Human evolution in running

Anthropology: A Look at the Evolution of Running Culture and Exercise Physiology

Why do humans run?

Running is a simple action and exercise that most if not all humans and athletes have engaged in. But what is the purpose of running? This article attempts to give the reader an overview of running evolution, history, and culture from the beginning to modern times. This article will also give the reader the fundamental understanding of exercise physiology behind running and how one can improve endurance. Furthermore I would like to attempt to elucidate the natural innate purpose behind running as this is where the heart of anthropology of running lies. Why do humans run?

Running during prehistoric times

Evolution of humans looking at skulls
Credit: Shutterstock

About 4 million years ago our human ancestors supposedly left the trees and evolved from the Australopithecus and began walking on two legs. They then evolved to Homo habilis and Homo Erectus who then started jogging and running to eventually become long-distance runners in order to track down and hunt prey. A 2004 study Endurance running and the evolution of Homo by Daniel E. Lieberman (a professor of human evolutionary biology at Harvard) and Dennis M. Bramble (a biologist at the University of Utah) identified some of the advantageous traits including:

  • Better tendons for better recoil and elasticity
  • The arch of the foot for absorbing energy and returning it like a spring
  • Longer stride length for increased speed
  • Larger buttocks for stabilizing the trunk during running
  • Better shoulder, arm, and hip rotation allowing for better counter-balancing movements while running
  • More sweat for dissipating heat through evaporative cooling
  • Less body hair for increased convection rate and dissipating heat
Abstract of article "Endurance running and the evolution of homo"
Credit: Bramble, D., Lieberman, D. Endurance running and the evolution of Homo. Nature 432, 345–352 (2004).

Lieberman and Bramble conclude that “It is reasonable to hypothesize that Homo evolved to travel long distances by both walking and running.”
Furthermore, a 2008 study by Karen L. Steudel-Numbers (a zoologist at the University of Wisconsin, Madison) and Cara M. Wall-Scheffler,
(a biologist at Seattle Pacific University) attempted to estimate the speed of locomotion of our distance-running ancestors and concluded that Homo was restricted to long periods of walking combined with surges of slow running.

Evolution of locomotion in Homo

Running in ancient history

Panathenaic amphora (480-470 BC) with four runners at the turning point. The position of the arms, barely moving and with clenched fists, is typical of a long-distance run.

Running however became a competitive sport around 600 to 700’s BCE in the first ever recorded Olympic Games took place in Greece in 776 BCE.

Running in the first Olympic Games

The ancient stadion of Nemea
The ancient stadion of Nemea. Credit: Michael F. Mehnert

Initially the ancient Olympic Games were a one-day event until 684 BC, and then they extended it to a three-day event. It was then extended to 5 days, in 5th century B.C. The ancient Olympic Games included many competitions including running, long jump, shot put, javelin, boxing, pankration and equestrian events.

The participants ran the events naked and barefoot on the ‘stadion’ (now called a running track). The Stadion consisted of sand and the start and finish lines were simply straight lines etched in the sand. Later more official lines were made out of stone slabs. In some stadia, a starting mechanism was made to hold back athletes with a small bar that would fall at the start signal. 

An excerpt from ancient texts, Philostratus, Gymnasticus 32-33 describes the ideal body of a runner.

“To be an excellent dolichos-runner one should have shoulders and a neck about as strong as those of a pentathlete, but he should have slender and light legs as a stadion-runner. These bring their legs into a sprint with their hands, as if their hands give them wings. Dolichos-runners do this only in the final sprint, the rest of the time it is almost as if they stride, with their hands in a fist, for which they need strong shoulders.

As stadion-runners – this is the lightest event – proportionate people are very well suited, but even better are tall people, not the very tall, but those who are just a little taller than the proportionate ones, for extraordinarily tall people lack stability, like overgrown plants. They should be build firmly, because the start of a goed sprint is a good posture. The proportions of their body should be as follows: the legs should be in equipoise with the shoulders, the chest smaller than the average and with healthy organs, they should have swift knees, straight shins and hands bigger than the average. The should have proportionate muscles, for excessive muscles are like chains for speed.
The contestants in the diaulos should be build stronger than stadion-runners, but lighter than contestants in the race in armour.”

As far as running goes, for the first 56 years of the festival, a footrace was its only athletic event. The station, was a sprint of 157-196 meters and was regarded as the central event of the Olympic Games throughout its history. It wasn’t until 720 BC that long distance running was added, a race called the dolichos. The dolichos was a race that was completed over the same track as the stadion by running from end to end until a distance of up to 5,400 meters was reached. During that time, the long distance race was not considered a main event by any means, and served more as a secondary event while spectators trickled in before the more exciting competitive events. In 450 BC, the festival was expanded to included the hoplitodromos, a race in which the athletes would carry various pieces of hoplite armor weighing up to 50 pounds total. 

The full list of Greek running competitions were as follows:

Event Description
Stadion a sprint the length of the stadion track, around 200 meters
Diaulos a two stadia sprint, around 400 meters
Hippios a four stadia race, around 800 meters
Dolichos an endurance race of 18-24 laps on the stadion – about 3 miles
Hoplitodromos an encumbered race in which athletes had to wear pieces of Hoplite armor
Pentathlon a fivefold event consisting of the discus toss, javelin throw, long jump, stadion sprint, and wrestling.

On the topic of Greek footraces, likely the most prominent example to come to mind is the Marathon. Its romanticized origin story relates that in 490 BC, a courier named Pheidippides ran a total of 153 miles (246 km) in a day and a half, first to request help from the Spartans when the Persians attacked Marathon, and then to announce their subsequent victory in Athens. The iconic 26-mile (42 km) distance from Marathon to Athens was the last leg of his journey, after which, legend relates, he collapsed and died of exhaustion. Ironically, the Marathon was never an event in the ancient Olympic Games, and has only held an official place since the new games were reignited in 1896. To commemorate Pheidippides’ entire 153 mile run, an annual race called the Spartathlon, which takes place over the same historical route, has been held in Greece since 1983.

Ancient Egyptian Footraces

Pharoah Djoser Running during the heb sed run
Credit: This file is licensed under the Creative Commons Attribution-Share Alike 3.0 Germany license. Flag of Germany.svg
Attribution: Nephiliskos at German Wikipedia

The Ancient Greeks weren’t the only one’s who participated in competitive running. Ancient Egypt hosted a small number of foot races, although more ceremonial than competitive. One ancient ceremonial festival run called the heb sed, took place as early as 3000 BC.  It was a celebration of the current pharaoh and a testament of the pharaoh’s physical fitness to rule. The heb sed took place 30 years into the pharaoh’s reign and every 3 years afterward. 

The history of ancient Egypt does host one great competitive race without name. During the 7th century BC, Pharaoh Taharqa organized a race among several military units from Memphis (the Egyptian capital) to the Faiyum Oasis and back, with a 2 hour break halfway through. The distance totaled approximately 62 miles (100 km), nearly two and a half times the distance of the modern marathon.

Running in modern times

Jamaica's Usain Bolt (R) crossing the finish line to win the men's 100m final at the National stadium as part of the 2008 Beijing Olympic Games.  Read more at:
Credit: AFP File Photo

Usain Bolt

Who is the fastest runner of all time? That title belongs to Usain Bolt. Looking at the photo above you can see how far ahead Usain Bolt is compared to his counterparts who are also elite Olympic runners. 

Who is the most famous runner in the world?

There have been many famous elite runners in recorded human history. Here is a list of some of the most famous runners in history

  • Jesse Ownes
  • Roger Bannister
  • Abebe Bikila
  • Lee Evans
  • Kathrine Switzer
  • Steve Prefontaine
  • Sebastian Coe
  • Florence-Griffith-Joyner
  • Linford Christie
  • Carl Lewis
  • Dean Karnazes
  • Michael Johnson
  • Paula Radcliffe
  • Scott Jurek
  • Haile Gebrselassie
  • Hicham El Guerrouj
  • Tyson Gay
  • Asafa Powell 
  • Mo Farah
  • Allyson Felix
  • Usain Bolt

State of Running 2019

The State of Running 2019 is a Global mapping and analysis of running participation that covers 107.9 million race results from more than 70 thousand events from 1986 to 2018. It  is by far the largest study of race results in history.

Key conclusions

  1. Event participation has declined by 13% since 2016, when it peaked with 9.1 million runners crossing the finish line. Growth continues in Asia.
  2. Runners have never been slower – male runners in particular. In 1986 the average finish time was 3:52:35, whereas today it’s 4:32:49 – a slowdown of 40 minutes and 14 seconds.
  3. Runners have never been older aging from 35.2 in 1986 to 39.3 in 2018.
  4. Spain has the fastest recreational runners on the marathon distance, Russia on the half marathon, Switzerland on the 10K and Ukraine on the 5K.  
  5. For the first time in history, there are more female than male runners. In 2018, 50.24% of runners were female.
  6. Traveling to race has never been more popular than today.
  7. The motives for participating in running are potentially changing from being acheivement-focused to being phycological, health and socially focused, which in part can be proved by more people traveling to race, slower finish times and how milestone-ages (30, 40, 50) now are much less dominant than 15 and 30 years ago. 
Graph of total participants overtime.
Graph of total participants overtime. Credit:

Running is huge and has a big following, but as we can see from the chart below it’s declining in the last two years. This is mainly due to a decline in participation in Europe and the US. Participation in Asia is very much on the rise, just not enough to compensate. 

The participation in running races has peaked in 2016 with a total of 9.1 million results and then it declined to 7.9 million (a decline of 13%) in 2018. If we look at the change in participation in the last 10 years there is an increase in participation of 57.8% (from 5 to 7.9 million participants).

5-kilometer races and half marathons have the highest numbers of participants (2.9 and 2.1 million participants in 2018, respectively). But also have suffered the biggest decline in participation in the last 2 years. Half marathoners have declined by 25% (from 2.9 million) in the last 2 years. and 5K participants have declined by 13% (from 3.4 million).

10Ks and Marathons have a more modest following – 1.8 and 1.1 million participants respectively in 2018. But it’s quite stable over the last few years – the participation rates have fluctuated less than 2% the last 2-3 years.


Participation trends by distance

How fast a runner are you compared to other runners? 

Compare running calculator

If you want to see how you compare to other fellow runners, a calculator on can be used. It is based on 35 million results collected in the last 20 years from more than 28,000 races. 

Exercise physiology: How to run faster and longer

Infographic that shows exercise physiology of running and VO2 max

With scientific discoveries in human physiology, we are now able to quantify why runners run faster or have higher endurance. In this section I would like to try to explain the significance of some of these physiologic numbers to help you understand the exercise physiology behind running. The basic question “How to run faster or longer?” can be answered through these fundamental physiologic principles. 

VO₂ Max

What is VO₂ max? VO₂ max is how much oxygen your body can consume and use during exercise.

How is VO₂ max measured? VO₂ max tests are usually done in a medical facility by a doctor such as a cardiologist or a specialist in exercise physiology or fitness. However there are submaximal exercise tests that are carried out by personal trainers or fitness instructors. These include but are not limited to:

VO2 Max Graph
VO2 Max Graph. Credit:

How can you increase your VO₂ max? There are typically 2 ways to improve VO₂ max: 1. Increasing the amount of blood your heart can pump and 2. Increasing the amount of oxygen your muscles can consume.

1. High intensity exercises

Exercising at high intensity can help increase your VO₂ max. Running coaches and fitness professionals recommend training at 90 to 95% of your maximum heart rate.

Your maximum heart rate can be estimated by subtracting your age from 220. 

2. Interval training

Studies have shown that interval training results in slightly improved Vo2 max than continuous aerobic exercise. Interval training is when you alternate between high intensity exercises and periods of rest.

Using a combination of continuous training and interval training also has shown to be beneficial in improving VO₂ max.

Cardiac Output

What is cardiac out? Cardiac output is the amount of blood your heart pumps in a minute. It can be calculated by multiplying the stroke volume and the heart rate. 

Stroke volume is the volume of blood your heart pumps with each heart beat. Heart rate is the number of heart beats per minute. 

Cardiac output = Stroke volume x Heart rate. 

Increased cardiac output graph when running

An average person will pump five liters of blood per minute. However, while running, cardiac output increases significantly. In order to run faster or for longer distances, a runner must improve their cardiac output. Cardiac output and VO₂ max work synergistically in enabling the body to consume oxygen at max capacity while running.

Cardiac output is increased over time due to a few things that increase stroke volume in runners. First, when you run, you increase the volume of blood filling your left ventricle, which causes it to stretch. Your body adapts by enlarging the ventricular chamber, which allows you to pump that much more blood with each heartbeat.

Second, the greater the stretch as blood fills your left ventricle, the greater the contraction strength when your heart beats. The Frank-Starling law explains this. What is the Frank-Starling law? The Frank-Starling Law states that the stroke volume of the left ventricle will increase as the left ventricular volume increases due to the myocyte stretch causing a more forceful systolic contraction.


Frank-Starling Law
Frank-Starling Relationship. Credit: reddit

In a nutshell, if there is more blood volume that goes back to the heart during ventricular diastole, which basically means when the ventricle is relaxed to allow blood to flow into the ventricle, there is a positive inotropic effect or increased strength of contractility of the heart muscle. 

Keep in mind that interval training is the workout of choice for improving stroke volume and this makes sense. Fast repetitions followed by recovery “intervals,” during which you jog or walk. The goal is to increase your heart rate during the repetition, then allow it to slow down during the recovery. While increased blood flow during the repetition is important, it’s the recovery interval that’s vital. During recovery, your heart rate drops more quickly than the corresponding drop in blood flow. This forces the ventricles to fill more fully, creating a brief increase in stroke volume. Repeated over multiple reps, this stimulus triggers an adaptation: Increased stroke volume. 

Type II Fast Twitch Muscle Fibers

What are type II fast twitch muscle fibers? Type II fast twitch muscle fibers are a type of muscle fiber that is used in high intensity exercises such as sprinting.

Fast-twitch type II muscle fibers are further divided into Type IIx and Type IIa. 

Typically, these have lower concentrations of mitochondria, myoglobin, and capillaries compared to our slow-twitch fibers, which means they are quicker to fatigue 

These larger-sized fibers are also produce a greater and quicker force, an important consideration for power activities 

Type IIX (also known as Type IIB) fibers produce the most force, but are incredibly inefficient based on their high myosin ATPase activity, low oxidative capacity, and heavy reliance on anaerobic metabolism 

Type IIA fibers, also known as intermediate muscle fibers, are a mix of type I and type IIx, with comparable tension. Able to use both aerobic and anaerobic energy systems, these fibers have a higher oxidative capacity and fatigue more slowly than type IIx 

Type I Slow Twitch Muscle Fibers

What are type I slow twitch muscle fibers? Type I slow twitch muscle fibers are muscle fibers that are used in endurance exercises such as marathons. Slow-twitch muscle fibers have high concentrations of mitochondria and myoglobin. Although they are smaller than the fast-twitch fibers, they are surrounded by more capillaries. This combination supports aerobic metabolism and fatigue resistance, particularly important for prolonged submaximal (aerobic) exercise activities. 

Chart showing muscle fiber type and characteristics

Red Blood Cells

How do red blood cells help in running? The main function of red blood cells in exercise is the transport of O2 from the lungs to the tissues and the delivery of metabolically produced CO2 to the lungs for expiration. Despite stimulated erythropoiesis (erythropoeitin is a hormone released by kidney capillaries to stimulate production of Red blood cells), exercise can decrease the red blood cells by breaking old red blood cells down, caused by mechanical rupture when red blood cells pass through capillaries in contracting muscles, and by compression of red cells e.g., in foot soles during running or in hand palms in weightlifters. Together, these adjustments cause a decrease in the average age of the population of circulating red blood cells in trained athletes. These younger red cells are characterized by improved oxygen release and deformability, both of which also improve tissue oxygen supply during exercise.

Oxygen dissociation curve

Basically runners have a relative anemia associated from two things. 1) Increased Plasma Volume of their blood causing their hematocrit to be low and 2) Intravascular hemolysis of “old” RBCs” which leads to an increase in concentration of “young” RBCs and inability to bind the oxygen as well, allowing for increased oxygen dissociation. 

Importance of Mitochondria in Running

What is the function of mitochondria in runners? If you’ve taken Biology class you know that the Mitochondria is the “powerhouse of the cell”. This is because the mitochondria is the most important player in aerobic respiration. It is the mitochondria that makes use of the oxygen to create energy for the body to use during exercise or running. So it would make sense then that if you increased the number of Mitochondria or enhanced it’s function, you’d end up with much more energy than the average person. This is exactly what happens with trained runners. 

  • Replication: Mitochondria can split to form more mitochondria.
  • Size increase: Individual mitochondria can get bigger and rev up their functionality

The key is mitochondrial genesis. Exercise helps promote mitochondrial genesis. In a 2011 paper on mitochondrial biogenesis Regulation of mitochondrial biogenesis and GLUT4 expression by exercise, Dr. John O. Holloszy concludes that exercise promotes mitochondrial biogenesis through GLUT4 expression due to exercise. 

Article for mitochondrial biogenesis

Types of Training

Base run

These short to moderate-length runs will make up the bulk of your weekly training mileage. They should be done at your natural pace and are not meant to be overly challenging. You can make big improvements in your endurance, aerobic capacity and running economy with base runs simply because you do them so often.

Example : 5 miles at your natural pace.

Progression run

This run is designed to begin at your natural pace and end at a faster pace, closer to your ideal speed during a race. 

Example : 5 miles at your natural pace, 1 mile at marathon pace.

Interval training

These are workouts that contain short or long bursts of intense effort separated by equal or slightly longer segments of slower running, jogging or walking. The intense segments should have you pushing yourself to a point where you are gasping for air and counting the seconds until you can stop. These running workouts help increase speed and boost running economy, efficiency and fatigue resistance.

Example : 1 mile of jogging to warm up, followed by 5 sets of 1000 meter runs at 5k pace with light jogging between intervals, followed by 1 mile of jogging to cool down.


These are base runs that mix in a few intervals of varying distance and duration. They are less structured than interval runs and give you a way to play with speed without the stress. For example, you may want to challenge yourself to run faster just until you get to a certain tree, sign or street. There are no rules with these workouts. It is more freestyle running and meant to be fun. 

Example : 5 miles at your natural pace with 10 spurts of increased speed ranging between 30 seconds and 1 minute each, followed by 1 minute recoveries.

Tempo run

The tempo run is done at the fastest pace you can sustain for a certain period of time. These workouts help you both increase sustained speed and the amount of time you can sustain that pace. These runs are challenging.

Example : 1 mile of jogging to warm up, followed by 3 miles at the fastest pace that can be sustained, followed by 1 mile of jogging to cool down.

Hill repeats

As the name implies, these are short segments of uphill running that you’ll repeat to increase aerobic power, high-intensity fatigue resistance, pain tolerance and strength. The best type of hill to run these on has a moderate gradient of about 4 – 6 percent.

Example : 2 miles of jogging to warm up, followed by 10 sets of 45 second hill runs with 1-2 minutes of recovery between sets, followed by 2 miles of jogging to cool down.

Long run

When you set out on a long run, your main goal is to increase the distance over what you can comfortably do on your base runs. By increasing your raw endurance, you won’t feel limited going the distance during a race. You don’t need to run faster than your normal pace during long runs, but if you want to challenge yourself, you can increase your pace or mix in some intervals.

Example : 10 miles at your natural pace.


Incorporating sprinting into your training will improve your muscular strength and power as well as your sprint finish. 

An example sprint training session:

  • 10-minute warmup jog & dynamic stretching
  • 4x100m
  • 3 minute rest between each interval
  • 10 minute cool down jog & static stretching

Recovery Run

Also called easy runs, these are short runs done at a relatively easy pace. They are best done after a hard workout, such as interval or tempo runs, so you can still add some mileage to your training routine without pushing your body too much. Feel free to do these as slow as you need to in order to overcome any lingering fatigue or soreness.

Example : 3 miles at an easy pace on the day after intervals, hill repeats or tempo runs.


Different types of running


Significance in Anthropology

The evolution of running and the purpose of running has gone from 1. Survival to 2. Competition in ancient times to 3. Psychosocial wellbeing, fitness, and sports in modern times. It is quite interesting to see what will happen as technology makes human movement and motion less essential. As the essential need for running in the hunter-gatherer sense decreases, the want for it seems to be increasing for its innate psychological and physiologic benefits. Running will probably never disappear from sports and will likely remain for the ‘long distance’ future. However, for what purpose we will run, as we evolve, is still unclear.

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