactivity, 6, 65–67, 82, 83 Optimization of energy allocation, 12, 16, 24
Industrial Revolution, 31 ovarian function, 53
Integrated lifestyle, 42 Ovarian steroids, 68
Integrative evolutionary model, 75 Ovarian suppression, 55, 56
Intervention, 83, 84 Ovarian supression, 55
K P
!Kung, 49 Paternal Provisioning Hypothesis, 59
Phylogeny analysis, 7
L Physical activity
Lactation, energy expenditure during, 56 Across populations, 30
Life history patterns among Tsimane’ population, 13
human uniqueness in, 22 benefits, 14
Index 89
costs and risks associated in, 14 Sedentary lifestyle, 20, 29, 41, 49, 50, 54, 57,
energetic costs of, 12 59, 73, 84
metabolism change and, 12 Self-efficacy and physical activity, 66
over time, 30 Sex-specific costs of physical activity, 53, 57,
Physical activity level (PAL), 27, 29, 31, 33 67
Physiological adaptations, 19 Sex differences and HPG axis, 73
Phylogenetic, 3, 7, 16, 21, 71, 75 Sexual selection, 5
Physiology/physiological, 3, 4, 8, 9, 11, 15, 19, Shuar forager-horticulturalists in Ecuador, 48
20, 21, 29, 31, 42, 51, 56, 58, 60, 65, 66, Sociosexual orientation index (SOI), 70
71, 73, 81 Sperm production, 58
Plasticity, 8 Storage, energy, 20, 21
Ponderal index, 55 Stress, energetic, 41–43, 55, 56, 59
Post-reproductive age Stress,psychosocial, 50
in men, 60 Survival, 5, 6, 13, 15, 16, 43, 66, 71, 82, 83
in women, 60
Postnatal development, 51 T
Precision in activity measurement, 27 Tamang, 55
Predictive adaptive responses (PARs), 41, 42 Tarahumara, 16
‘Preemptive ovarian suppression’ hypothesis, Testosterone (T), 59
55 Tinbergen, Nikolaas, 3
Pregnancy, 28, 53, 56–58, 68 Total energy expenditure (TEE), 27–29
Prenatal development, 51 in humans, 30
Primates, 19–21, 23 in primates, 30
Progesterone, 68, 70 Trade-offs, 42
Proximate mechanisms, 4, 65 energetic trade-offs, 47–49
Psychological variables, 65 Trade-offs in energy allocation, 11, 13
Psychology, 65 Tsimane’ population
Psychology/psychological, 3–6, 43, 58, 65, 72, physical activities among, 13
74, 76, 81–83
Psychological adaptation, 4, 5 U
Puberty, 48 Ultimate analysis, 3, 4, 6
Public and private policy, 82–84
V
R Variation in PAL, 27, 31, 33
Rational processing, 6, 82
Reaction norm, 8, 41, 42, 74 W
Reproductive effort, 11, 48, 50, 56 Women
in men, 59 HPG in, 67
Reproductive ecology, 53, 81 ovarian function, 53
Reproductive success, 3, 4, 6, 9, 11, 13, 15, 24, physical activity in, 53
31, 43, 49, 55, 58, 69, 75, 82, 83 reproductive ecology in, 53
S Y
Sedentary, 31 Yapu, 33

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SPRINGER BRIEFS IN ANTHROPOLOGY

HUMAN BEHAVIOR, BIOLOGY AND E VOLUTION

Human Behavior, Biology and Evolution

Human Physical Fitness

ISSN 2195-0806 ISSN 2195-0814 (electronic)

Library of Congress Control Number: 2016934017

© The Author(s) 2016

Printed on acid-free paper

This Springer imprint is published by Springer Nature

shape human biology to operate in an environment with an unlimited supply of

Part I Evolution and Life History Theory

2 Physical Activity and Life History Theory . . . . . . . . . . . . . . . . . . . 11

3 Phylogeny and Life History Patterns . . . . . . . . . . . . . . . . . . . . . . . 19

4 Physical Activity and Energy Expenditure in Humans . . . . . . . . . . 27

Part II Physical Activity and Energetic Trade-Offs Through

6 Energetic Trade-Offs and Physical Activity During

7 Physical Activity and Reproductive Ecology in Adults . . . . . . . . . . 53

Part III Summary of Proximate Mechanisms, an Integrated

9 Summary, Conclusions, and Applications . . . . . . . . . . . . . . . . . . . 81

Ann E. Caldwell received her Ph.D. in evolutionary psychology with distinction

Bolivian lowlands with a subsistence economy based on foraging and horticulture,

Examining physical activity from an evolutionary perspective can lead to novel

In 1963, evolutionary biologist Nikolaas Tinbergen speciﬁed four complementary

© The Author(s) 2016 3

Proximate and Ultimate

Traditional approaches to understand physical activity in health psychology or

Another category of analysis that would not be considered by traditional health or

Fig. 1.1 Net production

Finally, an evolutionary perspective includes an understanding of the develop-

We have broadly explored the advantages of using evolutionary theory to help us

Life history theory (LHT) is a branch of evolutionary theory aimed at explaining

© The Author(s) 2016 11

Energetic Costs of Physical Activity

Beneﬁts of Physical Activity

Summary and Applications

successfully. In humans, reproducing successfully involves not only mating effort,

Phylogenic or comparative analysis allows us to examine how traits or behaviors

Human Morphological, Physiological, and Metabolic

© The Author(s) 2016 19

Human Uniqueness in Life History Patterns

One way to utilize a life history approach to understanding physical activity is to

© The Author(s) 2016 27

TEE in Humans and Other Primates

Physical Activity in Humans over Time and Across

Dufour and Piperata (2008) highlighted the importance of understanding the

© The Author(s) 2016 41

hominin evolution, later environments predictably did resemble prenatal environ-

energy available to invest in immune system and growth simultaneously. Energetic

© The Author(s) 2016 47

that transitioning to young adulthood typically involves, particularly among girls.

As detailed in Chap. 3, women incur substantial energetic costs in order to

was once considered an on/off phenomenon measured by the presence or absence of

Therefore, human physiology preemptively decreases reproductive effort in antic-

Men’s reproductive physiology, in contrast to women’s, is generally less energet-

of early developomental resource availability) predicted metabolically costly

Post-reproductive Age Men and Women

An evolutionary, life history framework can provide useful interpretation of pre-

Interpreting Previous Findings: Why do Psychological

© The Author(s) 2016 65

Neuroendocrine System as Potential Mechanism

Rather than provide a summary of all potentially important neuroendocrine

Hypothalamo-Pituitary-Gonadal (HPG) Axis

The hypothalamo-pituitary-gonadal (HPG) axis controls the onset of reproductive