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By 1989 the market for cartridge-based console games was more than $2 billion, while that for disk-based computer games was less than $300 million.[29] Large computer-game companies such as Epyx, Electronic Arts, and LucasArts began devoting much or all of their attention on console games. Computer Gaming World warned that computer gaming could become a “cultural backwater,” similar to what had happened a few years earlier with 8-bit computers.[30] In 1990, Commodore and Amstrad entered the console market with their C64GS and GX4000 game machines respectively. These were both based on the 8-bit computers of their manufacturers, and had only limited success due to a lack of software support and the arrival of 16-bit machines. Amstrad’s GX4000 sold just over 15,000 units, with only 25 officially released game cartridges. Even though it was technically superior to the Master System and Nintendo Entertainment System, it was discontinued after 6 months.

This generation ended with the discontinuation of the NES in 1995.
A 16-bit generation of video game consoles starts in the late 1980s. The TurboGrafx-16, named the PC Engine in Europe and Japan, debuted in 1987 as the first commercial 16-bit game system. It had a large following in Japan, but, did poorly in North America and Europe because of its limited library of games and because of excessive distribution restrictions imposed by Hudson Soft. Sega’s Mega Drive/Genesis sold well world-wide early on after its debut in 1988. Nintendo responded with its own next generation system named the Super Nintendo Entertainment System (SNES), in 1990.

TurboGrafx-16 was billed as the first 16-bit system but its central processor was an 8-bit HuC6280, with only its HuC6270 graphics processor being a true 16-bit chip. Also, the much earlier Mattel Intellivision contained a 16-bit processor. Sega used the term “Blast Processing” throughout its marketing to describe the simple fact that their console’s CPU ran at a higher clock speed than that of the SNES (7.67 MHz vs 3.58 MHz).

In Japan, the PC Engine was a very successful competitor against the Famicom and a CD drive peripheral allowed it to fend off the Mega Drive in 1988, though it never really caught on to the same degree outside Japan. The PC Engine eventually lost out to the Super Famicom, but, due to its popular CD add-ons, retained enough of a user base to support new games well into the late 1990s.

CD-ROM drives were introduced in this generation, as add-ons for the PC Engine in 1988 and the Mega Drive in 1991. Nintendo experimented with optical media formats for the SNES in a joint venture with Sony, who would go on to develop this concept into the PlayStation and rise to prominence as a major competitor to Nintendo and Sega. Basic 3D graphics entered the mainstream with flat-shaded polygons enabled by added processors in game cartridges like Virtua Racing and Star Fox, while the Mega Drive managed to produce such graphics without special processors, on the ~8 MHz 68000 chip by using highly simplified polygon models, a slow frame rate (<4 fps), and reduced resolution.

Sonic the Hedgehog, released in 1991 for the Mega Drive/Genesis, gave the console mainstream popularity, and rivaled Nintendo’s Mario franchise, starting the so-called “console war.” Its namesake character became the mascot of Sega and one of the most recognizable video game characters in history.

SNK’s Neo-Geo was the most costly console by a wide margin when released in 1990, and would remain so for years. Its 2D graphics were of a quality years ahead of other consoles. The reason for this was that it contained the same hardware that was found in SNK’s arcade games. This was the first time since the home Pong machines that a true-to-the-arcade experience could be had at home, but the system was commercially inviable.

This era also saw a revival of handheld consoles, which were absent in the prior generation. Nintendo’s Game Boy, a portable released in 1989 with monochromatic 2D graphics and 35-hours battery life, became widely popular in the world and sold much more than its three competitors, the Atari Lynx, the Sega Game Gear and NEC’s Turbo Express, released in Japan in North America until 1991. Despite these three consoles had much more sophisticated 16-bit graphics (similar to home consoles of the time), graphic resource consumed too much battery life, what made the consoles unpopular and with scarce game libraries compared to that of the Game Boy that, with more than one thousand games released, inspired an entire line of portable machines that continued through the following two generations.

The 1990s were a decade of marked innovation in video gaming. It was a decade of transition from raster graphics to 3D graphics and gave rise to several genres of video games including first-person shooter, real-time strategy, and MMO. Handheld gaming began to become more popular throughout the decade, thanks in part to the release of the Game Boy in 1989.[31] Arcade games experienced a resurgence in the early-to-mid-1990s, followed by a decline in the late 1990s as home consoles became more common.

 

As arcade games declined, however, the home video game industry matured into a more mainstream form of entertainment in the 1990s, but their video games also became more and more controversial because of their violent nature, especially in games of Mortal Kombat, Night Trap, and Doom, leading to the formation of the Interactive Digital Software Association and their rating games by signing them their ESRB ratings since 1994.[32] Major developments of the 1990s include the popularizing of 3D computer graphics using polygons (initially in arcades, followed by home consoles and computers), and the start of a larger consolidation of publishers, higher budget games, increased size of production teams, and collaborations with both the music and motion picture industries. Examples of this include Mark Hamill’s involvement with Wing Commander III, the introduction of QSound with arcade system boards such as Capcom’s CP System II, and the high production budgets of titles such as Squaresoft’s Final Fantasy VII and Sega’s Shenmue.
Resurgence and decline of arcades

In North America, arcade games, which had seen a slow decline with the increase in popularity of home gaming, experienced a resurgence in the early-to-mid-1990s, with the 1991 release of Capcom’s Street Fighter II[33] popularizing competitive one-on-one fighting games and reviving the arcade industry to a level of popularity not seen since the days of Pac-Man.[34] Its success led to a wave of other popular fighting games, such as Mortal Kombat and The King of Fighters. Sports games such as NBA Jam also briefly became popular in arcades during this period.

 

 

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Human evolution is characterized by a number of morphological, developmental, physiological, and behavioral changes that have taken place since the split between the last common ancestor of humans and chimpanzees. 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.[55][page needed] Other significant morphological changes included the evolution of a power and precision grip, a change first occurring in H. erectus.[56]
Bipedalism

Bipedalism is the basic adaptation of the hominin and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominins. The earliest hominin, of presumably primitive bipedalism, is considered to be either Sahelanthropus[57] 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 somewhat later.[citation needed]

The early bipeds eventually evolved into the australopithecines and later 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,[58] 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 environment versus the previous forest habitat.[32][58][59] 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.[60][61] However, recent studies suggest that bipedality without the ability to use fire would not have allowed global dispersal.[62]

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.[63] 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.[64]

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;[65] 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.[66]

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.[67] 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.[59] The increased brain growth after birth and the increased dependency of children on mothers had a big effect upon the female reproductive cycle,[68] and the more frequent appearance of alloparenting in humans when compared with other hominids.[69] 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 of their own.
The human species developed a much larger brain than that of other primates—typically 1,330 cm3 in modern humans, over twice the size of that of a chimpanzee or gorilla.[71] The pattern of encephalization started with Homo habilis,[72] which at approximately 600 cm3 had a brain slightly larger than that of chimpanzees, and continued with Homo erectus (800–1,100 cm3), reaching a maximum in Neanderthals with an average size of (1,200–1,900 cm3), larger even than Homo sapiens. The pattern of human postnatal brain growth differs from that of other apes (heterochrony) and allows for extended periods of social learning and language acquisition in juvenile humans. However, the differences between the structure of human brains and those of other apes may be even more significant than differences in size.[73][74][75][76]

The increase in volume over time has affected areas within the brain unequally—the temporal lobes, which contain centers for language processing, have increased disproportionately, and seems to favor a belief that there was evolution after leaving Africa, as has the prefrontal cortex which has been related to complex decision-making and moderating social behavior.[71] Encephalization has been tied to an increasing emphasis on meat in the diet,[77][78][79] or with the development of cooking,[80] 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.[81] The human brain was able to expand because of the changes in the morphology of smaller mandibles and mandible muscle attachments to the skull into allowing more room for the brain to grow.[82]

The increase in volume of the neocortex also included a rapid increase in size of the cerebellum. Traditionally the cerebellum has been associated with a paleocerebellum and archicerebellum as well as a neocerebellum. Its function has also traditionally been associated with balance, fine motor control but more recently speech and cognition. The great apes including humans and its antecessors had a more pronounced development of the cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and assisting in learning complex muscular action sequences, the cerebellum may have underpinned the evolution of human’s technological adaptations including the preadaptation of speech.

The reason for this encephalization is difficult to discern, as the major changes from Homo erectus to Homo heidelbergensis were not associated with major changes in technology. It has been suggested that the changes have been associated with social changes, increased empathic abilities[87][88] and increases in size of social groupings

The reduced degree of sexual dimorphism 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 and bonobos are the only apes 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 during estrus).

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. 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.
A number of other changes have also characterized the evolution of humans, among them an increased importance on vision rather than smell; a smaller gut; 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.
Evidence
The evidence on which scientific accounts of human evolution are based comes from many fields of natural science. The main source of knowledge about the evolutionary process has traditionally been the fossil record, but since the development of genetics beginning in the 1970s, DNA analysis has come to occupy a place of comparable importance. The studies of ontogeny, phylogeny and especially evolutionary developmental biology of both vertebrates and invertebrates offer considerable insight into the evolution of all life, including how humans evolved. The specific study of the origin and life of humans is anthropology, particularly paleoanthropology which focuses on the study of human prehistory.

The closest living relatives of humans are bonobos and chimpanzees (both genus Pan) and gorillas (genus Gorilla).[93] With the sequencing of both the human and chimpanzee genome, current estimates of the similarity between their DNA sequences range between 95% and 99%.[93][94][95] By using the technique called the molecular clock which estimates the time required for the number of divergent mutations to accumulate between two lineages, the approximate date for the split between lineages can be calculated.

The gibbons (family Hylobatidae) and then orangutans (genus Pongo) were the first groups to split from the line leading to the hominins, including humans—followed by gorillas, and, ultimately, by the chimpanzees (genus Pan). The splitting date between hominin and chimpanzee lineages is placed by some between 4 to 8 million years ago, that is, during the Late Miocene.[3][96][97] Speciation, however, appears to have been unusually drawn-out. Initial divergence occurred sometime between 7 to 13 million years ago, but ongoing hybridization blurred the separation and delayed complete separation during several millions of years. Patterson (2006) dated the final divergence at 5 to 6 million years ago.[98]

Genetic evidence has also been employed to resolve the question of whether there was any gene flow between early modern humans and Neanderthals, and to enhance our understanding of the early human migration patterns and splitting dates. By comparing the parts of the genome that are not under natural selection and which therefore accumulate mutations at a fairly steady rate, it is possible to reconstruct a genetic tree incorporating the entire human species since the last shared ancestor.

Each time a certain mutation (Single-nucleotide polymorphism) appears in an individual and is passed on to his or her descendants a haplogroup is formed including all of the descendants of the individual who will also carry that mutation. By comparing mitochondrial DNA which is inherited only from the mother, geneticists have concluded that the last female common ancestor whose genetic marker is found in all modern humans, the so-called mitochondrial Eve, must have lived around 200,000 years ago.
Genetics
Main articles: Human evolutionary genetics and Human genetic variation

Human evolutionary genetics studies how one human genome differs from the other, the evolutionary past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical and forensic implications and applications. Genetic data can provide important insight into human evolution.
Evidence from the fossil record