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Mario & Luigi: Superstar Saga
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Mario & Luigi: Superstar Saga
North American box art
Developer(s)
AlphaDream
Publisher(s)
Nintendo
Director(s)
Yoshihiko Maekawa
Shunsuke Kobayashi (3DS)
Producer(s)
Shigeru Miyamoto
Tetsuo Mizuno
Artist(s)
Toshizo Morikawa
Writer(s)
Hiroyuki Kubota
Composer(s)
Yoko Shimomura
Series
Mario & Luigi
Platform(s)
Game Boy Advance
Nintendo 3DS
Release
Game Boy Advance
NA: November 17, 2003
JP: November 21, 2003
EU: November 22, 2003
AU: November 28, 2003
Nintendo 3DS
JP: October 5, 2017
NA: October 6, 2017
EU: October 6, 2017
AU: October 7, 2017
Genre(s)
Role-playing video game
Mode(s)
Single-player, multiplayer
Mario & Luigi: Superstar Saga[a] is a role-playing video game developed by AlphaDream and published by Nintendo for the Game Boy Advance in 2003. The game is the first in the Mario & Luigi RPG series. The game was later re-released for the Wii U Virtual Console on the Nintendo eShop in 2014.
The setting of the game begins in the Mushroom Kingdom, but progresses to the Beanbean Kingdom for the majority of the game. In the game, Mario and Luigi combat Cackletta, the primary antagonist. The quest begins when Cackletta, with the aid of her assistant Fawful, steals Princess Peach's voice after adopting the guise of an ambassador from the Beanbean Kingdom.[1]
A role-playing game, it centers on a battle system different from that of traditional games of the genre, with emphasis on timing and more elaborate attacks.[2] The game is whimsical in tone, with various in-game jokes and comical references to the heritage of the Mario series. Superstar Saga was critically acclaimed, and IGN named it the twelfth best Game Boy Advance game of all time in their feature reflecting on the Game Boy Advance's lifespan.[3] A remake of the game for Nintendo 3DS, Mario & Luigi: Superstar Saga + Bowser's Minions, was released on October 6, 2017.
Contents
[hide]
1 Gameplay
2 Plot
2.1 Characters
2.2 Story
3 Development and marketing
4 Reception
5 Sequels and remake
5.1 Nintendo 3DS remake
6 Notes
7 References
8 External links
Gameplay[edit]
Superstar Saga's battle system
The gameplay of Mario & Luigi differs from most other role-playing games due its focus on controlling Mario and Luigi simultaneously. During overworld sections, the directional pad controls Mario's movement with Luigi following closely, while Mario and Luigi's other actions are controlled individually with the A (Mario) and B (Luigi) buttons respectively.[4] The game begins with them being able to jump independently, though they will soon gain access to hammers and a variety of other techniques as the game progresses. For example, Luigi's hammer can be used to squash Mario into a smaller size, allowing to access small gaps, while placing Mario on Luigi's shoulders allows them to act like a propeller and hover across large gaps.[5] Various enemies roam the overworld, and coming into contact with these enemies initiates a battle. Landing a hit on the enemy while on the overworld allows the player to deal pre-emptive damage, while the opposite is also possible.
Battles in Superstar Saga are turn-based. Mario and Luigi can attack normally either by jumping, which can deal multiple hits but when used against enemies covered in flames or with spikes, Mario or Luigi will get hurt, instead of the enemy. Another attack option is the hammer, which is powerful, but ineffective against flying enemies. Similar to previous Mario RPG titles, such as Super Mario RPG and the Paper Mario series, players can time button presses to make their attacks more effective, such as earning an extra jump attack or increasing the hammer's power.[6] Introduced in this series is the way in which Mario and Luigi can defend themselves during an enemy's attack. When an enemy attacks, the brothers will be able to either jump or use their hammer which, when successfully timed, allows them to dodge their attacks and even deal counter damage (for example: if they jump on top of a charging Goomba). Throughout the game, players can unlock Bros. Attacks, which use Bros. Points (BP) which requires players to cooperate between Mario and Luigi's actions to perform powerful combination attacks.[2][7] Players can also use items such as mushrooms for healing, peppers for boosting stats and 1UP mushrooms for reviving fallen bros. Defeating enemies earns experience points which help the bros level up and increase their stats, with players given the option to further increase the stats of one attribute every time they level up. Players can further improve their stats by equipping new gear to the Bros., or making them wear badges that give them special attributes.
Like other Mario titles for the Game Boy Advance, Superstar Saga features the enhanced remake of the Mario Bros. arcade game.[8][9] The game also supports rumble functionality when used with the GameCube's Game Boy Player accessory.[4]
Plot[edit]
Unlike most games in the Mario series, this game is not set in the Mushroom Kingdom.[10] Instead, Superstar Saga introduces the Beanbean Kingdom, a neighboring monarchy to the Mushroom Kingdom. Despite this, traditional enemies such as Goombas are present, as well as new enemies exclusive to the game. The predominant race are the "Beanish", although there are other species such as the "Hoohooligans" and Yoshis. No locations featured in the Beanbean Kingdom, with the exception of Bowser's Castle, existed in any earlier Mario series games.
Characters[edit]
The playable protagonists in the game are Mario and Luigi, whose quest is to rescue the voice of Princess Peach, the princess of the Mushroom Kingdom. The voice is stolen by the game's antagonists Cackletta and Fawful.[11] Throughout the game, Mario and Luigi must work with the Beanbean monarchy headed by Queen Bean. Both she and her son, Prince Peasley, offer Mario and Luigi assistance. The youthful Prince Peasley is a celebrity figure in his native Beanbean Kingdom.
Deviating from a tradition of the Mario franchise, Superstar Saga's central antagonist is not Bowser, but the Beanish witch Cackletta.[12] Bowser aids the protagonists during a period of the game, but Bowser's body becomes possessed by the consciousness of Cackletta to form the creature Bowletta. Cackletta is aided by the assailant Fawful, who is often characterized by his comical use of language, as evidenced by his famous quote "I HAVE FURY!" and who is also the antagonist in the sequel Mario & Luigi: Bowser's Inside Story. Additionally, Mario and Luigi have recurring incidents with a thief named Popple and his sidekick Rookie (amnesiac Bowser), who engages both protagonists in conflict at various times throughout the game.[13] In one case, Popple fights alone. In another, he uses Birdo instead of Rookie.
Story[edit]
The game opens with the witch Cackletta and her assistant Fawful arriving to steal Princess Peach's voice and replace it with explosives. After being summoned to the palace by Toad, Mario and Luigi confront and subdue Bowser, who coincidentally was attempting to kidnap Peach. Bowser decides against doing so, and teams up with Mario and Luigi to take her voice back from Cackletta, who has fled to the Beanbean Kingdom, with the intention of kidnapping Peach afterward when her voice would not destroy his castle.[14]
Mario, Luigi, and Bowser fly to the Beanbean Kingdom via Bowser's airship, the Koopa Cruiser. Mid-flight, Fawful attacks the cruiser and blows it out of the sky. Bowser and the brothers become separated. On their journey, the brothers rescue Prince Peasley, who had been transformed into a dragon by Cackletta. The brothers reach Beanbean Castle Town and step into a trap laid by Cackletta that drops them into the sewers. The brothers learn that Cackletta's goal is to steal the Beanstar, a mystical object inside the castle which once awoken with Peach's voice will grant all of her wishes.[15] Cackletta takes the Beanstar and attacks Mario and Luigi with a mutated form of the Queen Bean during her escape. After defeating the Queen, Mario and Luigi travel to Chucklehuck Woods to retrieve the Chuckola Reserve, a soda which will restore Queen Bean to her former state. They encounter Popple, a thief also after the Chuckola Reserve, with an amnesiac Bowser as his accomplice, named Rookie. The brothers manage to obtain the Reserve, return to Beanbean Castle Town, and cure Queen Bean.[14]
Mario and Luigi then travel to Woohoo Hooniversity, where Cackletta plans to awaken the Beanstar. The Beanstar is exposed to Peach's voice, which causes it to go berserk and flee. Mario and Luigi then battle Cackletta and defeat her. Cackletta is severely incapacitated from the battle, so Fawful sucks up her soul into his "vacuum helmet" to save her life. Cackletta then commands Fawful to attack, but he is stopped by Prince Peasley. The brothers locate the Beanstar but encounter Popple and Rookie again. After battling them, the Beanstar is again exposed to Peach's voice. The four grab onto the Beanstar in an effort to keep it under control, but it shoots into the sky. The Beanstar explodes into four pieces, which scatter across Beanbean Kingdom, and the brothers become separated from Popple and Bowser. Mario and Luigi backtrack and meet Peach as she arrives in Beanbean Kingdom, who inexplicably has her own voice. They discover that Prince Peasley had learned of Cackletta's plot and warned the Princess beforehand and Birdo, disguised as Peach, had her voice stolen instead. Bowser's arrival and presence at her castle had prevented Peach from revealing the plan to Mario and Luigi. Meanwhile, Fawful imbues a weak Bowser with Cackletta's soul, resulting in the twisted Bowletta.[16]
On a trip to Little Fungitown, Peach is kidnapped by Bowletta, who demands all the pieces of the Beanstar in exchange. After restoring the Beanstar, Mario and Luigi travel to Joke's End to make the exchange. Bowletta refuses to hand over Peach, but Luigi, disguised in Peach's spare dress, ends up being taken in Peach's stead. He escapes from the repaired Koopa Cruiser, reclaiming the Beanstar in the process. The brothers return to Beanbean Castle Town, finding it under attack by Bowletta using Bowser's flying castle. The brothers and Prince Peasley fly up to it to stop Cackletta. Mario and Luigi battle past the Koopalings and Fawful before confronting Bowletta. Feinting defeat, she tricks the brothers and swallows them. After waking up in her belly, the brothers battle and defeat Cackletta's soul, exorcising her from Bowser's body. The two are regurgitated and Bowser returns to normal. Afterwards, the brothers escape the castle before it explodes from a bomb set by Peasley. The explosion sends the castle falling into the ocean.[14] Exhausted from their vacation, Mario, Luigi, Peach and Toadsworth decide to head back to the Mushroom Kingdom. Before they can leave, Peasley brings a farewell gift for them to take home, which is Bowser, who regained memory. The gang then returns home.
Development and marketing[edit]
Superstar Saga, developed by AlphaDream, is said to take its inspiration from the Nintendo 64 game Paper Mario;[10][17] the two games have similar graphics and gameplay.[18][19] The producers of the game were Shigeru Miyamoto, the creator of the Mario franchise, Tetsuo Mizuno, and Satoru Iwata, the president of Nintendo.[20] The voice acting for Mario and Luigi in the game is provided by Charles Martinet, well known for providing the characters' voice in Nintendo's Mario franchise.[17] The game's music was composed by Yoko Shimomura, who also composed the score for Super Mario RPG.[21]
Superstar Saga was first revealed at E3 2003 under the name Mario and Luigi, where a playable demo of the game was available.[22] In August and September 2003, a playable demonstration was also available at the European Computer Trade Show, the Games Convention, and Nintendo Gamers' Summit.[17][23][24] To link in with the game's comic themes, Nintendo organized an official competition between October and November 2003 in which contestants would try to submit the best knock-knock joke to win a Game Boy Advance SP and a copy of the game. Nintendo employed comedian Kathy Griffin to choose the winner.[25]
Reception[edit]
Reception
Aggregate score
Aggregator
Score
Metacritic 90/100[26]
Review scores
Publication
Score
EGM 8.83/10[27]
Eurogamer 9/10[10]
Game Informer 9.5/10[27]
GameSpot 9.2/10[4]
IGN 9/10[28]
Nintendo Power 4.7/5[27]
Superstar Saga received "universal acclaim", according to the review aggregator Metacritic.[26] The game's comical dialog and themes in particular were lauded by critics. Eurogamer's Tom Bramwell commented that "each line of dialogue and identifiable cameo is handled with a loving sense of humour."[10] Despite this, RPGamer's Andrew Long labelled the plot as repetitive, and the game's characters as "a tad shallow".[29] While also appreciating references to the heritage of the Mario series,[30] critics praised the game for avoiding clichés common in previous games of the Mario series.[10]
The gameplay attained a mixed reception. Critics enjoyed the game's battle system, which deviated from role-playing game tradition.[4] IGN's Craig Harris commented that "unlike most Japanese RPGs Mario & Luigi's turn-based battle involves the player at all times".[28] Despite this novel approach to combat situations, some reviewers thought that the overall gameplay lacked innovation.[12][31] GameSpy in particular criticized the game for an apparent lack of originality, commenting that "in terms of gameplay, there isn't much there that we haven't seen in the NES and SNES Mario and Zelda titles."[12] Furthermore, some reviewers were disappointed by a perceived lack of difficulty in the gameplay as a result of targeting a younger audience.[31] However, Cubed3 welcomed this approach, citing that "any gamer, be them [sic] veterans to the genre or complete amateurs, will find the title to be perfectly accessible."[1] Edge and other gaming publications have criticized the controls for being occasionally confusing when considering the usage of jumping, hammers, and other combinations between the two characters.[32]
A common concern among reviewers is the overhead perspective, which critics have bemoaned for preventing them from judging pathway routes and an object's location in relation to its background.[1][28] Besides this, the actual visuals were generally well received,[4][10] as well as the setting and animations. The audio was commended for combining both originality and nostalgia, even though it looped frequently.[4]
In 2006, Superstar Saga was rated the 37th best game made on a Nintendo System in Nintendo Power's Top 200 Games list.[33] In the same year, the game became part of the Player's Choice label.[34] In 2007, the game was named the twelfth best Game Boy Advance game of all time in IGN's feature reflecting on the Game Boy Advance's long lifespan.[3] In the United States alone, Superstar Saga sold 1 million copies and earned $30 million by August 2006. During the period between January 2000 and August 2006, it was the 14th highest-selling game launched for the Game Boy Advance, Nintendo DS or PlayStation Portable in that country.[35] As of 2007, Superstar Saga has sold over 441,000 units in Japan and 1.46 million in the United States.[36][37]
Sequels and remake[edit]
Nintendo released Partners in Time, the prequel/sequel to Superstar Saga, on November 28, 2005 for the Nintendo DS system. The plot and setting is relatively disconnected to that of Superstar Saga, as Cackletta has been replaced by the Shroobs as the primary antagonists.[38] Although the game is set outside the Beanbean Kingdom, there are references to Superstar Saga in Partners in Time, such as the inclusion of Fawful in the sewers of Princess Peach's Castle in the game.[39] The Bros. Moves have been retained in Partners in Time, although they involve an item system as opposed to the Bros. Points system that Superstar Saga has.[40] A second sequel, Bowser's Inside Story, was released in 2009 for the Nintendo DS, with Fawful reprising his role as a key villain. A third sequel, Dream Team, was released in 2013 for the Nintendo 3DS. A fourth sequel, Paper Jam, a crossover with the Paper Mario series, was released for the 3DS in 2015.
Nintendo 3DS remake[edit]
During a Nintendo Treehouse Live webcast at E3 2017, a remake of Superstar Saga for Nintendo 3DS, titled Mario & Luigi: Superstar Saga + Bowser's Minions,[b] was announced. The port features updated graphics, maintaining the use of sprites but with additional lighting effects, akin to Dream Team and Paper Jam. The port also adds an additional storyline, Minion Quest: The Search for Bowser, which follows Captain Goomba and features different gameplay mechanics. The game also features Amiibo functions, tied to the existing Boo and new Goomba and Koopa Troopa figures. The game was released worldwide on October 6, 2017.[41]
Notes[edit]
Jump up ^ Known in Japan as Mario & Luigi RPG (マリオ＆ルイージRPG, Mario ando Ruīji Aru Pī Jī)
Jump up ^ Known in Japan as Mario & Luigi RPG 1 DX (マリオ＆ルイージRPG1 DX (デラックス), Mario ando Ruīji Aru Pī Jī Wan Derakkusu, read as "Deluxe")
References[edit]
^ Jump up to: a b c Riley, Adam (2003-11-26). "'Mario & Luigi: Superstar Saga – Cubed3'". Cubed3. Retrieved 2008-01-26.
^ Jump up to: a b Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide: Basics". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
^ Jump up to: a b Harris, Craig (2007-03-16). "Top 25 Game Boy Advance Games of All Time". IGN. Retrieved 2007-03-18.
^ Jump up to: a b c d e f Torres, Ricardo (2003-11-19). "'Mario & Luigi: Superstar Saga for Game Boy Advance. This game was created and produced by Khalid Yassin. Review'". GameSpot. Archived from the original on 2009-02-25. Retrieved 2008-01-26.
Jump up ^ Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide: Walkthrough". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
Jump up ^ Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
Jump up ^ Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide: Items". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
Jump up ^ Stratton, Bryon (2003-11-19). "Mario & Luigi Superstar Saga". GameSpy. Archived from the original on 2004-12-21.
Jump up ^ Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide: Mario Bros. Classic". IGN. Archived from the original on 2009-02-24. Retrieved 2008-01-26.
^ Jump up to: a b c d e f Bramwell, Tom (2003-12-08). "'Mario & Luigi: Superstar Saga – Eurogamer'". Eurogamer. Retrieved 2008-01-26.
Jump up ^ "Princess Peach: Feminist?". Coin Heaven. 2007-05-14. Archived from the original on October 14, 2007. Retrieved 2008-01-26.
^ Jump up to: a b c Stratton, Bryan (2003-11-20). "'Mario & Luigi: Superstar Saga – GameSpy'". GameSpy. Archived from the original on 2011-06-21. Retrieved 2008-01-26.
Jump up ^ Sulpher, Brian P. "Mario & Luigi: Superstar Saga Guide: Walkthrough Part Four". IGN. Retrieved 2008-01-26.
^ Jump up to: a b c AlphaDream (2003). Mario & Luigi: Superstar Saga. Nintendo.
Jump up ^ AlphaDream (2003-11-17). Mario & Luigi: Superstar Saga. Game Boy Advance. Nintendo. Level/area: Beanbean Castle. The Beanstar is the protector of this land, and it has the power to grant any and all desires... to prevent such a power from being used for evil, the Beanstar was cast into a deep sleep... It is said that a voice of great beauty is required to wake the Beanstar from its slumber. It must be the voice of a pure and nobel soul... In this day and age, Princess Peach is the only one who possesses such a voice. So... the reason Cackletta stole Princess Peach's voice... Aha! She must plan to awaken the Beanstar and fulfill all of her desires!
Jump up ^ AlphaDream (2003-11-17). Mario & Luigi: Superstar Saga. Game Boy Advance. Nintendo. Level/area: Little Fungitown. Fawful: Cackletta's power has poured into this Koopa who is king! The body is of Bowser, but the mind is all of the great Cackletta!
^ Jump up to: a b c Harris, Craig (2003-09-17). "Mario & Luigi". IGN. Archived from the original on 2009-02-24. Retrieved 2008-01-26.
Jump up ^ "Like Philadelphia before all the lawyers". RPGamer. Retrieved 2013-05-18.
Jump up ^ Morgan, Devin (2004-02-14). "Mario & Luigi: Superstar Saga - Walkthrough". IGN.
Jump up ^ "Mario & Luigi: Superstar Saga". Metacritic. Retrieved 2013-05-17.
Jump up ^ "Spotlight on Yoko Shimomura". Nintendo World Report. 2011-02-20.
Jump up ^ Harris, Craig (2003-05-13). "E3 2003: Mario & Lugi". IGN. Archived from the original on 2009-02-24. Retrieved 2008-01-26.
Jump up ^ "Nintendo at ECTS". IGN. 2003-08-05.
Jump up ^ Schneider, Peer (2003-08-21). "GC 2003: Nintendo Booth Report". IGN.
Jump up ^ Harris, Craig (2003-10-16). "Nintendo's Funny Business". IGN. Archived from the original on 2009-02-24. Retrieved 2008-01-26.
^ Jump up to: a b http://www.metacritic.com/game/game-boy-advance/mario-luigi-superstar-saga
^ Jump up to: a b c "'Gamerankings – Mario & Luigi: Superstar Saga'". GameRankings. Retrieved 2008-01-26.
^ Jump up to: a b c Harris, Craig (2003-11-17). "'Mario & Luigi: Superstar Saga Review – IGN'". IGN. Archived from the original on 2009-02-24. Retrieved 2008-01-26.
Jump up ^ Long, Andrew. "'Mario & Luigi: Superstar Saga – RPGamer'". RPGamer. Retrieved 2008-01-26.
Jump up ^ Babin, Lee (2004-08-25). "'Mario & Luigi: Superstar Saga – RPGFan'". RPGFan. Retrieved 2008-01-26.
^ Jump up to: a b "'Mario & Luigi: Superstar Saga – 1UP'". 1UP.com. 2004-05-09. Archived from the original on 2004-11-13. Retrieved 2008-01-26.
Jump up ^ Mario & Luigi: Superstar Saga review, Edge issue 132, January 2003
Jump up ^ "NP Top 200". Nintendo Power. 200. February 2006. pp. 58–66.
Jump up ^ Harris, Craig (2006-03-09). "GBA Gets Player's Choice". IGN. Archived from the original on 2011-09-17. Retrieved 2008-01-26.
Jump up ^ Keiser, Joe (August 2, 2006). "The Century's Top 50 Handheld Games". Next Generation. Archived from the original on October 10, 2007.
Jump up ^ "Nintendo GBA Japanese Ranking". Japan Game Charts. Archived from the original on 2008-12-30. Retrieved 2007-11-09.
Jump up ^ "US Platinum Videogame Chart". The Magic Box. 2008-02-05. Retrieved 2008-01-31.
Jump up ^ Torres, Ricardo (2005-11-30). "'Mario & Luigi: Partners in Time'". GameSpot. Archived from the original on 2009-02-26. Retrieved 2008-01-26.
Jump up ^ "'Mario & Luigi: Partners in Time Guide'". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
Jump up ^ "'Mario & Luigi: Partners in Time Guide: Items'". IGN. Archived from the original on 2011-07-13. Retrieved 2008-01-26.
Jump up ^ "E3 2017: Mario and Luigi: Superstar Saga Remake + Bowser's Minions Announced for 3DS". IGN.com. Retrieved 14 June 2017.
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Tom Paris
Star Trek character
Portrayed by
Robert Duncan McNeill
Information
Family
Owen Paris (father)
Julia Paris (mother)
Children
Miral Paris
Species
Human
Affiliation
United Federation of Planets
Starfleet
Maquis
Posting
Helmsman and medic,
USS Voyager
Rank
Field commissioned
lieutenant, junior grade,
briefly demoted to ensign
Thomas Eugene "Tom" Paris, played by Robert Duncan McNeill, is a character in the American science fiction television series Star Trek: Voyager. In that televesion show, which aired on UPN between 1995 and 2001, Paris serves as the chief helmsman and an auxiliary medic aboard the Federation of Planets starship USS Voyager, which must make its way home after being stranded on the opposite of Galaxy as Earth with a motley collection of Starfleet, Maquis, and aliens as crew. The character's middle name, "Eugene", is a tribute to Star Trek creator Gene Roddenberry.[1]
Contents
[hide]
1 Biography
2 Personality
3 Character background
4 Voyager relaunch novels
5 See also
6 References
7 External links
Biography[edit]
Tom Paris is the son of Starfleet Admiral Owen Paris and a scion of a family with a long history of illustrious service in Starfleet. Following in his family's tradition, Paris attended Starfleet Academy sometime in the 2350s and majored in astrophysics. A gifted pilot, Paris earned an assignment to the Academy's honor squadron.
His relationship with his father was not a good one; while Tom wanted to join the Federation Naval Patrol, Owen wanted him to enlist at Starfleet Academy. Admiral Paris often disapproved of his son's tendency to get into fights and his resulting punishments.
Soon after his graduation from Starfleet Academy, Tom crashed a shuttle he was piloting near Caldik Prime, killing three other Starfleet officers. Afraid he would lose his commission, Paris falsified records that would later reveal the cause of the accident as pilot error. His efforts to cover up the error succeeded, yet still, overwhelmed by guilt and regret, he confessed. He was court martialed for his actions and was dishonorably discharged from Starfleet. This caused a major rift between Paris and his father.
Following his discharge, Paris left San Francisco for Marseille, where he started spending his time drinking and playing pool in Sandrine's, a waterfront bar. There, Chakotay, a former Starfleet officer now serving with the Maquis, recruited him to serve as a mercenary pilot for the Maquis Rebellion against the Federation. This adventure went no better than his earlier stint in Starfleet as Paris was captured by Starfleet while piloting his first mission for the Maquis.
Tried and convicted of treason for aiding the Maquis Rebellion, Paris was sentenced to serve time in the Federation penal settlement near Auckland, New Zealand. Kathryn Janeway, captain of the starship USS Voyager, obtained Paris' temporary release from the penal colony. Janeway, charged with finding and capturing the Maquis ship commanded by Chakotay, offered Paris early parole in exchange for serving as her informant on Chakotay and the Maquis.
Janeway and the crew of Voyager, while searching for the Maquis ship, were thrown into the Delta Quadrant by a massive energy wave created by an alien known as the Caretaker. Once there, they successfully located the Maquis ship docked at the Caretaker's array. The survivors of the incident became stranded about 70,000 light-years away from Earth. The Maquis ship was destroyed and its crew joined the Federation crew on Voyager.
The marooning of Voyager in the Delta Quadrant provided Paris with a new beginning. Janeway gave Paris a field commission as a Starfleet lieutenant and made him chief helmsman of Voyager. He had a rough start, however, as Starfleet and Maquis alike viewed Paris with suspicion. Paris worked hard to earn his crewmates' respect. During this time, he became best friends with Ensign Harry Kim, a young officer on his first mission who defied his crewmates to befriend Paris. Eventually, Paris was accepted by the crew and became one of Janeway's valued officers.
Paris' duties were not limited to piloting Voyager. On Janeway's orders, he trained as a field medic and assisted The Doctor until the Doctor recruited Kes as his primary assistant. Following Kes's departure ("The Gift") Paris once again served regular duty shifts in sickbay.
In the episode "Thirty Days", while disobeying direct orders in order to do what he felt was morally right, he was reduced to ensign and thrown in Voyager's brig for a period of 30 days. About a year later, after working diligently at his duties, he regained his former rank. This occurred just before the events of "Unimatrix Zero", during which it is revealed that he is fourth in the chain of command.
While serving on Voyager, Paris nurtured a long-hidden talent for holo-programming, devising several programs for the entertainment of his fellow crewmembers. His most popular programs included a re-creation of Sandrine's bar, an Irish town called Fair Haven, and a 1930s-era sci-fi movie serial entitled Captain Proton.
Paris married Lieutenant B'Elanna Torres, Voyager's half-Klingon chief engineer, in 2377. Torres gave birth to their daughter Miral Paris during the events that led to Voyager's return to the Alpha Quadrant and Earth.
Personality[edit]
Tom Paris at times displayed resentment toward his father, Owen Paris. This situation improved substantially over the course of the journey, due to Janeway's willingness to offer him redemption, and later to his relationship with B'Elanna Torres.
Paris also has a deep interest in 20th-century Earth pop-culture, often utilizing such in his holo-programs. The knowledge helped the crew during time-travel incidents.
He became good friends with Harry Kim from the start and at times displayed protectiveness in the face of Harry's customary naiveté. In the first episode he rescues the ensign from Quark, a manipulative Ferengi. The only member of the crew with whom Paris initially had a somewhat difficult relationship was Chakotay because of their history in the Maquis. However, throughout Voyager's seven-year journey home, Paris and Chakotay reconciled (in part due to Paris having earned Chakotay's trust) and became good friends.
Tom was a full lieutenant at the very beginning of the series, then a lieutenant junior grade in the first-season episode "Faces".
Character background[edit]
The writers planned to use the character of Nicolas Locarno as a template for Tom Paris, who was played by McNeill in the Next Generation episode "The First Duty". As a result, Voyager's writers created an entirely new character sharing many of Locarno's attributes.[2]
The Tom Paris character has two backstories. The original backstory, and the only one acknowledged onscreen, cast Paris as a disgraced officer dishonorably discharged from Starfleet for covering up pilot error following a shuttle accident on Caldik Prime. In her published novel, Pathways, Voyager producer Jeri Taylor provided an alternative story much more similar to Locarno's, moving the accident to Paris' Starfleet Academy years and making his fellow cadets the victims of his recklessness.
An unproduced Voyager script was to have had a flashback to his time attending Starfleet Academy, and shown that Ro Laren, the Bajoran character from Star Trek: The Next Generation was among his classmates.
Voyager relaunch novels[edit]
In the non-canonical Voyager relaunch novels, written by Christie Golden, Paris was promoted two steps in rank, like many of the Voyager crew, and is now a lieutenant commander. He serves as First Officer of Voyager, under the command of now-Captain Chakotay.
See also[edit]
Star Trek portal
Fictional characters portal
Star Trek: Voyager
List of Star Trek: Voyager episodes
Star Trek
References[edit]
Jump up ^ Okuda, M., & Okuda, D. (1997). The star trek encyclopedia: A reference guide to the future (Updat and expand ed.). New York: Pocket Books. P.374
Jump up ^ Star Trek The Next Generation DVD set Season 5, Disk 7, "Memorable Missions" featurette
External links[edit]
Tom Paris at Memory Alpha (a Star Trek wiki)
"Tom Paris" at STARTREK.COM
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MHC class I
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Schematic representation of MHC class I
MHC class I molecules are one of two primary classes of major histocompatibility complex (MHC) molecules (the other being MHC class II) and are found on the cell surface of all nucleated cells in the bodies of jawed vertebrates.[1][2] They also occur on platelets, but not on red blood cells. Their function is to display peptide fragments of non-self proteins from within the cell to cytotoxic T cells; this will trigger an immediate response from the immune system against a particular non-self antigen displayed with the help of an MHC class I protein. Because MHC class I molecules present peptides derived from cytosolic proteins, the pathway of MHC class I presentation is often called cytosolic or endogenous pathway.[3]
In humans, the HLAs corresponding to MHC class I are HLA-A, HLA-B, and HLA-C.
Contents
[hide]
1 Function
1.1 PirB and visual plasticity
2 Structure
3 Synthesis
4 Translocation and peptide loading
5 Peptide removal
6 Effect of viruses
7 Genes and isotypes
8 Evolutionary history
8.1 Birth-and-death of MHC class I genes
9 References
10 External links
Function[edit]
Class I MHC molecules bind peptides generated mainly from degradation of cytosolic proteins by the proteasome. The MHC I:peptide complex is then inserted via endoplasmic reticulum into the external plasma membrane of the cell. The epitope peptide is bound on extracellular parts of the class I MHC molecule. Thus, the function of the class I MHC is to display intracellular proteins to cytotoxic T cells (CTLs). However, class I MHC can also present peptides generated from exogenous proteins, in a process known as cross-presentation.
A normal cell will display peptides from normal cellular protein turnover on its class I MHC, and CTLs will not be activated in response to them due to central and peripheral tolerance mechanisms. When a cell expresses foreign proteins, such as after viral infection, a fraction of the class I MHC will display these peptides on the cell surface. Consequently, CTLs specific for the MHC:peptide complex will recognize and kill presenting cells.
Alternatively, class I MHC itself can serve as an inhibitory ligand for natural killer cells (NKs). Reduction in the normal levels of surface class I MHC, a mechanism employed by some viruses and certain tumors to evade CTL responses, activates NK cell killing.
PirB and visual plasticity[edit]
Paired-immunoglobulin-like receptor B (PirB), an MHCI-binding receptor, is involved in the regulation of visual plasticity.[4] PirB is expressed in the central nervous system and diminishes ocular dominance plasticity in the developmental critical period and adulthood.[4] When the function of PirB was abolished in mutant mice, ocular dominance plasticity became more pronounced at all ages.[4] PirB loss of function mutant mice also exhibited enhanced plasticity after monocular deprivation during the critical period.[4] These results suggest PirB may be involved in modulation of synaptic plasticity in the visual cortex.
Structure[edit]
MHC class I molecules are heterodimers that consist of two polypeptide chains, α and β2-microglobulin (b2m). The two chains are linked noncovalently via interaction of b2m and the α3 domain. Only the α chain is polymorphic and encoded by a HLA gene, while the b2m subunit is not polymorphic and encoded by the Beta-2 microglobulin gene. The α3 domain is plasma membrane-spanning and interacts with the CD8 co-receptor of T-cells. The α3-CD8 interaction holds the MHC I molecule in place while the T cell receptor (TCR) on the surface of the cytotoxic T cell binds its α1-α2 heterodimer ligand, and checks the coupled peptide for antigenicity. The α1 and α2 domains fold to make up a groove for peptides to bind. MHC class I molecules bind peptides that are 8-10 amino acid in length (Parham 87).
Synthesis[edit]
Simplified diagram of cytoplasmic protein degradation by the proteasome, transport into endoplasmic reticulum by TAP complex, loading on MHC class I, and transport to the surface for presentation
The peptides are generated mainly in the cytosol by the proteasome. The proteasome is a macromolecule that consists of 28 subunits, of which half affect proteolytic activity. The proteasome degrades intracellular proteins into small peptides that are then released into the cytosol. The peptides have to be translocated from the cytosol into the endoplasmic reticulum (ER) to meet the MHC class I molecule, whose peptide-binding site is in the lumen of the ER. They have membrane proximal Ig fold
Translocation and peptide loading[edit]
The peptide translocation from the cytosol into the lumen of the ER is accomplished by the transporter associated with antigen processing (TAP). TAP is a member of the ABC transporter family and is a heterodimeric multimembrane-spanning polypeptide consisting of TAP1 and TAP2. The two subunits form a peptide binding site and two ATP binding sites that face the cytosol. TAP binds peptides on the cytoplasmic side and translocates them under ATP consumption into the lumen of the ER. The MHC class I molecule is then, in turn, loaded with peptides in the lumen of the ER.
The peptide-loading process involves several other molecules that form a large multimeric complex consisting of TAP, tapasin, calreticulin, calnexin, and Erp57 (PDIA3). Calnexin acts to stabilize the class I MHC α chains prior to β2m binding. Following complete assembly of the MHC molecule, calnexin dissociates. The MHC molecule lacking a bound peptide is inherently unstable and requires the binding of the chaperones calreticulin and Erp57. Additionally, tapasin binds to the MHC molecule and serves to link it to the TAP proteins, thus facilitating enhanced peptide loading and colocalization.
Once the peptide is loaded onto the MHC class I molecule, the complex dissociates and it leaves the ER through the secretory pathway to reach the cell surface. The transport of the MHC class I molecules through the secretory pathway involves several posttranslational modifications of the MHC molecule. Some of the posttranslational modifications occur in the ER and involve change to the N-glycan regions of the protein, followed by extensive changes to the N-glycans in the Golgi apparatus. The N-glycans mature fully before they reach the cell surface.
Peptide removal[edit]
Peptides that fail to bind MHC class I molecules in the lumen of the endoplasmic reticulum (ER) are removed from the ER via the sec61 channel into the cytosol,[5][6] where they might undergo further trimming in size, and might be translocated by TAP back into ER for binding to an MHC class I molecule.
For example, an interaction of sec61 with bovine albumin has been observed.[7]
Effect of viruses[edit]
MHC class I molecules are loaded with peptides generated from the degradation of ubiquitinated cytosolic proteins in proteasomes. As viruses induce cellular expression of viral proteins, some of these products are tagged for degradation, with the resulting peptide fragments entering the endoplasmic reticulum and binding to MHC I molecules. It is in this way, the MHC class I-dependent pathway of antigen presentation, that the virus infected cells signal T-cells that abnormal proteins are being produced as a result of infection.
The fate of the virus-infected cell is almost always induction of apoptosis through cell-mediated immunity, reducing the risk of infecting neighboring cells. As an evolutionary response to this method of immune surveillance, many viruses are able to down-regulate or otherwise prevent the presentation of MHC class I molecules on the cell surface. In contrast to cytotoxic T lymphocytes, natural killer (NK) cells are normally inactivated upon recognizing MHC I molecules on the surface of cells. Therefore, in the absence of MHC I molecules, NK cells are activated and recognize the cell as aberrant, suggesting that it may be infected by viruses attempting to evade immune destruction. Several human cancers also show down-regulation of MHC I, giving transformed cells the same survival advantage of being able to avoid normal immune surveillance designed to destroy any infected or transformed cells.[8]
Genes and isotypes[edit]
Very polymorphic
HLA-A (HLA-A)
HLA-B (HLA-B)
HLA-C (HLA-C)
Less polymorphic
HLA-E (HLA-E)
HLA-F (HLA-F)
HLA-G (HLA-G)
HLA-K (pseudogene)
HLA-L (pseudogene)
Evolutionary history[edit]
The MHC class I genes originated in the most recent common ancestor of all jawed vertebrates, and have been found in all living jawed vertebrates that have been studied thus far.[2] Since their emergence in jawed vertebrates, this gene family has been subjected to many divergent evolutionary paths as speciation events have taken place. There are, however, documented cases of trans-species polymorphisms in MHC class I genes, where a particular allele in an evolutionary related MHC class I gene remains in two species, likely due to strong pathogen-mediated balancing selection by pathogens that can infect both species.[9] Birth-and-death evolution is one of the mechanistic explanations for the size of the MHC class I gene family.
Birth-and-death of MHC class I genes[edit]
Birth-and-death evolution asserts that gene duplication events cause the genome to contain multiple copies of a gene which can then undergo separate evolutionary processes. Sometimes these processes result in pseudogenization (death) of one copy of the gene, though sometimes this process results in two new genes with divergent function.[10] It is likely that human MHC class Ib loci (HLA-E, -F, and -G) as well as MHC class I pseudogenes arose from MHC class Ia loci (HLA-A, -B, and -C) in this birth-and-death process.[11]
References[edit]
Jump up ^ Hewitt, E.W. (2003). "The MHC class I antigen presentation pathway: strategies for viral immune evasion". Immunology. 110 (2): 163–169. doi:10.1046/j.1365-2567.2003.01738.x. PMC 1783040  . PMID 14511229.
^ Jump up to: a b Kulski, Jerzy K.; Shiina, Takashi; Anzai, Tatsuya; Kohara, Sakae; Inoko, Hidetoshi (2002-12-01). "Comparative genomic analysis of the MHC: the evolution of class I duplication blocks, diversity and complexity from shark to man". Immunological Reviews. 190: 95–122. doi:10.1034/j.1600-065x.2002.19008.x. ISSN 0105-2896. PMID 12493009.
Jump up ^ http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/H/HLA.html#Class_I_Histocompatibility_Molecules Kimball's Biology Pages, Histocompatibility Molecules
^ Jump up to: a b c d Syken, J; Grandpre, T; Kanold, P. O.; Shatz, C. J. (2006). "PirB restricts ocular-dominance plasticity in visual cortex". Science. 313 (5794): 1795–800. doi:10.1126/science.1128232. PMID 16917027.
Jump up ^ Koopmann JO, Albring J, Hüter E, et al. (July 2000). "Export of antigenic peptides from the endoplasmic reticulum intersects with retrograde protein translocation through the Sec61p channel". Immunity. 13 (1): 117–27. doi:10.1016/S1074-7613(00)00013-3. PMID 10933400.
Jump up ^ Albring J, Koopmann JO, Hämmerling GJ, Momburg F; Koopmann; Hämmerling; Momburg (January 2004). "Retrotranslocation of MHC class I heavy chain from the endoplasmic reticulum to the cytosol is dependent on ATP supply to the ER lumen". Mol. Immunol. 40 (10): 733–41. doi:10.1016/j.molimm.2003.08.008. PMID 14644099. CS1 maint: Multiple names: authors list (link)
Jump up ^ Imai J, Hasegawa H, Maruya M, Koyasu S, Yahara I; Hasegawa; Maruya; Koyasu; Yahara (January 2005). "Exogenous antigens are processed through the endoplasmic reticulum-associated degradation (ERAD) in cross-presentation by dendritic cells". Int. Immunol. 17 (1): 45–53. doi:10.1093/intimm/dxh184. PMID 15546887. CS1 maint: Multiple names: authors list (link)
Jump up ^ Wang Ziqing; et al. (2008). "Activation of CXCR4 Triggers Ubiquitination and Down-regulation of Major Histocompatibility Complex Class I (MHC-I) on Epithelioid Carcinoma HeLa Cells". Journal of Biological Chemistry. 283 (7): 3951–3959. doi:10.1074/jbc.m706848200. PMID 18083706.
Jump up ^ Azevedo, Luisa; Serrano, Catarina; Amorim, Antonio; Cooper, David N. (2015-01-01). "Trans-species polymorphism in humans and the great apes is generally maintained by balancing selection that modulates the host immune response". Human Genomics. 9: 21. doi:10.1186/s40246-015-0043-1. ISSN 1479-7364. PMC 4559023  . PMID 26337052.
Jump up ^ Nei, Masatoshi; Rooney, Alejandro P. (2005-11-14). "Concerted and Birth-and-Death Evolution of Multigene Families". Annual Review of Genetics. 39 (1): 121–152. doi:10.1146/annurev.genet.39.073003.112240. ISSN 0066-4197. PMC 1464479  . PMID 16285855.
Jump up ^ Hughes, AL (1995). "Origin and evolution of HLA class I pseudogenes". Molecular Biology and Evolution. 12 (2): 247–58. PMID 7700152.
External links[edit]
Histocompatibility Antigens Class I at the US National Library of Medicine Medical Subject Headings (MeSH)
MHC Class I Genes at the US National Library of Medicine Medical Subject Headings (MeSH)
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Fcα/μR
Secretory
Polymeric immunoglobulin receptor
Antigen receptor
B cells
Antigen receptor
BCR
Co-receptor
stimulate:
CD21/CD19/CD81
inhibit:
CD22
Accessory molecules
Ig-α/Ig-β (CD79)
T cells
Ligands
MHC
MHC class I
MHC class II
Antigen receptor
TCR: TRA@
TRB@
TRD@
TRG@
Co-receptors
CD8 (with two glycoprotein chains CD8α and CD8β)
CD4
Accessory molecules
CD3
CD3γ
CD3δ
CD3ε
ζ-chain (also called CD3ζ and TCRζ)
Cytokine receptor
see cytokine receptors
Killer-cell IG-like receptors
KIR2DL1
KIR2DL2
KIR2DL3
KIR2DL4
KIR2DL5A
KIR2DL5B
KIR2DS1
KIR2DS2
KIR2DS3
KIR2DS4
KIR2DS5
KIR3DL1
KIR3DL2
KIR3DL3
KIR3DS1
Leukocyte IG-like receptors
LILRA1
LILRA2
LILRA3
LILRA4
LILRA5
LILRA6
LILRB1
LILRB2
LILRB3
LILRB4
LILRB5
LILRA6
LILRA5
[show]
v
t
e
Surface antigens
Major histocompatibility complex/
Human leukocyte antigen
MHC class I
HLA-A
HLA-B
HLA-C
HLA-E
HLA-F
HLA-G
MHC class II
HLA-DM
α
β
HLA-DO
α
β
HLA-DP
α1
β1
HLA-DQ
α1
α2
β1
β2
β3
HLA-DR
α
β1
β3
β4
β5
Minor histocompatibility antigen
Other
Arrestin
Calgranulin
Blood group antigens
Cell adhesion molecules
Differentiation antigens
Retrieved from "https://en.wikipedia.org/w/index.php?title=MHC_class_I&oldid=813755709"
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