Knuckle-walking

A Western Lowland Gorilla knuckle-walking.

Knuckle-walking is a form of quadrupedal walking in which the forelimbs hold the fingers in a partially flexed posture that allows body weight to press down on the ground through the knuckles.

Gorillas, common chimpanzees and bonobos use this style of locomotion as do anteaters and platypuses.

Anthropologists once thought that the common ancestor of chimpanzees and humans engaged in knuckle-walking, and humans developed upright walking from knuckle-walking.

Since then, scientists discovered Ardipithecus ramidus, a human-like hominid descended from the common ancestor of chimpanzees and humans. Ar. ramidus engages in upright walking, but not knuckle-walking. This leads scientists to conclude that chimpanzees and apes developed knuckle-walking after they split from humans 6 million years ago, and humans developed upright walking without knuckle-walking.^[1]

Apes

Common chimpanzees and bonobos (Pan) and gorillas engage in knuckle-walking.^[2] This form of hand-walking posture allows these tree climbers to use their hands for terrestrial locomotion while retaining long fingers for climbing. It may also allow small objects to be carried in the fingers while walking on all fours.

Their knuckle walking involves flexing the tips of their fingers and carrying their body weight down on the dorsal surface of their middle phalanges (middle segments of their fingers). The outer fingers are held clear on the ground. The wrist is held in a stable, locked position during the support phase of knuckle-walking by means of strongly flexed interphalangeal joints, and extended metacarpophalangeal joints. The palm as a result is positioned perpendicular to the ground and in-line with the forearm.^[3][2] The wrist and elbow are extended throughout the last period in which the knuckle-walker's hand carried body weight.^[4]

There are differences between chimpanzees and gorillas: juvenile chimpanzees and bonobos engage in less knuckle-walking than juvenile gorillas. Another difference is that the hand bones of gorillas lack key features that were once thought to limit the extension of the wrist during knuckle-walking in chimpanzees and bonobos. For example, the ridges and concavities features of the capitate and hamate bones have been interpreted to enhance stability of weight-bearing; on this basis, they have been used to identify knuckle-walking in fossils. These are found in all chimpanzees and bonobos but in only two out of five gorillas. They are also less prominent when found in gorillas. They are however found in primates that do not knuckle-walk.^[5]

It has been suggested that chimpanzee/bonobo and gorilla knuckle walking are biomechanically and posturally distinct. Gorilla use a form of knuckle-walking which is ‘‘columnar’’. In this forelimb posture the hand and wrist joints are aligned in a relatively straight, neutral posture. In contrast, chimpanzees and bonobos use an extended wrist posture. These differences underlie the different characteristics of their hand bones.^[5]

The difference has been attributed to the greater locomotion of chimpanzees/bonobos in trees, compared to gorillas. The former frequently engage in both knuckle-walking and palm-walking branches. As a result, to preserve their balance in trees chimpanzees and bonobos, like other primates in trees, often extended their wrists. This need has produced different wrist bone anatomy and, through this, a different form of knuckle-walking.^[5]

Knuckle-walking has been reported in some baboons.^[6] It has also been suggested that fossils attributed to Australopithecus anamensis and A. afarensis had specialized wrist morphology that was retained from an earlier knuckle-walking ancestor.^[7][2]

Nonprimates

Giant anteaters^[8] and platypuses^[9] are also knuckle-walkers. Pangolins also sometimes walk on their knuckles. Another possible knuckle-walking taxon was the extinct chalicotheres, which looked something like a cross between a horse and a gorilla.^[10] The ground sloths may have also walked on their knuckles.

Advantages

Knuckle-walking tends to evolve when the fingers of the forelimb are specialized for tasks other than locomotion on the ground. In the gorilla the fingers are used for the manipulation of food, and in chimpanzees for the manipulation of food and for climbing. In anteaters and pangolins the fingers have large claws for opening the mounds of social insects. Platypus fingers have webbing that extend past the fingers to aid in swimming, thus knuckle-walking is used to prevent stumbling.^[9]

It has been argued that knuckle-walking of chimpanzees and gorillas arose originally from fist-walking as found in orangutans.^[11]

Human evolution

One theory of the origins of human bipedality is that it evolved from a terrestrial knuckle-walking ancestor. This theory is opposed to the theory that such bipedalism arose from a more generalized arboreal ape ancestor. The terrestrial knuckle-walking theory argues that early hominin wrist and hand bones retain morphological evidence of early knuckle-walking.^[7][2] The argument is not that they were knuckle-walkers themselves but that it is an example of "phylogenetic “lag”".^[2] "The retention of knuckle-walking morphology in the earliest hominids indicates that bipedalism evolved from an ancestor already adapted for terrestrial locomotion. ... Pre-bipedal locomotion is probably best characterized as a repertoire consisting of terrestrial knuckle-walking, arboreal climbing and occasional suspensory activities, not unlike that observed in chimpanzees today".^[7]

It has however been suggested that knuckle-walking evolved independently and separately in Pan and gorillas and so was not present in the human ancestors.^[12][5] This is supported by the evidence that gorilla and chimpanzee differ in their knuckle-walking related wrist anatomy and in the biomechanics of their knuckle-walking.^[5]

Indeed as Tracy L. Kivell and Daniel Schmitt note "Features found in the hominin fossil record that have traditionally been associated with a broad definition of knuckle-walking are more likely reflecting the habitual Pan-like use of extended wrist postures that are particularly advantageous in an arboreal environment. This, in turn, suggests that human bipedality evolved from a more arboreal ancestor occupying a generalized locomotor and ecological niche common to all living apes".^[5]

Related forms of hand-walking

Primates can walk on their hands in other ways than on their knuckles. They can walk on fists such as orangutans. In this body weight is borne on the back of the proximal phalanges.^[13]

Quadrupedal primate walking can be done on the palms. This occurs in many primates when walking on all fours on tree branches.^[14][15] It is also the method used by human infants when crawling on their knees or engaged in a "bear-crawl" (in which the legs are fully extended and weight in taken by the ankles). A few older children and some adults retain the ability to walk quadrupedally, even after acquiring bipedally.^[16] A BBC2 and NOVA in The Family That Walks On All Fours reported on the Ulas family in which five individuals grew up walking normally upon the palms of their hands and fully extended legs due to a recessive genetic mutation that causes a non-progressive congenital cerebellar ataxia that impairs the balance need for bipedality.^[17] Not only did they walk on their palms of their hands but could do so holding objects in their fingers.^[17]

Primates can also walk on their fingers.^[14][15] In Olive baboons, Rhesus Macaques, and Patas Monkeys such finger walking turns to palm walking when animals start to run.^[15] This has been suggested to spread the forces better across the wrist bones to protect them.^[15]

References