Bipedalism

Eduzendium article

Bipedalism is the condition of having or using only two feet for locomotion. This form of movement is found in a few groups of animals on Earth. Throughout the course of evolutionary history, the use of bipedal movement came forward several times as an adaptation, including at what many consider to be an influential point in human evolution. The particular selective pressures that acted to bring bipedalism to the forefront time and time again are most likely diverse and unique to each group for which this type of locomotion was expressed. However, the prevalence of bipedal movement in animals both extant and extinct shows that this form of locomotion is sometimes advantageous.

The use of bipedal movement is thought to be a key element in human evolutionary history, as it is part of what separates humans and their ancestors from the other Great Apes. By virtue of similar musculoskeletal organization of human and ape bodies it is possible to trace common ancestry. Humans today bear many anatomical resemblances to their closest relatives. Residual anatomical traits that include but are not limitied to dorsally-placed scapulae and a wide range of motion in the shoulder, arm, and wrist point to an origin of bipedal locomotion that is very recent in evolutionary history of the Genus Homo. These residual traits suggest suspensory adaptations in the forelimbs of a primarily terrestrial group of animals that usually bear the bulk of its mass on their hindlimbs.

Species displaying bipedal locomotion

There are many species on Earth that use bipedal movement as their primary means of terrestrial locomotion, and also some that can use bipedal movement when pressed to escape danger.

Birds

In nearly every species of bird, a form of terrestrial bipedal movement shows itself. Because of their wings many birds do not use bipedalism as their primary means of locomotion in life, but when they land they hop or walk on two feet. Birds whose lives are spent in a mostly terrestrial capacity such as chickens, kiwi birds, and larger forms like ostriches and emus have reduced wings and larger, stronger legs. These birds are often highly-efficient bipeds, able to run at high speeds and relatively agile for their size. Ostriches have been known to run at speeds in excess of 40 mph (65 km/h)when chased by predators.

Birds are thought to be descendants of theropod dinosaurs, and are thought to have split from that lineage before the event that cause the extinction of much of land vertebrates 65 million years ago at the end of the Cretaceous period.

Dinosaurs

The type of bipedalism seen in birds is similar to that seen in bipedal dinosaurs, such as the Tyrannosaurus rex. A recent study that used chickens as analogs for dinosaur locomotion showed it was possible to change the weight-bearing bones in the legs to a more robust, non-avian dinosaur-like form through adding weights to the back of a chicken.(Reference will be added) The weight added was to symbolize a tail, which changed the form of the femur of the birds in the experiment.

Thus, dinosaur bipedal locomotion may be similar to that of birds, but the addition of the tail is a key morphological trait that changes it slightly.

Lizards

Several extant species of lizard walk or run on their hind legs when trying to escape predators. Lophognathus longirostris is an example that is closely related to the frill-necked lizards, and runs on its back legs to gain speed and agility over short distances. Physignathus lesueurii, another relative, is capable of running for short distances on the surface of water.

Anatomical Correlates of Bipedalism in Humans and Recent Human Ancestors

Through looking at the osteology of humans and their close relatives in the fossil record, a few key anatomical traits for the particular bipedalism associated with humans have come forward. A combination of these traits makes it possible for humans to walk upright and allows scientists to diagnose whether a fossil likely used bipedal movement or not.

Medially-Angled Femur

One trait that is often used to diagnose bipedal movement in hominins is the angle of the femur. In humans this bone angles toward the mid-line of the body, allowing for better balancing when walking or running and the load of the weight to be better distributed. When balanced on the distal (distant) end of the bone on the medial and lateral condyles, the angle created between the it and the flat surface it is balanced on is around 10 degrees. This bi-condylar angle is nearly 14 degrees in TM1513, an Australopithecine fossil found in 1938, as compared to the human average of 10 degrees and the near-nonexistance of such an angle in chimpanzees. In the few fossilized femora that have been unearthed this angle has been helpful to show some form of bipedalism.

In addition, the medial condyle on the distal end of the femur of a human-like biped is considerably larger than in other extant apes. This is an adaption that increases the stability of the knee joint and prevents the distal end of the femur from slipping out of alignment.

Sigmoid Spine Shape

The "S" shape of the spines of humans and their closest bipedal relatives is another characteristic that helps diagnose a biped. This shape is the caused by progressive dorsal wedging. This morphological trait is the result of lumbar lordosis and thoracic kiphosis, both of which are different curves of the spine. Lumbar lordosis is the forward curve of the lower portion of the back, and thoracic kiphosis is the slight backward curve of the vertebrae to which the ribs attach. These shapes are likely developmental in nature and not genetic, as they develop in infants after they begin to walk upright and not before.

Some recent work on the lumbar vertebrae of female humans and our closest relatives proposes that lumbar lordosis is greater in females because they need to be able to carry the weight of a fetus when they are pregnant. Women appear to have one or two more lumbar vertebrae that are capable of slipping forward a bit and creating more of a curve in the lower back. Without this adaptation, women who became pregnant might fall over constantly (to the detriment of their fetus) because they would not be able to balance very well.

Low/Broad Illium in Os Coxae

The Os Coxae are formed of three fused bones that form one half of the pelvis. In chimpanzees, these bones are elongated and somewhat gracile due to the distribution of weight in a quadruped. An animal that moves like a human biped, on the other hand, places a lot of weight on the os coxae and therefore those bones have become short, thick, and broad. The illium in particular, the bone that is at the top of the pelvis. The bones need to be compact in order for the muscles of the legs to attach strongly to hold up a person as they walk.

In studies of biomechanics, there appears to be a balance between a need for width and more efficient narrow hip for a biped's pelvis In modern human women, the pelvis is a compromise between the demands of walking, running, and jumping bipedally and the difficulty of delivering a fetus with a large brain size

Foramen Magnum

Upon finding the Taung Child in 1925, Raymond Dart faced harsh criticism in the attempt to prove that this was actually a bipedal human ancestor. As there were no post-cranial remains, the only venue open to him was to try to infer the position of its foramen magnum. In human-like bipeds, this opening that the brain stem goes through is anteriorly placed. It is directly under the top of the skull, to allow the head to be held straight up and down. Dart inferred the position of the foramen magnum on the skull and believed it was placed anteriorly and that this was therefore a bipedal creature. With the lack of post-crania for Australopithecus africanus at the time, few accepted the view.

In fact, it was not until 1947 when Broom and Robinson found a partial skeleton, STS 14, that the world began to accept that A. africanus was indisputably bipedal.

Distinctly Arched Feet

In 1978, Mary Leakey and her team were looking for fossil hominids in the Laetoli region of Tanzania when they stumbled upon a find unlike any other. Over 100 footprints of many species, preserved in volcanic ash, captured a moment in time dated 3.7-3.5 million years ago. Probably accumulated over a period of hours, the footprints give invaluable information about behavior in Paleolithic animals, including elephants, sabertooth cats, and most stunningly of all, bipedal hominids.

The footprints at Laetoli show a pair of hominids walking side by side, possibly even arm in arm by some estimations. These footprints are clearly those of a biped and show an arch in the foot. This is a structure that helps bear the weight of the body when walking. This element of the Laetoli prints (along with the non-opposable big toes discussed below) is what diagnosed some of the earliest evidence of bipedalism known to science.

The energy cost of bipedal locomotion

In humans, viewed as a system, walking and running emerge as system behaviors (no subsystem of the human organism itself walks or runs). The energy cost to the system of those locomotor behaviors defines a property of the system applicable to those behaviors. The energy cost owes to the appropriate forces the system must generate to support itself against gravity and to swing the locomoting limbs to achieve forward motion.

Researchers find that the rate at which the system produces those forces — viz., ‘force production’—provides a correlate of the system’s energy cost of locomotion. Thus, if one could develop a mathematical model that predicts force production from readily determined values of variables related to anatomy (e.g., limb length) and motion (e.g., forward speed), that model could then predict the system property of energy cost of bipedal locomotion.

Based on the findings of earlier studies, Harvard anthropologist Herman Pontzer developed a mathematical model — viz., an equation — that justified force production as a function of three variables:

• the rate of muscular force production in the vertical direction
• the rate of muscular force production in the horizontal direction
• the rate of muscular force production required to swing the limbs.

From empirical data, knowledge of trigonometry and physics (force mechanics) and of muscle physiology, Pontzer identified the measureable anatomical and motor variables that allowed estimation of those required three force variables. They were length and proportion of limbs, speed, frequency of stride, and angle of excursion. Following earlier studies that linked force production with cost of locomotion, he generated the model — the equation — that he hoped would predict the latter from the former. He found that the model (equation) well predicted the observed cost of locomotion. It appears that the length of the transporting limbs (hip height) inversely inversely predicted energy cost, and that body mass has no independent effect on locomotor cost.

Subsequently, Professor Pontzer tested the model in quadrupeds as well as humans. The model proved superior to previous models and confirmed the predictive ability of considering the proposed anatomical variables in estimating the rate of force production and energy cost of locomotion.

With the development of quadruped and biped robots for human service, Professor Pontzer's model might help make decisions on energy-cost-effective robot locomotor anatomy and dynamics.

Bipedalism in Human Ancestry

Lucy

In one of the most stunning finds in the previous century, a near complete skeleton of a female Australopithecus emerged from the Hadar region of Eastern Africa.

Evolution of Bipedalism

Consider:

• The energetic costs of load-carrying and the evolution of bipedalism

• Gene copy number variation spanning 60 million years of human and primate evolution