Sonic the Hedgehog[a] is a video game series and media franchise created by the Japanese developers Yuji Naka, Naoto Ohshima, and Hirokazu Yasuhara for Sega. The franchise follows Sonic, an anthropomorphic blue hedgehog who battles the evil Doctor Eggman, a mad scientist. The main Sonic the Hedgehog games are platformers mostly developed by Sonic Team; other games, developed by various studios, include spin-offs in the racing, fighting, party and sports genres. The franchise also incorporates printed media, animations, feature films, and merchandise.
The Sonic franchise is known for its large cast of characters;[194] Sonic the Fighters (1996) producer Yu Suzuki joked that anyone who makes a Sonic game has the duty to create new characters.[195] The first game introduced Sonic, a blue hedgehog who can run at incredible speeds, and Doctor Eggman, a rotund mad scientist.[4] During the Genesis era, Eggman was referred to as Doctor Ivo Robotnik in Western territories.[196][197] Sega of America's Dean Sitton made the change[198] without consulting the Japanese developers, who did not want a single character to have two different names. Since Sonic Adventure, the character has been referred to as Eggman in all territories,[196] although the Robotnik name is still acknowledged.[197][199][200]
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Much of the supporting cast was introduced in the succeeding games for the Genesis and its add-ons. Sonic 2 introduced Sonic's sidekick Miles "Tails" Prower, a fox who can fly using his two tails.[209] Sonic CD introduced Amy Rose, a pink hedgehog and Sonic's self-proclaimed girlfriend, and Metal Sonic, a robotic doppelgnger of Sonic created by Eggman.[210] Sonic 3 introduced Sonic's rival Knuckles, a red echidna and the guardian of the Master Emerald.[211] The Master Emerald, introduced in Sonic & Knuckles,[212] controls the power of the Chaos Emeralds.[202] Knuckles' Chaotix introduced the Chaotix, a group comprising Vector the Crocodile, Espio the Chameleon and Charmy Bee.[213] Three characters introduced during this period, Mighty the Armadillo and Ray the Flying Squirrel from SegaSonic the Hedgehog and Fang the Sniper from Sonic Triple Trouble (1994), faded into obscurity,[37][214] but became prominent characters again in Sonic Mania and Superstars.[215]
During Sonic Adventure's development, Sonic Team discovered that the relatively simple character designs did not suit a 3D environment. The art style was modernized to alter the characters' proportions and make them appeal to Western audiences.[72] Since Sonic Adventure, the series' cast has expanded.[194] Notable characters include Big,[216] a large cat who fishes for his pet frog Froggy;[217][218] the E-100 Series of robots;[219] Shadow, a brooding black hedgehog;[216] Rouge, a treasure-hunting bat;[220] Blaze, a cat from an alternate dimension;[221] and Silver, a telekinetic hedgehog from the future.[216] The Chao creatures function as digital pets and minor gameplay elements,[222] and Wisp creatures function as power-ups.[223] Flicky, the blue bird from Sega's 1984 arcade game, appears in several Sonic games, most notably 3D Blast.[224]
Shogakukan published a Sonic the Hedgehog manga series in its Shogaku Yonensei magazine, beginning in 1992. Written by Kenji Terada and illustrated by Sango Norimoto, it follows a sweet but cowardly young hedgehog named Nicky whose alter ego is the cocky, heroic Sonic.[363] According to character artist Kazuyuki Hoshino, the manga was part of Sega's promotional strategy to appeal to primary school children.[364] The Sonic design team worked with Shogakukan to create new characters; Amy Rose and Charmy Bee originated in the manga before appearing in the games.[37][364]
The sonic hedgehog gene, first identified for its role in fruit fly embryonic development, was named after Sonic.[555][556] Robert Riddle, then a postdoctoral fellow at the Tabin Lab,[557] came up with the name after his wife bought a magazine containing an advert for Sonic.[558][559] A mutation in the gene causes fly larvae to bear spiky denticles, reminiscent of Sonic.[555]
Sonic the Hedgehog[c] is a 1991 platform game developed by Sonic Team and published by Sega for the Genesis/Mega Drive. It was released in North America on June 23 and in PAL regions and Japan the following month. Players control Sonic the Hedgehog, who can run at near supersonic speeds; Sonic sets out on a quest to defeat Dr. Robotnik, a scientist who has imprisoned animals in robots and seeks the powerful Chaos Emeralds. The gameplay involves collecting rings as a form of health, and a simple control scheme, with jumping and attacking controlled by a single button.
Development began in 1990 when Sega ordered its developers to create a game featuring a mascot for the company. The developers chose a blue hedgehog designed by Naoto Oshima after he won an internal character design contest, and named themselves Sonic Team to match their character. It uses a novel technique that allows Sonic's sprite to roll along curved scenery which was based on a concept by Oshima from 1989.[2] Sonic the Hedgehog, designed for fast gameplay, was influenced by games by Super Mario series creator Shigeru Miyamoto. The music was composed by Masato Nakamura, bassist of the J-pop band Dreams Come True.
The team developed ideas for characters, an engine, and gameplay mechanics. Development emphasized speed, so Sega considered fast creatures such as kangaroos and squirrels and eliminated designs not associated with fast animals.[3] One idea, a rabbit able to grasp objects with prehensile ears, showed promise but was too complex for the Genesis hardware. The team narrowed its search to animals that could roll into a ball, their idea for an attacking move, and considered armadillos and hedgehogs.[12] The hedgehog character, proposed by Naoto Ohshima,[15] prevailed. Ohshima went on vacation to New York, taking sketches with him. He went to Central Park and asked locals for their opinions on them, and Sonic was the favorite. A man with a moustache, who eventually became Dr. Robotnik, was in second place.[16]
Ohshima said that "Sonic" was chosen because it represented speed.[22] Ideas proposed to flesh out the character included placing him in a rock band, giving him vampire fangs, and giving him a human girlfriend named Madonna.[23] Sega of America scrapped these ideas to keep his identity simple. Sega of America also expressed concerns that most Americans would not know what a hedgehog is.[15] According to Mark Cerny, who worked in Tokyo as an intermediary between the Japanese and American Sega offices, the American staff felt the character was "unsalvageable". They made plans to educate Sonic Team on character design with the work of Will Vinton, the creator of the California Raisins.[24] They proposed a full-scale recreation of the character, but compromised with Sonic Team to simply make design changes for western audiences.[15] The antagonist was named "Dr. Eggman" in Japan and "Dr. Robotnik" in other regions as a result of a dispute between Sega's American and Japanese divisions.[18]
Sonic the Hedgehog was developed by a team of seven: two programmers, two sound engineers, and three designers,[27] although it began with just Ohshima and Naka.[22] People came onto the team as the need for content increased.[22] After being assigned a project with the code name "Defeat Mario", Ohshima and Naka began work, but encountered problems: Ohshima's Rabbit proved hard to program. Catching items and throwing them caused the action's rhythm to break. Naka stated that the rabbit was not suitable for his game engine, and he also wanted the game to be playable with only one button. Hirokazu Yasuhara came onto the team to supervise Ohshima and Naka and develop levels. He became the lead designer due to his greater experience, and found the way to make the game playable with only one button by having Sonic do damage by jumping. The trio came up with the idea of him rolling into a ball. After the hedgehog character was chosen, many characters were redrawn, and the team agreed on the environments' visual complexity, with particular focus on the colors. After this, four people came onto the team to speed development up.[28]
Sonic hedgehog (SHH) as a secreted protein is important for neuronal development in the central nervous system (CNS). However, the mechanism about SHH release remains largely unknown. Here, we showed that SHH was expressed mainly in the synaptic vesicles of hippocampus in both young postnatal and adult rats. High, but not low, frequency stimulation, induces SHH release from the neurons. Moreover, removal of extracellular Ca2+, application of tetrodotoxin (TTX), an inhibitor of voltage-dependent sodium channels, or downregulation of soluble n-ethylmaleimide-sensitive fusion protein attachment protein receptors (SNAREs) proteins, all blocked SHH release from the neurons in response to HFS. Our findings suggest a novel mechanism to control SHH release from the hippocampal neurons.
Among the many morphogenic signaling cascades in embryonic and adult development18,34,35, hedgehog signaling remains one of the least understood. Though SHH has been best known as a key regulator during embryonic development, components of SHH signaling pathway continue to express in the adult brain5,36,37. Its transcripts are found in GABAergic neurons located in various basal forebrain nuclei38. Moreover, the subcellular localization of SHH in primary neurons has been shown15. However, where SHH signaling takes place in the adult brain remains largely unknown. The preferential distribution of PTCH1 and SMO in the post-synapse of hippocampal neurons7 and our current results showing the synaptic localization of SHH in the postnatal and adult rat hippocampus, are consistent with the reported roles of SHH in axonal or dendritic growth39,40,41. ff782bc1db
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