Human evolution

Human evolution is the evolutionary process that led to the emergence of anatomically modern humans, beginning with the evolutionary history of primates—in particular genus Homo—and leading to the emergence of Homo sapiens as a distinct species of the hominid family, the great apes. This process involved the gradual development of traits such as human bipedalism and language,[1] as well as interbreeding with other hominins, which indicate that human evolution was not linear but a web.[2][3][4][5]

The study of human evolution involves several scientific disciplines, including physical anthropology, primatology, archaeology, paleontology, neurobiology, ethology, linguistics, evolutionary psychology, embryology and genetics.[6] Genetic studies show that primates diverged from other mammals about 85 million years ago, in the Late Cretaceous period, and the earliest fossils appear in the Paleocene, around 55 million years ago.[7]

Within the Hominoidea (apes) superfamily, the Hominidae family diverged from the Hylobatidae (gibbon) family some 15–20 million years ago; African great apes (subfamily Homininae) diverged from orangutans (Ponginae) about 14 million years ago; the Hominini tribe (humans, Australopithecines and other extinct biped genera, and chimpanzee) parted from the Gorillini tribe (gorillas) between 8–9 million years ago; and, in turn, the subtribes Hominina (humans and biped ancestors) and Panina (chimps) separated 4–7 million years ago.[8]

Anatomical changes

The hominoids are descendants of a common ancestor

Human evolution from its first separation from the last common ancestor of humans and chimpanzees is characterized by a number of morphological, developmental, physiological, and behavioral changes. The most significant of these adaptations are bipedalism, increased brain size, lengthened ontogeny (gestation and infancy), and decreased sexual dimorphism. The relationship between these changes is the subject of ongoing debate.[9][page needed] Other significant morphological changes included the evolution of a power and precision grip, a change first occurring in H. erectus.[10]

Bipedalism

Bipedalism is the basic adaptation of the hominid and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominids. The earliest hominin, of presumably primitive bipedalism, is considered to be either Sahelanthropus[11] or Orrorin, both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the gorilla and chimpanzee, diverged from the hominin line over a period covering the same time, so either of Sahelanthropus or Orrorin may be our last shared ancestor. Ardipithecus, a full biped, arose approximately 5.6 million years ago.[12]

The early bipeds eventually evolved into the australopithecines and still later into the genus Homo. There are several theories of the adaptation value of bipedalism. It is possible that bipedalism was favored because it freed the hands for reaching and carrying food, saved energy during locomotion,[13] enabled long distance running and hunting, provided an enhanced field of vision, and helped avoid hyperthermia by reducing the surface area exposed to direct sun; features all advantageous for thriving in the new savanna and woodland environment created as a result of the East African Rift Valley uplift versus the previous closed forest habitat.[14][13][15] A new study provides support for the hypothesis that walking on two legs, or bipedalism, evolved because it used less energy than quadrupedal knuckle-walking.[16][17] However, recent studies suggest that bipedality without the ability to use fire would not have allowed global dispersal.[18] This change in gait saw a lengthening of the legs proportionately when compared to the length of the arms, which were shortened through the removal of the need for brachiation. Another change is the shape of the big toe. Recent studies suggest that Australopithecines still lived part of the time in trees as a result of maintaining a grasping big toe. This was progressively lost in Habilines.

Anatomically, the evolution of bipedalism has been accompanied by a large number of skeletal changes, not just to the legs and pelvis, but also to the vertebral column, feet and ankles, and skull.[19] The femur evolved into a slightly more angular position to move the center of gravity toward the geometric center of the body. The knee and ankle joints became increasingly robust to better support increased weight. To support the increased weight on each vertebra in the upright position, the human vertebral column became S-shaped and the lumbar vertebrae became shorter and wider. In the feet the big toe moved into alignment with the other toes to help in forward locomotion. The arms and forearms shortened relative to the legs making it easier to run. The foramen magnum migrated under the skull and more anterior.[20]

The most significant changes occurred in the pelvic region, where the long downward facing iliac blade was shortened and widened as a requirement for keeping the center of gravity stable while walking;[21] bipedal hominids have a shorter but broader, bowl-like pelvis due to this. A drawback is that the birth canal of bipedal apes is smaller than in knuckle-walking apes, though there has been a widening of it in comparison to that of australopithecine and modern humans, permitting the passage of newborns due to the increase in cranial size but this is limited to the upper portion, since further increase can hinder normal bipedal movement.[22]

The shortening of the pelvis and smaller birth canal evolved as a requirement for bipedalism and had significant effects on the process of human birth which is much more difficult in modern humans than in other primates. During human birth, because of the variation in size of the pelvic region, the fetal head must be in a transverse position (compared to the mother) during entry into the birth canal and rotate about 90 degrees upon exit.[23] The smaller birth canal became a limiting factor to brain size increases in early humans and prompted a shorter gestation period leading to the relative immaturity of human offspring, who are unable to walk much before 12 months and have greater neoteny, compared to other primates, who are mobile at a much earlier age.[15] The increased brain growth after birth and the increased dependency of children on mothers had a big effect upon the female reproductive cycle,[24] and the more frequent appearance of alloparenting in humans when compared with other hominids.[25] Delayed human sexual maturity also led to the evolution of menopause with one explanation providing that elderly women could better pass on their genes by taking care of their daughter's offspring, as compared to having more children of their own.[26]

Encephalization

Brain size and tooth size in hominins

The human species eventually developed a much larger brain than that of other primates—typically 1,330 cm3 (81 cu in) in modern humans, nearly three times the size of a chimpanzee or gorilla brain.[27] After a period of stasis with Australopithecus anamensis and Ardipithecus, species which had smaller brains as a result of their bipedal locomotion,[28] the pattern of encephalization started with Homo habilis, whose 600 cm3 (37 cu in) brain was slightly larger than that of chimpanzees. This evolution continued in Homo erectus with 800–1,100 cm3 (49–67 cu in), and reached a maximum in Neanderthals with 1,200–1,900 cm3 (73–116 cu in), larger even than modern Homo sapiens. This brain increase manifested during postnatal brain growth, far exceeding that of other apes (heterochrony). It also allowed for extended periods of social learning and language acquisition in juvenile humans, beginning as much as 2 million years ago.

Furthermore, the changes in the structure of human brains may be even more significant than the increase in size.[29][30][31][32]

Three students hold three different skulls in front of their faces, to show the difference in size and shape compared to the modern head
The size and shape of the skull changed over time. The leftmost, and largest, is a replica of a modern human skull.

The temporal lobes, which contain centers for language processing, have increased disproportionately, as has the prefrontal cortex, which has been related to complex decision-making and moderating social behavior.[27] Encephalization has been tied to increased meat and starches in the diet,[33][34][35] and the development of cooking,[36] and it has been proposed that intelligence increased as a response to an increased necessity for solving social problems as human society became more complex.[37] Changes in skull morphology, such as smaller mandibles and mandible muscle attachments, allowed more room for the brain to grow.[38]

The increase in volume of the neocortex also included a rapid increase in size of the cerebellum. Its function has traditionally been associated with balance and fine motor control, but more recently with speech and cognition. The great apes, including hominids, had a more pronounced cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and learning complex muscular actions, the cerebellum may have underpinned human technological adaptations, including the preconditions of speech.[39][40][41][42]

The immediate survival advantage of encephalization is difficult to discern, as the major brain changes from Homo erectus to Homo heidelbergensis were not accompanied by major changes in technology. It has been suggested that the changes were mainly social and behavioural, including increased empathic abilities,[43][44] increases in size of social groups,[45][46][47] and increased behavioural plasticity.[48] Encephalization may be due to a dependency on calorie-dense, difficult-to-acquire food.[49]

Sexual dimorphism

The reduced degree of sexual dimorphism in humans is visible primarily in the reduction of the male canine tooth relative to other ape species (except gibbons) and reduced brow ridges and general robustness of males. Another important physiological change related to sexuality in humans was the evolution of hidden estrus. Humans are the only hominoids in which the female is fertile year round and in which no special signals of fertility are produced by the body (such as genital swelling or overt changes in proceptivity during estrus).[50]

Nonetheless, humans retain a degree of sexual dimorphism in the distribution of body hair and subcutaneous fat, and in the overall size, males being around 15% larger than females.[51] These changes taken together have been interpreted as a result of an increased emphasis on pair bonding as a possible solution to the requirement for increased parental investment due to the prolonged infancy of offspring.[52]

Ulnar opposition

The ulnar opposition—the contact between the thumb and the tip of the little finger of the same hand—is unique to the genus Homo,[53] including Neanderthals, the Sima de los Huesos hominins and anatomically modern humans.[54][55] In other primates the thumb is short and unable to touch the little finger.[54] The ulnar opposition facilitates the precision grip and power grip of the human hand, underlying all the skilled manipulations.

Other changes

A number of other changes have also characterized the evolution of humans, among them an increased importance on vision rather than smell; a longer juvenile developmental period and higher infant dependency; a smaller gut; faster basal metabolism;[56] loss of body hair; evolution of sweat glands; a change in the shape of the dental arcade from being u-shaped to being parabolic; development of a chin (found in Homo sapiens alone); development of styloid processes; and the development of a descended larynx.