Did the Human Big Toe Evolve for Walking or Running? The Evolution of Our Toes, Foot Arch and Plantar Fascia
24 September 2026
By Mark Reyneker, BTech (Podiatry), MSc (Palaeontology), Registered Podiatrist, Foot & Gait Specialist | Founder & Clinical Director, Family Podiatry Centre
Published: 24 September 2026 | Last reviewed: 17 September 2026
Did the human big toe evolve for walking or running?
The evidence suggests that walking started the transformation of the human big toe, while running may have helped refine the modern human foot.
Early hominins were walking on two legs millions of years before modern humans appeared. At 4.4 million years ago, Ardipithecus ramidus could walk bipedally while retaining a divergent, grasping big toe. By 3.66 million years ago, the Laetoli footprints show much more human-like foot function, including evidence of an aligned hallux participating in propulsion.
Running becomes particularly interesting when we consider our unusually short toes. Experimental research found little mechanical penalty from longer toes during walking, but substantially greater digital-flexor demands during running. This raises the possibility that toe shortening was favoured as our ancestors became increasingly capable endurance runners.
The modern human foot therefore probably does not represent a single adaptation. Its basic architecture was shaped by millions of years of bipedal walking, while some later features may have made walking—and especially running—more economical.
Did the Human Big Toe Evolve for Walking or Running?
The human big toe is one of the clearest anatomical differences between our feet and those of the other great apes. But why did it change? Was it primarily an adaptation for walking efficiently on two legs, or did running help shape the short, straight and powerful big toe we have today?
As both a podiatrist and a palaeoanthropologist, I find this question particularly fascinating because the answer takes us far beyond the big toe itself.
Look down at your foot.
Your big toe — the hallux — points forwards, approximately in line with your other toes. It is relatively short, robust and connected to a highly specialised mechanical system involving the first metatarsal, two sesamoid bones, powerful muscles and the plantar fascia.
Now compare that with the foot of a chimpanzee.
The difference is striking.
The chimpanzee hallux diverges away from the other toes and is capable of participating in grasping. That is extremely useful when moving through trees.
Somewhere during our evolutionary history, however, the hominin foot began abandoning much of this grasping ability and became increasingly specialised for moving the body across the ground.
The big toe moved into line.
The toes became shorter.
The foot developed increasingly human-like arches and stiffness.
And eventually our lineage acquired a foot capable not only of remarkably efficient walking, but also sustained running.
This raises an intriguing question:
Did walking create the human big toe — or did running?
The evidence suggests that the answer is not one or the other.
Walking appears to have started the transformation. Running may have helped refine it.
Did the human big toe evolve for walking or running?
The earliest major changes to the big toe were probably associated with habitual bipedal walking.
Early hominins were walking on two legs millions of years before modern humans appeared. At approximately 4.4 million years ago, Ardipithecus ramidus could move bipedally while retaining a remarkably divergent, grasp-capable big toe.
By approximately 3.66 million years ago, however, the Laetoli footprints show substantially more human-like foot function, including an aligned hallux participating in terrestrial bipedal locomotion.
This happened long before the appearance of the body proportions commonly associated with endurance running in later Homo.
Running becomes particularly interesting when we consider another characteristic of the modern human foot:
our toes are unusually short.
Experimental biomechanics suggests that longer toes impose relatively little additional mechanical demand during walking but substantially increase the work required of the toe flexor muscles during running.
This raises the possibility that once our ancestors had already become effective walkers, running exerted additional evolutionary pressure on the foot.
The modern human foot may therefore contain layers of evolutionary history.
Its basic architecture reflects millions of years of bipedal walking.
Some of its later specialisations may have made both walking and, particularly, running more economical.
Before we walked on two legs, our feet could grasp
To understand the human big toe, we need to start with the ancestral primate foot.
Primates evolved in environments where grasping was extremely useful.
Hands could grasp.
Feet could grasp.
In many living primates, the big toe diverges considerably from the other digits. Together with long, often curved toes, this allows the foot to wrap around branches and other objects.
Chimpanzees provide a useful living comparison.
Their feet are not simply inferior versions of human feet.
They are specialised for a different locomotor repertoire.
A divergent hallux is extremely useful when climbing.
But once a lineage becomes increasingly committed to terrestrial bipedalism, the mechanical requirements change.
A foot designed partly for grasping must increasingly become a structure capable of:
- accepting body weight,
- supporting balance,
- transferring forces,
- becoming sufficiently stiff during propulsion,
- and interacting efficiently with the ground during repeated steps.
This transition did not happen overnight.
And one of our earliest hominin relatives captures an extraordinary stage in the process.
Ardipithecus: walking with a grasping big toe
Ardipithecus ramidus lived approximately 4.4 million years ago in what is now Ethiopia.
Its foot is remarkable.
The hallux remained widely abducted — much more like the grasping big toe of an ape than the straight hallux of a modern human.
Yet other aspects of the skeleton indicate that Ardipithecus was capable of terrestrial bipedal locomotion.
Lovejoy and colleagues (2009) described this unusual combination as a foot capable of combining prehension and propulsion.
In other words, this was not a modern human foot.
It was not simply an ape foot either.
It represented a different evolutionary solution.
And it tells us something extremely important:
our ancestors began walking on two legs before they possessed a modern human big toe.
That immediately challenges a common misconception about human evolution.
We did not first evolve a modern foot and then start walking.
Bipedalism began first.
The foot continued changing afterwards.
Then something remarkable happened
Move forward roughly 700,000 years.
At Laetoli in Tanzania, early hominins walked across wet volcanic ash.
Soon afterwards, additional ash covered the surface.
The footprints were preserved.
Approximately 3.66 million years later, we can still examine them.
The Laetoli footprints are among the most extraordinary pieces of evidence in palaeoanthropology because bones tell us what an animal's anatomy looked like.
Footprints can tell us something about what the foot was actually doing.
They are generally associated with Australopithecus afarensis, the species that includes the famous partial skeleton known as Lucy.
And compared with Ardipithecus, something significant had changed.
Laetoli: the big toe joins the direction of travel
Crompton and colleagues (2012) analysed the Laetoli footprints using topographic measurements, experimentally generated footprints and computer simulation.
Their results suggested surprisingly human-like external foot function.
The footprint makers appear to have demonstrated characteristics consistent with:
- upright bipedal walking,
- an adducted hallux,
- a functional longitudinal arch,
- lateral-to-medial transfer of loading,
- and propulsion involving the big toe.
We should be careful not to imagine that an Australopithecus afarensis foot functioned exactly like ours.
It probably did not.
Fossil anatomy indicates that important differences remained between australopith and modern human feet.
Nevertheless, the overall evolutionary change is difficult to miss.
By approximately 3.66 million years ago, the big toe had become substantially incorporated into terrestrial bipedal locomotion.
And that date is crucial.
It is far earlier than the appearance of the classic long-legged Homo body form usually discussed in relation to endurance running.
This provides our first major clue.
Running cannot easily explain the initial alignment of the human big toe.
Walking came first.
Why would walking favour a straight big toe?
Think about what happens during a normal step.
The heel contacts the ground.
The body progresses over the foot.
Eventually the heel begins to rise while the forefoot remains in contact with the ground.
Pressure progresses forwards.
The toes dorsiflex.
A big toe positioned alongside the other digits provides a very different mechanical arrangement from a divergent grasping hallux.
Instead of functioning primarily to wrap around something, the hallux can participate in supporting and transmitting forces beneath the medial forefoot.
This becomes particularly important during late stance.
But something else happens when the toes dorsiflex.
They interact with the plantar fascia.
The big toe became connected to a mechanical windlass
In 1954, anatomist J.H. Hicks described what became known as the windlass mechanism of the foot.
The plantar fascia runs from the heel toward the forefoot and toes.
As the toes dorsiflex, the plantar fascia wraps around the metatarsal heads.
This increases tension within the structure and contributes to changes in the shape and stiffness of the foot.
The analogy is a windlass: a cable being wound around a drum.
The hallux therefore does much more than simply provide a surface from which we “push off”.
Its movement interacts mechanically with structures extending all the way back toward the heel.
Modern research has shown that the foot is considerably more complicated than a purely passive windlass. Muscles, ligaments, joint geometry and both the longitudinal and transverse arches contribute to foot stiffness.
Nevertheless, the principle remains important.
The big toe became incorporated into an increasingly integrated propulsive system.
But walking does not explain everything
If walking explains why the hallux became aligned with the other toes, another question remains.
Why did our toes become so short?
Modern humans have unusually short toes compared with many other primates.
One obvious explanation would be that once our ancestors stopped depending heavily on their feet for grasping, long toes were no longer necessary.
That is probably part of the story.
But it does not tell us whether shorter toes provided an actual locomotor advantage.
Campbell Rolian and colleagues tested precisely this question.
And the results are fascinating.
Did running help make our toes shorter?
Rolian and colleagues (2009) investigated how toe length influences the mechanical demands placed upon the digital flexor muscles during walking and running.
Imagine making the toes longer.
The point at which ground forces act beneath the toes moves farther from the metatarsophalangeal joints.
That increases the external lever arm acting upon the toes.
The muscles controlling the digits must therefore work harder to resist those forces.
The researchers wanted to know whether that additional mechanical demand mattered equally during walking and running.
It did not.
During walking, longer toes produced relatively little additional digital-flexor demand.
During running, the effect was dramatically greater.
Their modelling suggested that increasing relative toe length by only 20% could approximately double peak digital-flexor impulse and mechanical work during running.
That is an extraordinary difference.
It suggests that simply becoming a habitual walker may not fully explain why modern human toes became so short.
Running provides another possible selective pressure.
Long toes become expensive when you run
This makes biomechanical sense.
Running produces a very different mechanical environment from walking.
Ground-contact times become shorter.
Forces become larger.
The foot must absorb, store, control and return energy rapidly.
Longer toes create longer external lever arms around the metatarsophalangeal joints.
The digital flexor muscles therefore have to work harder to stabilise them.
Shortening the toes reduces this demand.
This does not prove that humans evolved short toes because our ancestors ran.
Evolutionary hypotheses rarely permit such simple conclusions.
But it does demonstrate something important:
short toes provide a much more obvious mechanical advantage during running than during ordinary walking.
That makes running an extremely plausible part of the evolutionary story.
Then Homo appears
This brings us to one of the most influential ideas in modern human evolutionary biology.
In 2004, Dennis Bramble and Daniel Lieberman proposed that endurance running played an important role in the evolution of the genus Homo.
Their argument was not based upon one anatomical characteristic.
They identified a suite of features that potentially improve running performance.
These included changes involving:
- relatively long legs,
- shorter toes,
- enlarged lower-limb joint surfaces,
- improved trunk stabilisation,
- enlarged gluteus maximus,
- the nuchal ligament,
- elastic tendons,
- and spring-like structures within the foot.
Many of these features become particularly evident with Homo, beginning roughly around 2 million years ago.
Humans are not particularly impressive sprinters compared with many quadrupedal mammals.
But we possess another unusual ability.
We can run for a surprisingly long time.
Why would our ancestors need to run?
This remains debated.
One hypothesis is persistence hunting, in which hunters repeatedly pursue an animal over long distances, particularly in hot environments, eventually exhausting it.
Another possibility involves travelling efficiently between dispersed resources.
Running may also have been useful for reaching animal carcasses before competitors or scavengers.
But we need to distinguish an attractive evolutionary narrative from what the fossil record can actually demonstrate.
We cannot watch a two-million-year-old hominin running.
Nor can we determine precisely how frequently different species ran.
What we can examine is anatomy.
And later Homo increasingly possessed anatomical characteristics that would have made endurance running more economical than it was for earlier hominins.
The exact selective pressures responsible remain open to scientific debate.
The chronology gives us an important answer
Put the dates together.
Ardipithecus ramidus:
~4.4 million years ago
Bipedal capability, but still a strongly divergent, grasping hallux.
Laetoli:
~3.66 million years ago
Human-like aspects of bipedal foot function and a substantially adducted hallux.
Later Homo:
~2 million years ago onwards
Increasing evidence of body proportions and anatomical characteristics compatible with efficient endurance running.
This sequence matters.
It tells us that the evolution of the human foot probably occurred in stages.
Stage 1 — climbing plus early bipedalism
Early hominins retained considerable arboreal capability.
The foot still needed to grasp.
Stage 2 — increasingly committed terrestrial walking
The hallux became increasingly aligned with the other toes.
The foot became better adapted to supporting and propelling the body during repeated bipedal walking.
Stage 3 — increasingly economical terrestrial locomotion
Later modifications, including shorter toes and enhanced elastic mechanisms, may have reduced the energetic and muscular costs of walking and particularly running.
Holowka and Lieberman (2018) have argued for a broadly similar staged interpretation of human foot evolution.
It makes much more biological sense than imagining that the modern human foot appeared as a complete package.
The plantar fascia adds another twist
There is another structure beneath your foot that becomes extremely important to this story.
The plantar aponeurosis, commonly called the plantar fascia.
It is tempting to assume that humans developed this strong band of connective tissue specifically when we developed an arch.
Comparative anatomy suggests otherwise.
Sichting and colleagues (2020) examined plantar aponeurosis anatomy across primates.
Their work indicates that the basic anatomical configuration of the human plantar aponeurosis has much deeper evolutionary origins.
Chimpanzees possess a broadly comparable arrangement.
So our ancestors probably did not suddenly “invent” a plantar fascia when they became terrestrial bipeds.
Instead, evolution appears to have modified and repurposed an existing structure.
This is common in evolution.
Old anatomy acquires new functions.
The plantar fascia becomes part of a spring
As the human foot evolved increasingly pronounced arch architecture, the plantar aponeurosis could contribute to something particularly useful.
Elastic energy storage.
When the longitudinal arch deforms under load, elastic structures within the foot stretch.
Energy can temporarily be stored within them.
As those tissues recoil, some of that energy can be returned.
The foot therefore behaves partly like a spring.
Ker and colleagues demonstrated this elastic behaviour of the human arch in classic experimental work in the 1980s.
But later research revealed something particularly interesting about running.
What happens if we stop the arch from working normally?
Stearne and colleagues (2016) experimentally restricted normal longitudinal-arch compression while people walked and ran.
The result was striking.
When arch compression was restricted, the energetic cost of running increased by approximately 6%.
The same significant energetic penalty was not observed during walking.
Think about what that means.
The arch certainly has important functions during walking.
But its ability to deform and recoil appears to provide a particularly valuable energy-saving mechanism during running.
Once again, we see the same evolutionary pattern.
An anatomical system already useful during walking becomes particularly advantageous when humans start running.
And the big toe is connected to this spring
This is why the evolution of the big toe cannot really be separated from the evolution of the rest of the foot.
The plantar fascia extends towards the toes.
The hallux interacts with it as the first metatarsophalangeal joint dorsiflexes.
Behind the hallux sits the first metatarsal.
Beneath that sit the sesamoids.
Behind them are the midfoot and arches.
The intrinsic foot muscles help regulate stiffness.
The Achilles tendon and calf muscles contribute enormous forces from above.
The entire system interacts.
Evolution was therefore not simply changing a toe.
It was changing a locomotor system.
Why the sesamoids matter
There is another fascinating part of this system sitting directly underneath the first metatarsal head.
Two small bones: the sesamoids.
These bones sit within the flexor hallucis brevis apparatus beneath the first metatarsophalangeal joint.
They help maintain the mechanical advantage of the muscles acting upon the hallux and participate in transmitting substantial loads beneath the medial forefoot.
Their position tells us something about the mechanical specialisation of the human first ray.
The modern hallux is not simply aligned with the other toes.
It sits at the end of a remarkably reinforced mechanical pathway.
That becomes clinically obvious when something goes wrong with the sesamoids.
Despite being tiny, sesamoid injuries can dramatically affect walking and running.
Was the big toe really the star of this evolutionary story?
Perhaps not by itself.
This is an important point.
It is easy to turn the hallux into the hero of human locomotion.
But evolution did not simply create a powerful big toe.
It created a foot in which the hallux could work effectively because everything behind it was changing as well.
The longitudinal arch changed.
The transverse arch contributed stiffness.
The toes shortened.
The plantar tissues changed.
The intrinsic muscles participated in regulating foot mechanics.
The heel, ankle and Achilles tendon interacted with the system.
And the rest of the lower limb changed too.
The modern human foot is therefore better understood as an integrated mechanical structure than as a collection of individual bones.
My own research made this particularly interesting
My MSc research at the University of the Witwatersrand investigated high-gear and low-gear propulsion in human gait and its relationship to metatarsal diaphyseal cross-sectional geometry (Reyneker, 2022).
Essentially, I was interested in how humans distribute forces across the forefoot and whether habitual loading might leave structural signatures within the metatarsals themselves.
This work reinforced something I think is important both clinically and evolutionarily.
There is no reason to assume that every human uses the forefoot in precisely the same way.
The evolution of an aligned hallux created the anatomical capacity for powerful medial forefoot propulsion.
But humans retain considerable variation in how forces travel across the foot.
Different environments, activities, anatomy and locomotor strategies may all influence forefoot loading.
The big toe is important.
But it functions as part of a five-ray forefoot.
That distinction becomes important whenever evolutionary anatomy is translated into modern clinical advice.
Evolution does not prescribe how you should run today
There is a trap in evolutionary medicine that we should avoid.
Humans evolved without modern shoes.
Therefore, some people conclude:
Humans should run barefoot.
That conclusion does not automatically follow.
Understanding how our ancestors evolved tells us why certain structures exist.
It does not necessarily tell us what every modern person should do.
A modern runner may be:
- 20 or 70 years old,
- highly trained or completely sedentary,
- accustomed to shoes or accustomed to barefoot activity,
- running on trails or concrete,
- carrying more body mass,
- recovering from injury,
- living with arthritis,
- or possessing very different foot morphology from another runner.
Evolution provides context.
It is not a universal treatment prescription.
This distinction is particularly important when discussing barefoot and minimalist footwear.
Evolution also does not create perfect anatomy
Another misconception is that because a structure evolved, it must be perfectly designed.
It isn't.
Evolution works with existing anatomy.
Structures are modified rather than redesigned from scratch.
And adaptations frequently involve compromises.
Our hallux lost much of its grasping capability.
Our toes became shorter.
Our foot became increasingly specialised for terrestrial locomotion.
Those changes provided enormous advantages.
But the same mechanically specialised structures can also develop problems.
Hallux valgus.
Hallux rigidus.
Sesamoid injuries.
Plantar fascia disorders.
Metatarsal stress injuries.
Modern pathology does not mean that evolution “failed”.
It reflects the reality that biological structures operate under competing demands, considerable individual variation and environments that can be very different from those in which they evolved.
Walking probably started the story
So let us return to the original question.
Did the human big toe evolve for walking or running?
The earliest part of the answer is relatively clear.
Walking came first.
Habitual bipedalism began millions of years before the appearance of modern humans.
At 4.4 million years ago, Ardipithecus demonstrates that early bipedal locomotion could coexist with a grasping hallux.
By 3.66 million years ago, the Laetoli footprints indicate a much more human-like relationship between the big toe and terrestrial propulsion.
The alignment of the hallux therefore appears deeply connected to the evolution of habitual bipedal walking.
But the story did not stop there.
Running may have helped finish the transformation
The modern human foot differs from those earlier hominin feet in more than hallucal alignment.
Our toes are exceptionally short.
Our arch can behave as an elastic structure.
Our plantar tissues participate in energy storage and return.
And our entire lower limb possesses characteristics that allow sustained running.
The Rolian experiments provide a particularly intriguing clue.
Longer toes appear relatively inexpensive during walking.
During running, however, they become substantially more mechanically demanding.
That makes toe shortening much easier to understand if running formed part of the selective environment affecting later hominins.
Likewise, experimental restriction of arch compression appears to make running measurably more metabolically expensive.
These findings do not prove that running created the modern human foot.
But they make it increasingly difficult to explain every aspect of our foot using walking alone.
A 4.4-million-year experiment beneath your feet
Look at your big toe again.
What you are looking at is not simply a digit.
It represents the surviving result of millions of years of evolutionary experimentation.
At approximately 4.4 million years ago, one of our early relatives could walk upright while retaining a grasping hallux.
By 3.66 million years ago, another hominin was walking across volcanic ash with a much more human-like foot.
Later, the toes shortened.
The arch became increasingly capable of storing and returning energy.
The plantar fascia became integrated into an extraordinary spring-like system.
And eventually members of our genus acquired the capacity to travel extraordinary distances across the landscape.
Some of that travelling was walking.
Some of it may have been running.
And the anatomy beneath our feet still contains evidence of both.
The answer is not walking versus running
Evolution rarely gives us the satisfaction of a simple answer.
The human big toe did not suddenly appear because our ancestors needed to run.
Nor can every feature of the modern human foot be explained solely by walking.
Instead, the evidence suggests a sequence.
Walking began the transformation.
Habitual terrestrial bipedalism progressively converted a grasping primate foot into a propulsive hominin foot.
Running may then have refined it.
Shorter toes reduced mechanical demands during running. Elastic structures within the arch improved energy storage and return. Changes elsewhere in the body made sustained running increasingly economical.
The modern human foot therefore represents layers of evolutionary history superimposed upon one another.
And perhaps that is the most interesting answer of all.
We did not evolve a walking foot or a running foot.
We inherited a grasping primate foot, transformed it into a walking foot, and then continued modifying it until it became remarkably good at running too.
Book An Appointment
Disclaimer: The word "treatment" in this article refers to the care and management of a patient’s health to prevent, cure, or improve a condition. Treatment results vary and do not necessarily indicate a cure. This article is for informational and educational purposes only and does not constitute medical advice.
About the Author
Mark B. Reyneker, BTech (Podiatry), MSc (Palaeontology) is a podiatrist and Founder & Clinical Director of Family Podiatry Centre, with more than 25 years of clinical experience across South Africa, Malaysia and Singapore. His clinical interests include foot and lower-limb pain, gait and biomechanics, sports-related foot conditions, orthotic therapy and footwear.
Alongside his clinical practice, Mark conducts research into human gait and foot biomechanics. His MSc research at the University of the Witwatersrand investigated human propulsion and the structural properties of the metatarsals. He is also the inventor of A Foot Orthotic, an orthotic technology developed through an international patent family.
References
- Bramble, D.M. and Lieberman, D.E. (2004) ‘Endurance running and the evolution of Homo’, Nature, 432, pp. 345–352.
Available at: https://doi.org/10.1038/nature03052 - Crompton, R.H., Pataky, T.C., Savage, R., D’Août, K., Bennett, M.R., Day, M.H., Bates, K., Morse, S. and Sellers, W.I. (2012) ‘Human-like external function of the foot, and fully upright gait, confirmed in the 3.66 million year old Laetoli hominin footprints by topographic statistics, experimental footprint-formation and computer simulation’, Journal of the Royal Society Interface, 9(69), pp. 707–719.
Available at: https://doi.org/10.1098/rsif. - Hicks, J.H. (1954) ‘The mechanics of the foot. II. The plantar aponeurosis and the arch’, Journal of Anatomy, 88(1), pp. 25–30.
Available at: https://pubmed.ncbi.nlm.nih.gov/13129168/ - Holowka, N.B. and Lieberman, D.E. (2018) ‘Rethinking the evolution of the human foot: insights from experimental research’, Journal of Experimental Biology, 221(17), jeb174425.
Available at: https://doi.org/10.1242/jeb.174425 - Ker, R.F., Bennett, M.B., Bibby, S.R., Kester, R.C. and Alexander, R.McN. (1987) ‘The spring in the arch of the human foot’, Nature, 325, pp. 147–149. Available at: https://doi.org/10.1038/325147a0
- Lovejoy, C.O., Latimer, B., Suwa, G., Asfaw, B. and White, T.D. (2009) ‘Combining prehension and propulsion: the foot of Ardipithecus ramidus’, Science, 326(5949), pp. 72e1–72e8.
Available at: https://pubmed.ncbi.nlm.nih.gov/19810198/ - Reyneker, M.B. (2022) An Investigation into High Gear and Low Gear Propulsion in Human Gait and its Relation to Metatarsal Diaphyseal Geometric Cross-Sectional Properties. MSc dissertation. Johannesburg: University of the Witwatersrand.
Available at: https://wiredspace.wits.ac.za/bitstreams/77c0af20-51ed-4347-b71f-6474d01d8a44/download - Rolian, C., Lieberman, D.E., Hamill, J., Scott, J.W. and Werbel, W. (2009) ‘Walking, running and the evolution of short toes in humans’, Journal of Experimental Biology, 212, pp. 713–721.
Available at: https://doi.org/10.1242/jeb.019885 - Sichting, F., Holowka, N.B., Ebrecht, F. and Lieberman, D.E. (2020) ‘Evolutionary anatomy of the plantar aponeurosis in primates, including humans’, Journal of Anatomy, 237(1), pp. 85–104.
Available at: https://doi.org/10.1111/joa.13173 - Stearne, S.M., McDonald, K.A., Alderson, J.A., North, I., Oxnard, C.E. and Rubenson, J. (2016) ‘The foot’s arch and the energetics of human locomotion’, Scientific Reports, 6, 19403.
Available at: https://doi.org/10.1038/srep19403 - Venkadesan, M. et al. (2020) ‘Stiffness of the human foot and evolution of the transverse arch’, Nature, 579, pp. 97–100.
Available at: https://doi.org/10.1038/s41586-020-2053-y
Frequently Asked Questions
Humans possess several anatomical characteristics that make sustained running possible, including relatively long legs, short toes, elastic tendons, an arched foot and effective thermoregulation. However, this does not mean every feature of the human body evolved specifically for running. Bipedal walking appeared millions of years earlier (Bramble and Lieberman, 2004).
We cannot determine an exact date from fossils. Bramble and Lieberman (2004) argued that a suite of characteristics associated with endurance running becomes particularly apparent with the genus Homo, roughly around 2 million years ago. This remains an evolutionary hypothesis rather than evidence for a precise moment when endurance running began.
Shorter toes reduce the external lever acting around the metatarsophalangeal joints. Rolian et al. (2009) found that longer toes imposed relatively little additional digital-flexor demand during walking but substantially greater demands during running, suggesting that running may have contributed to selection for shorter toes.
A divergent hallux is useful for grasping during climbing. As hominins became increasingly committed to terrestrial bipedalism, the big toe became aligned with the other digits and increasingly incorporated into weight bearing and propulsion.
The approximately 3.66-million-year-old Laetoli footprints preserve direct evidence of early hominin foot-ground interaction. Detailed analysis indicates human-like aspects of bipedal foot function, including an adducted hallux and medial forefoot propulsion (Crompton et al., 2012).
Possibly, but fossils cannot establish how often or how effectively Australopithecus ran. Evidence for habitual bipedal walking is much stronger. The more extensive anatomical suite associated with endurance running becomes particularly apparent later in Homo.
Not exclusively. The arches contribute to foot mechanics during both walking and running. However, Stearne et al. (2016) found that restricting normal arch compression increased the energetic cost of running by approximately 6%, suggesting that elastic arch behaviour is particularly valuable during running.
Probably not from scratch. Comparative anatomy indicates that the basic configuration of the plantar aponeurosis predates the modern human foot. Existing primate anatomy appears to have been modified and incorporated into the increasingly specialised hominin foot (Sichting et al., 2020).
The hallux is important in both. During walking it participates in late-stance forefoot loading and propulsion. Running increases the magnitude and rate of loading and places greater demands upon the toes and elastic structures of the foot. Its modern anatomy probably reflects multiple locomotor pressures rather than a single behaviour.
Humans evolved before modern footwear, but that does not mean every modern person should run barefoot. Evolutionary history explains anatomy; it does not automatically prescribe modern footwear or treatment. Conditioning, anatomy, surface, training history, age and pathology all matter.