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WHY YOUR
E VOLUTION

BRAIN
NEEDS
EXERCISE
Key transitions in the evolutionary history
of humans may have linked body and mind
in ways that we can exploit to slow brain aging
By David A. Raichlen and Gene E. Alexander
Illustration by Bryan Christie Design

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I
David A. Raichlen is a professor of biological sciences and
director of the evolutionary biology of exercise laboratory
at the University of Southern California. His research focuses
on the biomechanics and physiology of exercise from
an evolutionary perspective.

Gene E. Alexander is a professor of psychology and

psychiatry and director of the brain imaging, behavior
and aging laboratory at the University of Arizona.
He studies the aging brain in both healthy adults and
those suffering from neurodegenerative disease.

n the 1990s researchers announced a series of discoveries that would upend

a bedrock tenet of neuroscience. For decades the mature brain was under­
stood to be incapable of growing new neurons. Once an individual reached
adulthood, the thinking went, the brain began losing neurons rather than
gaining them. But evidence was building that the adult brain could, in fact,
generate new neurons. In one particularly striking experiment with mice, sci­
entists found that simply running on a wheel led to the birth of new neurons
in the hippocampus, a brain structure that is associated with memory. Since then, other studies
have established that exercise also has positive effects on the brains of humans, especially as we
age, and that it may even help reduce the risk of Alzheimer’s disease and other neurodegenera­
tive conditions. But the research raised a key question: Why does exercise affect the brain at all? 

Physical activity improves the function of many F LEXING THE BRAIN

organ systems in the body, but the effects are usually To explore why e xercise benefits the brain, we need to
linked to better athletic performance. For example, first consider which aspects of brain structure and
when you walk or run, your muscles demand more cognition seem most responsive to it. When re­­search­
IN BRIEF oxygen, and over time your cardiovascular system ers at the Salk Institute for Biological Studies in La
It is by now w  ell responds by increasing the size of the heart and build­ Jolla, Calif., led by Fred Gage and Henriette Van Praag,
established that ing new blood vessels. The cardiovascular changes are showed in the 1990s that running increased the birth
exercise has positive primarily a response to the physical challenges of of new hippocampal neurons in mice, they noted that
effects on the brain, exercise, which can enhance endurance. But what this process appeared to be tied to the production of
especially as we age. challenge elicits a response from the brain? a protein called brain-derived neurotrophic factor
Less clear has been Answering this question requires that we rethink (BDNF). BDNF is produced throughout the body and
why physical activi- our views of exercise. People often consider walking in the brain, and it promotes both the growth and the
ty affects the brain
and running to be activities that the body is able to survival of nascent neurons. The Salk group and oth­
in the first place.
Key events in the perform on autopilot. But research carried out over ers went on to demonstrate that exercise-induced
evolutionary h istory the past decade by us and others would indicate that neurogenesis is associated with improved perfor­
of humans may have this folk wisdom is wrong. Instead exercise seems to mance on memory-related tasks in rodents. The re­­
forged the link be­­ be as much a  cognitive activity as a physical one. In sults of these studies were striking because atrophy of
tween exercise and fact, this link between physical activity and brain the hippocampus is widely linked to memory difficul­
brain function. health may trace back millions of years to the origin of ties during healthy human aging and occurs to a
Cognitively chal- hallmark traits of humankind. If we can better under­ greater extent in individuals with neurodegenerative
lenging e xercise
stand why and how exercise engages the brain, per­ diseases such as Alzheimer’s. The findings in rodents
may benefit the
brain more than haps we can leverage the relevant physiological path­ provided an initial glimpse of how exercise could
physical activity that ways to design novel exercise routines that will boost counter this decline.
makes fewer cogni- people’s cog­nition as they age—work that we have Following up on this work in animals, researchers
tive demands. begun to undertake. carried out a series of investigations that determined

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that in humans, just like in rodents, aerobic exercise leads to
the production of BDNF and augments the structure—that is,
the size and connectivity—of key areas of the brain, including New Neurons
the hippocampus. In a randomized trial conducted at the Uni­
versity of Illinois at Urbana-Champaign by Kirk Erickson and
in Aging Brains
Arthur Kramer, 12 months of aerobic exercise led to an increase Exercise leads to beneficial changes in the adult brain,
in BDNF levels, an increase in the size of the hippocampus and including the birth of new neurons and increased connections
improvements in memory in older adults. among existing neurons. One of the ways in which physical
Other investigators have found associations between exer­ activity seems to induce this neuroplasticity is by increasing
cise and the hippocampus in a variety of observational studies. production of a protein called brain-derived neurotrophic
In our own study of more than 7,000 middle-aged to older factor (BDNF), which promotes neuron growth and survival.
adults in the U.K., published in 2019 in Brain Imaging and Recent research suggests that cognitively engaging the brain
Behavior, we demonstrated that people who spent more time during physical activity enhances this process.
engaged in moderate to vigorous physical activity had larger
hippocampal volumes. Although it is not yet possible to say
whether these effects in humans are related to neurogenesis or
other forms of brain plasticity, such as increasing connections
among existing neurons, together the results clearly indicate
that exercise can benefit the brain’s hippocampus and its cogni­ Cognitively engaged
tive functions. exercise may aug­
ment neuroplasticity
Researchers have also documented clear links be­­tween aero­ by exploiting
bic exercise and benefits to other parts of the brain, including physi­o­logical
expansion of the prefrontal cortex, which sits just behind the pathways between
Hippocampus
forehead. Such augmentation of this region has been tied to the body and the
sharper executive cognitive functions, which involve aspects of brain that were
forged in our hunter-
planning, decision-making and multitasking—abilities that,
gatherer forebears,
like memory, tend to decline with healthy aging and are further who had to multitask
Stem cell
degraded in the presence of Alz­heimer’s. Scientists suspect that to find food.
increased connections be­­tween existing neurons, rather than
Stem cells
the birth of new neurons, are responsible for the beneficial
give rise to
effects of exercise on the prefrontal cortex and other brain cells destined
regions outside the hippocampus. to become
new neurons Spatial
U PRIGHT AND ACTIVE memory and
navigation
With mounting evidence t hat aerobic exercise can boost brain Nascent
health, especially in older adults, the next step was to figure out neuron
exactly what cognitive challenges physical activity poses that Motor Executive
trigger this adaptive response. We began to think that examin­ system and function
control (decisions and
ing the evolutionary relation between the brain and the body
planning)
might be a good place to start. Hominins (the group that
includes modern humans and our close extinct relatives) split
from the lineage leading to our closest living relatives, chim­ New cell BDNF
panzees and bonobos, between six million and seven million migrates and
years ago. In that time, hominins evolved a number of anatomi­ develops into
an immature Sensory and
cal and behavioral adaptations that distinguish us from other new neuron
primates. We think two of these evolutionary changes in partic­ attention
systems
ular bound exercise to brain function in ways that people can
make use of today.
First, our ancestors shifted from walking on all fours to
walking upright on just their hind legs. This bipedal posture
means that there are times when our bodies are precariously
balanced over one foot rather than two or more limbs like in New
neuron is
other apes. To accomplish this task, our brains must coordinate active and
a great deal of information and, in the process, make adjust­ wired into
ments to muscle activity throughout the body to maintain our a learning
balance. While coordinating these actions, we must also watch network
out for any environmental obstacles. In other words, simply
because we are bipedal, our brains may be more cognitively
challenged than those of our quadrupedal ancestors.

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 Bipedalism
By around six million to seven million years ago
human ancestors had abandoned walking on all
fours for striding upright on their hind limbs like us.
The shift from quadrupedal to bipedal locomotion
Up and at ’Em introduced balance challenges that may have placed
new demands on the brain.
In the six million to seven million years since the human lineage diverged from
that of the chimpanzees and bonobos, our kind has evolved a host of character- Hunting and Gathering
istics that set us apart from other apes. The beneficial effects of cognitively Some two million years ago our ancestors
engaged exercise on the brain may stem from two evolutionary changes in began to forage in a new way, hunting animals
particular that made humans more physically active than our ape cousins and and gathering plant foods. This strategy
supercharged our ability to multitask: the shift to upright walking and the adop- involves far more aerobic activity than is seen
in other apes, which subsist mainly on plants.
tion of hunting and gathering as a subsistence strategy.
And it requires that the brain carry out an
array of cognitive tasks while on the move.

Homo erectus Homo sapiens

Orangutans Gorillas Bonobos Chimpanzees

Common ancestor

Second, the hominin way of life changed to incorporate ed by the hippocampus and the prefrontal cortex, among other
higher levels of aerobic activity. Fossil evidence indicates that in regions. Hunter-gatherers also often forage in groups, in which
the early stages of human evolution, our ancestors were proba­ case they may have conversations while their brains are main­
bly relatively sedentary bipedal apes who ate mainly plants. By taining their balance and keeping them spatially located in
some two million years ago, however, as habitats dried out their environment. All of this multitasking is controlled, in part,
under a cooling climate, at least one group of ancestral humans by the prefrontal cortex, which also tends to diminish with age.
began to forage in a new way, hunting animals and gathering Although any foraging animal must navigate and figure out
plant foods. Hunting and gathering dominated human subsis­ where to find food, hunter-gatherers have to perform these
tence strategies for nearly two million years until the advent of functions during fast-paced treks that can extend over more
farming and herding around 10,000 years ago. With Herman than 20 kilometers. At high speeds, multitasking becomes even
Pontzer of Duke University and Brian Wood of the University of more difficult and requires faster information processing. From
California, Los Angeles, we have shown that because of the long an evolutionary perspective, it would make sense to have a
distances traversed in search of food, hunting and gathering brain ready to respond to an array of challenges during and af­
involves much more aerobic activity than seen in other apes. ter foraging to maximize the chances of success in finding food.
Increased demands on the brain accompanied this shift to­ But the physiological resources required to build and maintain
ward a more physically active routine. When out foraging afar, such a brain—including those that support the birth and sur­
hunter-gatherers must survey their surroundings to make sure vival of new neurons—cost the body energy, meaning that if
they know where they are. This kind of spatial navigation relies we do not regularly make use of this system, we are likely to lose
on the hippocampus, the same brain region that benefits from these benefits.
exercise and that tends to atrophy as we get older. In addition, This evolutionary neuroscience perspective on exercise and
they have to scan the landscape for signs of food, using sensory the brain, which we detailed in an article published in 2017 in
information from their visual and auditory systems. They must Trends in Neurosciences, has profound implications for humans
remember where they have been before and when certain kinds today. In our modern society, we do not need to engage in aero­
of food were available. The brain uses this information from bic physical activity to find food for survival. The brain atrophy
both short- and long-term memory, allowing people to make de­ and attendant cognitive declines that commonly occur during
cisions and plan their routes—cognitive tasks that are support­ aging may be partly related to our sedentary habits.

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 But simply exercising more may not realize the full potential players spatially navigate and complete attention and memory
of physical activity for keeping brain decline at bay. Indeed, our tasks while cycling at a moderate aerobic intensity level. To eval­
model suggests that even people who already get a lot of aerobic uate the potential of this approach to boost cognitive perfor­
activity may want to rethink their routines. It is possible that we mance in healthy older adults, we are comparing a group exer­
might not always exercise in ways that take full advantage of our cising while playing the game with a group exercising without
evolved mechanisms for sustaining brain performance. the game, a group playing the game without exercising, and a
Think about the ways in which many of us get our aerobic control group that only watches nature videos. The results to
ex­­ercise. Often we go to gyms and use a stationary exercise ma­­­ date are promising.
chine; the most cognitively demanding task in such a workout Many other research groups are testing combinations of
might be deciding what channel to watch on the built-in televi­ exercise and cognitive tasks. In the near future, we will proba­
sion. What is more, these machines remove some of the de­­ bly have a better idea of how best to deploy them to support and
mands of maintaining balance and adjusting speed, among enhance cognition in both healthy individuals and those expe­
many other intrinsic cognitive challenges of movement through riencing disease-related cognitive decline.
a changing environment. In addition to specially designed interventions similar to the
What if this form of exercise is shortchanging us? Our ances­ ones described here, it is possible that participation in sports
tors evolved in an unpredictable world. What if we could modi­ that require combinations of cognitive and aerobic tasks may
fy our exercise routines to include cognitive challenges like be a way to activate these brain benefits. For example, we
those faced by our hunter-gatherer forebears? If we can aug­ recently showed that collegiate cross-country runners who
ment the effects of exercise by including a cognitively demand­ train extensively on outdoor trails have increased connectivity
ing activity, then perhaps we can increase the efficacy of exer­ among brain regions associated with executive cognitive func­
cise regimens aimed at boosting cognition during aging and tions compared with healthy but more sedentary young adults.
potentially even alter the course of neurodegenerative diseases Future work will help us understand whether these benefits are
such as Alzheimer’s. also greater than those seen in runners who train in less com­
plex settings—on a treadmill, for instance.
M OVE AND THINK Much remains to be discovered. Although it is still too early
In fact, a growing body of research suggests that exercise that is to make specific prescriptions for combining exercise and cog­
cognitively stimulating may indeed benefit the brain more than nitive tasks, we can say with certainty that exercise is a key play­
exercise that does not make such cognitive demands. For exam­ er in preserving brain function as we age. The U.S. Department
ple, Gerd Kempermann and his colleagues at the Center for of Health and Human Services guidelines suggest that people
Regenerative Therapies Dresden in Germany explored this pos­ should engage in aerobic exercise for at least 150 minutes a
sibility by comparing the growth and survival of new neurons week at a moderate intensity or at least 75 minutes a week at a
in the mouse hippocampus after exercise alone or after exercise vigorous intensity (or an equivalent combination of the two).
combined with access to a cognitively enriched environment. Meeting or exceeding these exercise recommendations is good
They found an additive effect: exercise alone was good for the for the body and may improve brain health.
hippocampus, but combining physical activity with cognitive Clinical trials will tell us much more about the efficacy of
demands in a stimulating environment was even better, leading cognitively engaged exercise—what kinds of mental and physi­
to even more new neurons. Using the brain during and after cal activities are most impactful, for example, and the optimal
exercise seemed to trigger enhanced neuron survival. intensity and duration of exercise for augmenting cognition.
We and others have recently begun to extend these studies But in light of the evidence we have so far, we believe that with
from animals to humans—with encouraging results. For example, continued careful research we can target physiological path­
researchers have been exploring combining exercise and cogni­ ways linking the brain and the body and exploit our brain’s
tive challenges in individuals experiencing notable cognitive evolved adaptive capacity for exercise-induced plasticity during
decline. Cay Anderson-Hanley of Union College in Schenectady, aging. In the end, working out both the body and the brain dur­
N.Y., has tested simultaneous exercise and cognitive interven­ ing exercise may help keep the mind sharp for life. 
tions in people with mild cognitive impairment, a condition asso­
ciated with increased risk for Alzheimer’s. More work certainly
MORE TO EXPLORE
needs to be done in populations such as this one before we can
Exercise, APOE Genotype, and the Evolution of the Human Lifespan. David A.
draw any firm conclusions, but the results so far suggest that peo­ Raichlen and Gene E. Alexander in T rends in Neurosciences, Vol. 37, No. 5, pages 247–
ple who are already experiencing some cognitive decline may 255; May 2014.
benefit from exercising while playing a mentally demanding vid­ Adaptive Capacity: An Evolutionary Neuroscience Model Linking Exercise,
eo game. In studies of healthy adults, Anderson-Hanley and her Cognition, and Brain Health. D  avid A. Raichlen and Gene E. Alexander in Trends
in Neurosciences, V  ol. 40, No. 7, pages 408–421; July 2017.
colleagues have also shown that simultaneously exercising and Differential Associations of Engagement in Physical Activity and Estimated
playing a cognitive challenging video game may elicit a greater Cardiorespiratory Fitness with Brain Volume in Middle-Aged to Older Adults.
increase in circulating BDNF than exercise alone. These findings David A. Raichlen et al. in Brain Imaging and Behavior. Published online June 17, 2019.
further bolster the idea that BDNF is instrumental in bringing https://link.springer.com/article/10.1007%2Fs11682-019-00148-x
about exercise-induced brain benefits. FROM OUR ARCHIVES
In our own work, we have developed a game designed to spe­ Evolved to Exercise. H
 erman Pontzer; January 2019.
cifically challenge aspects of cognition that tend to decline with
s c i e n t if i c a m e r i c a n . c o m /m a g a zi n e /s a
age and that are probably needed during foraging. In the game,

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