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Department of Podiatry

Evolution of Human Locomotion

 

Evolution of human locomotion outline

geologic timescale

why study evolution?

evolution of locomotion

primitive lifeforms

primates

bipedalism

modern foot disorders


Geologic timescale

4600 million years ago: formation of earth's crust - subsequent evolution of life

600 m.y. : (Palaeozoic) marine animals eg. trilobites, molluscs

450 m.y: (Silurian) primitive fish, corals

250 m.y: (Triassic/Jurassic): amphibians and reptiles dominant land animals, dinosaurs, conifers

150 m.y.: (Cretaceous) reptiles peak

65 m.y : (Tertiary) Mass extinction of dinosaurs, rise of modern animals, shrubs, grasses

55 m.y.: Mammals dominant, carnivores, horses, whales

25 m.y: more and more moden mammals: dogs, bears, modern birds - Ramapithecus appears

5 m.y: Australopithecines appear

2 m.y: Ice ages, Homo Habilis appears

1.0 m.y: Homo Erectus appears

0.5 m.y: Homo Sapiens (archaic)

0 m.y: Homo Sapiens

tallest building analogy: "If you can imagine a 10¢ piece balanced on top of the Rialto building (or the Grollo supertower for that matter), the thickness of the coin is equivalent to the time that homo sapiens has walked the earth. The rest of the height of the building is equivalent to the total period of time that life has existed on the earth"


Why study evolution ?

bipedal gait is unique to man

human locomotion is a relatively recent phenomenon


Why study evolution?

Recapitulation (Haekel's Law)

'during embryological development, and organism passes through stages which resemble the structural form of several ancestral types of the species as it evolved'

Atavism

the appearance in an organism of features similar to that of a distant evolutionary ancestor


Why study evolution ?

'Man's foot is all his own. It is unlike any other foot. It is a human specialisation, and whether he be proud of it or not, it is his hallmark. So long as man has been man, and so long as he remains man, it is by his feet that he will be known from all other members of the animal kingdom' (Buettner-Janusch, 1966)

'A major part of the podiatrist's contribution to foot health is to determine the most efficient structural posture of the supportive bony elements of the foot...a knowledge of the development of upright bipedal locomotion contributes greatly to such an understanding' (Tax, 1985)


Evolution defined:

'Evolution is a series of partial or complete and irreversible transformations of the genetic composition of populations, based principally upon altered interactions with their environment' (Dobzhansky et al, 1977).

Example: changes in ratio of black to white moths following change in environment


Two models of evolution

Gradualistic model: no such thing as a species; all organisms are a 'snap shot' in time of a gradually changing group of organisms problem is 'missing links' in the fossil record

Punctuated model: production of new species as a result of short 'bursts' of evolutionary change each new species is an addition to the previous one, not a replacement explains 'missing links', as chances of an organism being fossilized from a short 'burst' of evolution are remote.


Primitive lifeforms

aquatic environment

amoeba, jellyfish, amphioxus,

paired fins

parallel to the long axis of the body

femur,tibia, fibula and digits

 

Amphibians

five toes (pentadactylous)

limbs perpendicular to body

mechanism to push body forward

poor weightbearing and locomotor structures

Reptiles

terrestrial environment

limbs underwent rotation to be fully weightbearing structures

'oars of a boat' analogy

less tarsal bones

 

Mammals

Appeared 265 million years ago

digitigrade: bear weight on ends of metatarsals (eg. cat or dog) - B

unguligrade: walk on tips of toes (eg. horse) - C

plantigrade: foot flat on the ground (human) - A


Primate Evolution

includes monkeys, apes and man

derived from a common ancestor

appeared 70 million years ago


Primate development

Dryopithecus

tree ape that lived 12-20m years ago

major 'missing link' in fossil record

dryopithecus ' ramapithecus ' man

dryopithecus ' d. africanus ' apes

brachiators capable of upright posture but not locomotion

opposable hallux necessary to wrap around a tree limb: still evident in man's thumb

 

Ramapithecus to modern man

appeared 12-6 million years ago

first ancestor to leave the trees and live in a terrestrial environment

most important evolutionary changes occurred from this point

through Australapithecus ('Lucy'), Homo Habilis, Homo erectus, Homo sapiens

 

Australapithecus afarensis

appeared 3.5m years ago

the first habitual biped

40% skeleton nicknamed 'Lucy'

hips and knees built for upright walking

footprints in Tanzania indistinguishable from modern man (slowly walking)

 

Homo habilis

descended from Ramapithecus

fossil foot bones found in the Olduvai Gorge in East Africa

ankle joint articular surface extended anteriorly to enable greater dorsiflexion

realignment of 1st tarso-metatarsal joint

persistent bipedal walker

 

Homo erectus

desceded from H. Habilis

appeared 1-2m years ago

completely upright walking man

fossils found in Java in 1891 then South Africa, Algeria, China

responsible for the spread of man from Africa to Europe and Asia

 

Neanderthal man

'spin-off' from the evolutionary chain of man

appeared 100-40,000 years ago (?)

became extinct by unknown mechanism

foot bones found are very similar to modern man, but more robust


Bipedalism

habitual bipedalism is unique to man (as is cerebral cortex development & vocal cord structure)

energy efficiency, food gathering, free the hands, predator avoidance

longer stride length

rigidity to act as a lever for propulsion

improved balance and shock absorption

muscular efficiency


Bipedalism

Bipedalism does not require:

grasping ability of hallux

fine dexterity of digits

bulky intrinsic musculature

Hand vs. foot function

 


Bipedal adaptations

vertical pelvis and spine

increased femoral inclination (towards midline of body)

structurally longer legs

full knee extension

increased ankle dorsiflexion

loss of hallux opposability

decreased muscle bulk(esp. intrinsic)

formation of medial longitudinal arch


Recapitulation

'during embryological development, an organism passes through stages which resemble the structural form of several ancestral types of the species as it evolved' (Haekel's Law)

evident in the early stages of development of the human embryo and foetus

possible explanation for the existence of structural anomalies in the lower limb


4 Weeks foetal development

 embryo is 5mm long

tail present

limb buds arise

limbs parallel to the trund

legs externally rotated

foot plate is inverted

all digits webbed - syndactyly

recapitulation: aquatic stage

 


6 weeks

 embryo is 12mm long

tail begins to disappear

webbing regresses to form digits

limbs perpendicular to body, foot in equinus and inverted

recapitulation: amphibian stage

 


8 weeks

 23mm long

digits are distinct

limbs undergo rotation and torsion

limbs internally rotated

elbows and knees drawn to each other

recapitulation - reptilian stage

 


9 weeks

metatarsals adducted

large IMA (opposable hallux)

tibia and fibula form ankle mortise

foot inverted, parallel to sagittal plane

subtalar joint forms

recapitulation: reptilian stage


12 weeks

56mm

foot begins to dorsiflex

nails begin to appear

limbs begin to fold into thigh/leg and arm/forearm components

recapitulation: mammalian


16 weeks - birth

feet continue to evert

bones begin to ossify, starting with the calcaneus

at birth:

genu varum

low medial longitudinal arch

ligamentous laxity

external limb position

recapitulation: primate


16 weeks - birth

NB: changes which occur from birth to beginning walking are very similar to primate evolutionary changes


Atavism

appearance in an individual of traits or characteristics more like those of an earlier ancestor

commonly referred to as 'phylogenetic throwbacks'

Examples:

metatarsus primus adductus

genu varum ('bow legs')

syndactyly / polydactyly

pes planus

ligamentous laxity


Summary

bipedalism is unique to man and relatively recent

significant structural changes in the lower limb

evolutionary development of locomotion has a number of parallels with embryological development (recapitulation)




References

Tax HR. Podopaediatrics (2nd ed.) Williams & Wilkins, Baltimore 1985 (Bundoora 617.585 TAX)

Dobzhansky T, Ayala F, Stebbins G et al Evolution WH Freeman and Company, San Francisco 1977

Leakey R. Looking ahead to the past (Video) Carolina Biological Supply Company, North Carolina 1988 (Bundoora AV:573.3)

Moorhead J and Wobeskya L 'Evolutionary aspects of foot disorders' Journal of the American Podiatric Medical Association 85(4):209, 1995


Content Approved by: Head of Podiatry
Page maintained by: Podiatry Webmaster
Last Updated: August 20th, 2002