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Masters of the Universe (film)
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Masters of the Universe
Theatrical release poster
Directed by
Gary Goddard[1]
Produced by
Yoram Globus
Menahem Golan
Written by
David Odell
Starring
Dolph Lundgren
Frank Langella
Courteney Cox
James Tolkan
Christina Pickles
Meg Foster
Robert Duncan McNeill
Narrated by
Peter Brooks
Music by
Bill Conti
Cinematography
Hanania Baer
Edited by
Anne V. Coates
Production
company
Golan-Globus
Distributed by
Cannon Films[2]
Release date
August 7, 1987 (1987-08-07)
Running time
106 minutes[3]
Country
United States
Language
English
Budget
$22 million[4]
Box office
$17.3 million[5]
Masters of the Universe is a 1987 American science fantasy action film directed by Gary Goddard, produced by Yoram Globus and by Menahem Golan and written by David Odell. The film stars Dolph Lundgren, Frank Langella, Jon Cypher, Chelsea Field, Billy Barty, Courteney Cox and Robert Duncan McNeill. It is based on the Mattel toyline of the same name and brings the story of two teenagers who meet the mighty warrior He-Man, who arrived on Earth from planet Eternia and now goes on a mission to save the universe from the villianous Skeletor, his arch-nemesis.
Masters of the Universe was released theatrically in the United States on August 7, 1987. It was a critical and commercial failure, grossing $17 million worldwide against a budget of $22 million, but is now regarded as a cult film.[6][7][8]
Contents
[hide]
1 Plot
2 Cast
3 Production
3.1 Jack Kirby inspiration
4 Music
5 Release
5.1 Home media
6 Reception
7 Legacy
8 Reboot
9 References
10 External links
Plot[edit]
On the planet Eternia, at the center of the Universe, Skeletor's army seizes Castle Grayskull, scatters the remaining Eternian defenders, and captures the Sorceress of Grayskull, planning to add her power to his own by the next moonrise.
Skeletor's archenemy, the warrior He-Man, veteran soldier Man-At-Arms, and his daughter Teela rescue Gwildor from Skeletor's forces. Gwildor, a Thenorian locksmith, reveals that Skeletor has acquired his invention: a "Cosmic Key" that can open a portal to anywhere by utilizing sound keys. The device was stolen by Skeletor's second-in-command, Evil-Lyn, allowing Skeletor to breach Castle Grayskull.
With Gwildor's remaining prototype of the Key in hand, He-Man and his friends travel to the Castle. They attempt to free the Sorceress but are overwhelmed by Skeletor's army and forced to flee through Gwildor's hastily opened portal, transporting them to Earth. The Key is misplaced on their arrival and discovered by two New Jersey teenagers, orphaned high school girl Julie Winston and her boyfriend Kevin Corrigan. The pair attempt to figure out what the Key is and accidentally send a signal that allows Evil-Lyn to track the Key and send her henchmen – Saurod, Blade, Beast Man and Karg – to recover it.
Kevin, an aspiring musician, mistakes the Key for a synthesizer and takes it to a music store run by his friend Charlie. Karg's team arrives and chases Julie until He-Man comes across her and rescues her. Karg's team returns to Grayskull where, incensed by their failure, Skeletor kills Saurod and sends the others back to Earth, with a larger force under Evil-Lyn's command. Unable to find Julie, Kevin is taken to Julie's house by Lubic, a detective investigating the disturbance created by Karg's team. Suspecting the Key is stolen, Lubic confiscates it from Kevin and leaves. Immediately afterward, Evil-Lyn captures and interrogates Kevin for the Key's location with a mind control collar, before pursuing Lubic.
Julie and the Eternians release Kevin from the collar before they go after Lubic. They manage to get hold of it in Charlie's store, but Skeletor's forces catch up with them, and a pitched battle ensues. Evil-Lyn recovers the key and summons Skeletor to Earth. Skeletor's forces overwhelm the Eternians and Julie is mortally wounded by Skeletor's lightning blast, which simultaneously erases the memory storage of Gwildor's Key. He-Man surrenders to save his comrades, and is returned to Eternia as Skeletor's slave. Skeletor demands that He-Man kneel before him for all of Eternia to witness, before he is killed. He-Man refuses and is lashed by Blade's laser whip in an attempt to make him kneel. He-Man is still standing when the moon rises and Skeletor absorbs the powers of the universe. Declaring himself the Master of the Universe, Skeletor asserts his victory and continues to torture He-Man with energy blasts.
Back on Earth, Gwildor repairs the Cosmic Key, and Kevin recreates the tones necessary to create a gateway to Eternia. The group, including Lubic who attempts to arrest them, are transported to Castle Grayskull, where they begin battling Skeletor's forces. Resenting that Skeletor absorbed the power of the Universe without sharing it with her, Evil-Lyn deserts him along with the other henchmen. Skeletor accidentally frees He-Man who then reclaims the Sword of Grayskull. The pair battle until He-Man shatters Skeletor's staff, removing his new powers and restoring him to his normal state. He-Man offers mercy but Skeletor draws a concealed sword and attempts to kill He-Man; He-Man knocks Skeletor from the throne room into a towering pit below. The freed Sorceress heals Julie, and a portal is opened to send the Earthlings home. Hailed as a hero for his bravery, Lubic decides to remain on Eternia.
Julie awakens on the morning of her parents' deaths by plane crash. She prevents them from taking the ill-fated flight by taking their keys, and runs outside to find Kevin. Kevin confirms that their experiences were real, producing a souvenir from Eternia: a small blue sphere containing a scene of He-Man in front of Castle Grayskull with his sword raised above his head.
In a post-credits scene, Skeletor's head emerges from the water at the bottom of the pit, saying "I'll be back!".
Cast[edit]
Frank Langella in 2012. Langella chose to portray Skeletor because of his son's love of the character.
Dolph Lundgren – He-Man
Frank Langella – Skeletor
Courteney Cox – Julie Winston
James Tolkan – Detective Lubic
Christina Pickles – Sorceress
Meg Foster – Evil-Lyn
Chelsea Field – Teela
Jon Cypher – Man-At-Arms
Billy Barty – Gwildor
Robert Duncan McNeill – Kevin Corrigan
Anthony De Longis – Blade
Tony Carroll – Beast Man
Pons Maar – Saurod
Robert Towers – Karg
Peter Brooks – Narrator
Production[edit]
One of the original drafts from the script by David Odell (whose previous writing credits include Supergirl and The Dark Crystal) was reviewed in the third episode of the He-Man and She-Ra podcast, Masters Cast. The original draft included more time spent on Eternia and Snake Mountain, had Beast man in a speaking role, and even revealed that He-Man's mother was originally from Earth, as per the character Queen Marlena from the Filmation animated series He-Man and the Masters of the Universe, thus linking the two planets.[9] Describing her character, Foster said that Evil-Lyn is not villainous, "she is just doing her job and she knows how to get results, even if it means being harsh." Langella agreed, calling Evil-Lyn a female more dedicated to Skeletor's cause than any man; she is obsessive around Skeletor because she is slightly lovelorn.[10] The filmmakers considered having Foster wear eye-lenses to mask her naturally pale-blue eyes, but decided that her natural eyes fit the character better. However, they did augment Foster's chest, fitting cleavage into the character's costume. Foster wanted the character to have a large hairstyle, rather than the short style featured in the film.[11]
When offered the role, Langella said that he "didn’t even blink ... I couldn't wait to play him." Langella cited his then-four-year-old son's love of Skeletor while running around his house yelling He-Man's battle cry "I have the power!", as the reason he chose to play He-Man's archenemy.[12]
Jack Kirby inspiration[edit]
Comic book writer/artist John Byrne compared the film to Jack Kirby's comic book metaseries Fourth World, stating in Comic Shop News #497:
The best New Gods movie, IMHO, is Masters of the Universe. I even corresponded with the director, who told me this was his intent, and that he had tried to get [Jack] Kirby to do the production designs, but the studio nixed it.
Check it out. It requires some bending and an occasional sex change (Metron becomes an ugly dwarf, The Highfather becomes the Sorceress), but it's an amazingly close analog, otherwise. And Frank Langella's Skeletor is a dandy Darkseid!
Director Gary Goddard clarified this in a letter appearing in John Byrne's Next Men #26, in which he stated:
As the director of Masters of the Universe, it was a pleasure to see that someone got it. Your comparison of the film to Kirby’s New Gods was not far off. In fact, the storyline was greatly inspired by the classic Fantastic Four/Doctor Doom epics, The New Gods and a bit of Thor thrown in here and there. I intended the film to be a "motion picture comic book," though it was a tough proposition to sell to the studio at the time. 'Comics are just for kids,' they thought. They would not allow me to hire Jack Kirby who I desperately wanted to be the conceptual artist for the picture…
I grew up with Kirby's comics (I've still got all my Marvels from the first issue of Fantastic Four and Spider-Man through the time Kirby left) and I had great pleasure meeting him when he first moved to California. Since that time I enjoyed the friendship of Jack and Roz and was lucky enough to spend many hours with Jack, hearing how he created this character and that one, why a villain has to be even more powerful than a hero, and on and on. Jack was a great communicator, and listening to him was always an education. You might be interested to know that I tried to dedicate Masters of Universe to Jack Kirby in the closing credits, but the studio took the credit out.
Brian Cronin, author of the "Comic Book Urban Legends Revealed" column, concludes that "the film itself was not intended to be literally a reworked Fourth World, although the intent WAS to make the film a tribute to Jack Kirby — just a tribute to ALL of his work, not just the Fourth World."[13]
Music[edit]
The musical score of Masters of the Universe was composed by Bill Conti. It was recorded by several European orchestras, chiefly the Graunke Orchestra of Munich (the only one to be credited on the soundtrack album) and conducted by a number of conductors, chiefly Bruce Miller and Harry Rabinowitz (Rabinowitz received sole credit). Conti did not conduct his score because it could not be recorded in the United States as "there was a musicians strike or something like that... So it went to various places." He and the score mixer Dan Wallin assembled the score from the various recorded takes, because there were problems with the orchestral performances ("We didn't have anything that went from beginning to end without a problem").[14]
The soundtrack album was released on record, cassette and compact disc by Varese Sarabande in 1987; it was subsequently issued in an expanded version by Edel. In 2008, La-La Land Records released a two-disc edition with the complete score and the original album presentation; in 2012, Intrada Records issued the complete score (the entirely of disc one and tracks 1–5 on disc two) on one disc.
Release[edit]
Prior to releasing it, the Cannon Group touted Masters of the Universe as the Star Wars of the 1980s. Despite releasing alongside the height of the success of the toy line, animated series, and related merchandise, Masters of the Universe began as the third-highest-grossing film of the weekend in North America on August 7, 1987, earning $4,883,168, behind Stakeout ($5,170,403) and The Living Daylights ($7,706,230). The film quickly left the charts altogether with a North American gross of $17,336,370.[15][2][5][16]
Home media[edit]
Masters of the Universe was released on DVD October 23, 2001. A 25th Anniversary Edition Blu-ray Disc was released by Warner Home Video on October 2, 2012.[17]
Reception[edit]
Masters of the Universe received negative reviews from critics and holds a 17% rating on Rotten Tomatoes based on 23 reviews.[18] Variety called it a "Conan-Star Wars hybrid ripoff" that is "a colossal bore".[19] Walter Goodman of The New York Times wrote, "If you liked the toy, you'll love the movie."[20] Michael Wilmington of the Los Angeles Times called it "a misfiring, underdone epic".[21] Johanna Steinmetz of the Chicago Tribune wrote that the film, while predictable and derivative, entertains audiences through its side plots set on Earth.[22]
In a retrospective review, Glenn Heath, Jr. of Slant Magazine called it a "jarring mix of corny screwball comedy and choppy action heroics".[23] Chris Eggertsen of HitFix, in an article identifying the film's campy, positive qualities, called it "an objectively bad film with a big heart".[24] Joshua Winning of Digital Spy wrote, "...beloved of '80s kids but scorned by critics, it's a high camp oddity that we should celebrate on its own terms".[4]
Billy Barty received a Golden Raspberry Award nomination for Worst Supporting Actor for his performance in the film, but 'lost' to David Mendenhall for Over the Top.
Legacy[edit]
The commercial failure of Masters of the Universe, among other films such as Superman IV: The Quest for Peace and Lifeforce, contributed to the eventual closure of Cannon Films.[11] Cannon Films had intended to create a Masters of the Universe sequel, indicated by the end credits with a surprise reveal that Skeletor survives his fall. The sequel titled "Masters of the Universe 2: Cyborg" was written and followed He-Man, who returned to Earth to battle Skeletor, who has left Earth as a post-apocalyptic wasteland and the film was to feature Trap Jaw and She-Ra. Pro surfer Laird Hamilton was originally to replace Dolph Lundgren as He-Man and the only aspect known about the sequel's screenplay was that He-Man would have returned to Earth and was disguised as a professional quarterback.[25] With a low budget of $4.5 million, the sequel was to be directed by Albert Pyun, consecutively with the aborted Spider-Man movie. The project was abandoned when Cannon would not pay Mattel's fees. The production instead utilized the already-made costumes and sets for the low-budget sci-fi film Cyborg.[26][27][28]
Masters of the Universe was Lundgren's first leading role in a feature film following his success in Rocky IV, and he later labeled it as his least favorite film role.[10] Conversely, Langella considers playing Skeletor to be one of his favorite roles.[12]
Reboot[edit]
A new He-Man film directed by John Woo was reportedly being developed in 2007, but despite many rumors circulating the Internet regarding the film's production status and casting, the project was never officially green-lit. The film rights to He-Man have reportedly since reverted to Mattel.[29]
In September 2009, Sony took over the rights from Warner Bros. to produce the live action adaptation after Mattel and producer Joel Silver, who was previously involved with a potential film, couldn't agree on creative direction for the film.[30] Sony and Escape Artists' Todd Black, Jason Blumenthal, and Steve Tisch will now start developing the project from scratch for Columbia Pictures. In April 2010, Sony hired screenwriters Mike Finch and Alex Litvak to draft a new script.[31] Warner announced that John Stevenson, director of Kung Fu Panda will direct the upcoming feature. On May 12, 2009, it was announced that the scripting duties had been handed to Evan Daugherty, with Stevenson still attached to direct.[32]
In late 2012, Deadline reported that Jon M. Chu was in talks to direct the film.[33] Original He-Man actor Lundgren did an interview with IGN about a possible role in the film as King Randor.[34] October 12, 2012, Richard Wenk is hired to rewrite the script for the film.[35] On March 28, 2013, Chu spoken that the film is still early is in the experimentation and also it won't be campy but an origin story.[36][37] On October 7, The Hollywood Reporter reported that Terry Rossio would script and that Black, Blumenthal, and Tisch would produce the film and it will be set on Eternia; the site also reported that Chu will not direct the film.[38]
On January 10, 2014, Schmoes Know reported that Joe Cornish (Attack the Block), Rian Johnson (Looper), Andrés Muschietti (Mama), Kirk DeMicco and Chris Sanders (The Croods), and Phil Lord and Chris Miller (The Lego Movie) were named as frontrunners to direct the film.[39] On February 26, it was reported that directors Mike Cahill, Jeff Wadlow, Harald Zwart, and Chris McKay are on the short list to direct.[40][41] On April 9, Schmoes Know reported that Wadlow will direct the film, but The Hollywood Reporter announced that he's rewriting the script of the film.[42][43]
On August 19, 2015, it was announced that Chris Yost had been hired to write the film.[44]
On January 22, 2016, Deadline reports that McG will direct the film and will also oversee a rewrite of the latest script by Alex Litvak and Mike Finch, while Escape Artists’ Todd Black, Jason Blumenthal and Steve Tisch are on board as producers.[45]
On June 24, 2016, Kellan Lutz tweeted that he has a meeting with both McG and Mary Viola about having the role of He-Man.[46][47]
On April 27, 2017, it was announced that the film will be released on December 18, 2019. At the same time, McG left the film and David S. Goyer has been hired to rewrite the script.[48]
References[edit]
Jump up ^ "`Masters` A Lesson In More Thrills For Less". Chicago Tribune. 1987-08-13. Retrieved 2010-08-28.
^ Jump up to: a b Arts, ' (1987-08-11). "Weekend Box Office". The Los Angeles Times. Retrieved 2010-11-17.
Jump up ^ "Masters of the Universe (1987)". bbfc.co.uk. British Board of Film Classification. September 16, 1987. Archived from the original on July 12, 2014. Retrieved July 12, 2014.
^ Jump up to: a b Winning, Joshua (2015-05-03). "In Defense Of... Masters of the Universe, Dolph Lundgren's fantasy flop". Digital Spy. Retrieved 2015-09-03.
^ Jump up to: a b "Masters of the Universe". Box Office Mojo. Amazon.com. July 12, 2014. Archived from the original on April 6, 2014. Retrieved July 12, 2014.
Jump up ^ Kroll, Justin (2015-08-19). "'Thor: Ragnarok' Scribe to Pen 'Masters of the Universe' for Sony". Variety. Retrieved 2016-04-01. spawned a movie... that subsequently became a cult hit
Jump up ^ Miller, Leon (2017-09-03). "Masters Of The Universe: 15 Shocking Things You Didn't Know About The Movie". Screen Rant. Retrieved 2017-09-10. it’s since gone on to become a cult classic
Jump up ^ Colburnon, Randall (2017-04-28). "Believe it or not, a Masters of the Universe reboot is slated for a 2019 release". Consequence of Sound. Retrieved 2017-09-10. the colossal flop turned campy cult classic that was 1987’s Masters of the Universe
Jump up ^ "Masters Cast – Episode 3". Masters Cast. Retrieved 2012-07-23.
^ Jump up to: a b SFX 2013, p. 84.
^ Jump up to: a b SFX 2013, p. 85.
^ Jump up to: a b "Frank Langella calls Skeletor "One of my Very Favourite Parts"". ifc.com. IFC. August 7, 2012. Archived from the original on May 10, 2013. Retrieved July 13, 2014.
Jump up ^ "Comic Book Urban Legends Revealed #75". Goodcomics.comicbookresources.com. Retrieved 2010-04-28.
Jump up ^ "Bill Conti's Music of the Universe," pg. 8, liner notes, Masters of the Universe: Complete Original Motion Picture Soundtrack, La-La Land Records LLLCD 1071.
Jump up ^ SFX 2013, p. 82.
Jump up ^ "August 7–9, 1987". Box Office Mojo. Amazon.com. July 12, 2014. Archived from the original on April 3, 2014. Retrieved July 12, 2014.
Jump up ^ "Masters Of The Universe: 25th Anniversary (BD) | WBshop.com | Warner Bros". Wbshop.com. Retrieved 2012-07-24.
Jump up ^ "Masters of the Universe". Rotten Tomatoes. Flixster. Retrieved September 7, 2014.
Jump up ^ "Review: 'Masters of the Universe'". Variety. 1987. Retrieved 2015-09-03.
Jump up ^ Goodman, Walter (1987-08-08). "Masters of the Universe (1987)". The New York Times. Retrieved 2015-09-03.
Jump up ^ Wilmington, Michael (1987-08-12). "Movie Review : 'Masters Of The Universe' Misfires". Los Angeles Times. Retrieved 2015-09-03.
Jump up ^ Steinmetz, Johanna (1987-08-12). "Surprise! 'Masters' Isn't Bad". Chicago Tribune. Retrieved 2015-09-03.
Jump up ^ Heath, Glenn (2012-07-29). "Summer of '87: Masters of the Universe: He-Man, Voice of Reason". Slant Magazine. Retrieved 2015-09-03.
Jump up ^ Eggertson, Chris (2015-08-25). "'Masters of the Universe' wasn't all bad: 13 things to actually admire about the 1987 flop". HitFix. Retrieved 2015-09-03.
Jump up ^ Lambie, Ryan (July 6, 2016). "How a He-Man sequel and a Spider-Man movie became Cyborg". Den of Geek.
Jump up ^ Brian Cronin (2013-01-30). "Movie Legends Revealed: He-Man & Spider-Man Films Became Cyborg? – Spinoff Online – TV, Film, and Entertainment News Daily". Spinoff.comicbookresources.com. Retrieved 2013-09-28.
Jump up ^ "Masters of the Universe DVD (1987)". BBC. Retrieved 2010-08-08.
Jump up ^ "Sequel Baiting Endings That Didn't Work". Empire. Retrieved 2012-06-11.
Jump up ^ Clint Morris, Fox2000 no longer has the Power of Grayskull, Moviehole, November 30, 2006.
Jump up ^ "Barbie's a living doll at Universal – Entertainment News, Film News, Media". Variety. 2009-09-23. Retrieved 2010-04-28.
Jump up ^ drees, Rich. "MASTERS OF THE UNIVERSE Gets Screenwriters". filmbuffonline.com. Retrieved 2010-04-12.
Jump up ^ Kit, Borys. "'Grayskull' lands new writer". Hollywoodreporter.com. Archived from the original on May 16, 2009. Retrieved 2009-05-14.
Jump up ^ Fleming, Mike. "Jon M. Chu In 'Masters Of The Universe' Talks". Deadline Hollywood. Retrieved 2012-07-30.
Jump up ^ Fleming, Mike. "He-Man and the Masters of the Universe – Lundgren on He-Man Reboot". IGN. Retrieved 2012-08-03.
Jump up ^ Sneider, Jeff (2012-10-12). "Wenk to pen 'Masters of the Universe'". Variety. Retrieved 2012-10-12.
Jump up ^ "He-Man and the Masters of the Universe - Jon M. Chu Production Update - IGN Video".
Jump up ^ "TR Interview: Jon M. Chu Talks G.I. Joe: Retaliation...and Masters of the Universe! - The Robot's Voice". March 28, 2013.
Jump up ^ "'Lone Ranger' Co-Writer Tapped to Write 'Masters of the Universe' (Exclusive)".
Jump up ^ The Phantom (2014-01-10). "He-Man and the Masters of the Universe Close to Happening; Looking at New Directors!". Schmoes Know. Retrieved 2014-01-10.
Jump up ^ Kroll, Justin (February 26, 2014). "BTW, Mike Cahill and Jeff Wadlow are on the short list for MASTERS OF THE UNIVERSE".
Jump up ^ "He-Man Film 'Masters of the Universe' Director Short-List Includes 'Lego Movie' Editor (Exclusive)". The Wrap. 2014-02-26. Retrieved 2014-02-26.
Jump up ^ "A PHANTOM EXCLUSIVE: Jeff Wadlow Will Direct MASTERS OF THE UNIVERSE!". Schmoes Know. April 9, 2014. Retrieved 2014-04-09.
Jump up ^ "Jeff Wadlow to Rewrite 'Masters of the Universe'". The Hollywood Reporter. April 10, 2014. Retrieved 2014-04-10.
Jump up ^ "Christopher Yost to Write 'Masters of the Universe' Film". Comicbookresources.com. August 19, 2015. Retrieved August 19, 2015.
Jump up ^ Fleming, Jr, Mike (January 22, 2016). "McG Orbiting 'Masters Of The Universe' At Sony". Deadline.
Jump up ^ Lutz, Kellan. "Hey @HeManTheMovie and all MOTU fans, Had an amazing meeting with @McGsWonderland and @ItsMaryViola talking MOTU! ITS IN GREAT HANDS!". line feed character in |title= at position 38 (help)
Jump up ^ Burlingame, Russ (June 24, 2016). "Kellan Lutz Met With McG About Masters Of The Universe Movie". ComicBook.com.
Jump up ^ Chavez, Danette. "Masters Of The Universe gets a release date, but loses its director".
"Masters of the Universe". SFX. Future plc (240): 82–85. November 2013.
External links[edit]
Wikiquote has quotations related to: Masters of the Universe
Masters of the Universe on IMDb
Masters of the Universe at Box Office Mojo
Masters of the Universe at Rotten Tomatoes
Masters of the Universe on YouTube
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Chromosomal crossover
From Wikipedia, the free encyclopedia
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Crossing over occurs between prophase 1 and metaphase 1 and is the process where homologous chromosomes pair up with each other and exchange different segments of their genetic material to form recombinant chromosomes. It can also happen during mitotic division,[1] which may result in loss of heterozygosity. Crossing over is essential for the normal segregation of chromosomes during meiosis.[citation needed] Crossing over also accounts for genetic variation, because due to the swapping of genetic material during crossing over, the chromatids held together by the centromere are no longer identical. So, when the chromosomes go on to meiosis II and separate, some of the daughter cells receive daughter chromosomes with recombined alleles. Due to this genetic recombination, the offspring have a different set of alleles and genes than their parents do. In the diagram, genes B and b are crossed over with each other, making the resulting recombinants after meiosis Ab, AB, ab, and aB.
Thomas Hunt Morgan's illustration of crossing over (1916)
A double crossing over
Chromosomal crossover (or crossing over) is the exchange of genetic material between homologous chromosomes that results in recombinant chromosomes during sexual reproduction. It is one of the final phases of genetic recombination, which occurs in the pachytene stage of prophase I of meiosis during a process called synapsis. Synapsis begins before the synaptonemal complex develops and is not completed until near the end of prophase I. Crossover usually occurs when matching regions on matching chromosomes break and then reconnect to the other chromosome.
Crossing over was described, in theory, by Thomas Hunt Morgan. He relied on the discovery of the Belgian Professor Frans Alfons Janssens of the University of Leuven who described the phenomenon in 1909 and had called it "chiasmatypie". The term chiasma is linked if not identical to chromosomal crossover. Morgan immediately saw the great importance of Janssens' cytological interpretation of chiasmata to the experimental results of his research on the heredity of Drosophila. The physical basis of crossing over was first demonstrated by Harriet Creighton and Barbara McClintock in 1931.[2]
The linked frequency of crossing over between two gene loci (markers) is the crossing-over value . For fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant and the same is then true for the crossing-over value which is used in the production of genetic maps.[3][4]
Contents
[hide]
1 Origins
1.1 DNA repair theory
1.2 Links to bacterial transformation
2 Chemistry
2.1 MSH4 and MSH5
2.2 Chiasma
3 Consequences
4 Non-homologous crossover
5 See also
6 References
Origins[edit]
There are two popular and overlapping theories that explain the origins of crossing-over, coming from the different theories on the origin of meiosis. The first theory rests upon the idea that meiosis evolved as another method of DNA repair, and thus crossing-over is a novel way to replace possibly damaged sections of DNA.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating diversity.[6]
DNA repair theory[edit]
Crossing over and DNA repair are very similar processes, which utilize many of the same protein complexes.[5][7][8] Recombinases and primases lay a foundation of nucleotides along the DNA sequence. One such particular protein complex that is conserved between processes is RAD51, a well conserved recombinase protein that has been shown to be crucial in DNA repair as well as cross over.[9] Several other genes in D. melanogaster have been linked as well to both processes, by showing that mutants at these specific loci cannot undergo DNA repair or crossing over. Such genes include mei-41, mei-9, hdm, spnA, and brca2.[5] This large group of conserved genes between processes supports the theory of a close evolutionary relationship. Furthermore, DNA repair and crossover have been found to favor similar regions on chromosomes. In an experiment using radiation hybrid mapping on wheat’s (Triticum aestivum L.) 3B chromosome, crossing over and DNA repair were found to occur predominantly in the same regions.[10] Furthermore, crossing over has been correlated to occur in response to stressful, and likely DNA damaging, conditions [11][12]
Links to bacterial transformation[edit]
The process of bacterial transformation also shares many similarities with chromosomal cross over, particularly in the formation of overhangs on the sides of the broken DNA strand, allowing for the annealing of a new strand. Bacterial transformation itself has been linked to DNA repair many times.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating genetic diversity.[6] .[13] Thus, this evidence suggests that it is a question of whether cross over is linked to DNA repair or bacterial transformation, as the two do not appear to be mutually exclusive. It is likely that crossing over may have evolved from bacterial transformation, which in turn developed from DNA repair, thus explaining the links between all three processes.
Chemistry[edit]
A current model of meiotic recombination, initiated by a double-strand break or gap, followed by pairing with an homologous chromosome and strand invasion to initiate the recombinational repair process. Repair of the gap can lead to crossover (CO) or non-crossover (NCO) of the flanking regions. CO recombination is thought to occur by the Double Holliday Junction (DHJ) model, illustrated on the right, above. NCO recombinants are thought to occur primarily by the Synthesis Dependent Strand Annealing (SDSA) model, illustrated on the left, above. Most recombination events appear to be the SDSA type.
Meiotic recombination may be initiated by double-stranded breaks that are introduced into the DNA by exposure to DNA damaging agents[5] or the Spo11 protein.[14] One or more exonucleases then digest the 5’ ends generated by the double-stranded breaks to produce 3’ single-stranded DNA tails (see diagram). The meiosis-specific recombinase Dmc1 and the general recombinase Rad51 coat the single-stranded DNA to form nucleoprotein filaments.[15] The recombinases catalyze invasion of the opposite chromatid by the single-stranded DNA from one end of the break. Next, the 3’ end of the invading DNA primes DNA synthesis, causing displacement of the complementary strand, which subsequently anneals to the single-stranded DNA generated from the other end of the initial double-stranded break. The structure that results is a cross-strand exchange, also known as a Holliday junction. The contact between two chromatids that will soon undergo crossing-over is known as a chiasma. The Holliday junction is a tetrahedral structure which can be 'pulled' by other recombinases, moving it along the four-stranded structure.
Holliday Junction
Molecular structure of a Holliday junction.
Molecular structure of a Holliday junction. From PDB: 3CRX.
MSH4 and MSH5[edit]
The MSH4 and MSH5 proteins form a hetero-oligomeric structure (heterodimer) in yeast and humans.[16][17][18] In the yeast Saccharomyces cerevisiae MSH4 and MSH5 act specifically to facilitate crossovers between homologous chromosomes during meiosis.[16] The MSH4/MSH5 complex binds and stabilizes double Holliday junctions and promotes their resolution into crossover products. An MSH4 hypomorphic (partially functional) mutant of S. cerevisiae showed a 30% genome wide reduction in crossover numbers, and a large number of meioses with non exchange chromosomes.[19] Nevertheless, this mutant gave rise to spore viability patterns suggesting that segregation of non-exchange chromosomes occurred efficiently. Thus in S. cerevisiae proper segregation apparently does not entirely depend on crossovers between homologous pairs.
Chiasma[edit]
The grasshopper Melanoplus femur-rubrum was exposed to an acute dose of X-rays during each individual stage of meiosis, and chiasma frequency was measured.[20] Irradiation during the leptotene-zygotene stages of meiosis (that is, prior to the pachytene period in which crossover recombination occurs) was found to increase subsequent chiasma frequency. Similarly, in the grasshopper Chorthippus brunneus, exposure to X-irradiation during the zygotene-early pachytene stages caused a significant increase in mean cell chiasma frequency.[21] Chiasma frequency was scored at the later diplotene-diakinesis stages of meiosis. These results suggest that X-rays induce DNA damages that are repaired by a crossover pathway leading to chiasma formation.
Consequences[edit]
The difference between gene conversion and chromosomal crossover.
In most eukaryotes, a cell carries two versions of each gene, each referred to as an allele. Each parent passes on one allele to each offspring. An individual gamete inherits a complete haploid complement of alleles on chromosomes that are independently selected from each pair of chromatids lined up on the metaphase plate. Without recombination, all alleles for those genes linked together on the same chromosome would be inherited together. Meiotic recombination allows a more independent segregation between the two alleles that occupy the positions of single genes, as recombination shuffles the allele content between homologous chromosomes.
Recombination results in a new arrangement of maternal and paternal alleles on the same chromosome. Although the same genes appear in the same order, some alleles are different. In this way, it is theoretically possible to have any combination of parental alleles in an offspring, and the fact that two alleles appear together in one offspring does not have any influence on the statistical probability that another offspring will have the same combination. This principle of "independent assortment" of genes is fundamental to genetic inheritance.[22] However, the frequency of recombination is actually not the same for all gene combinations. This leads to the notion of "genetic distance", which is a measure of recombination frequency averaged over a (suitably large) sample of pedigrees. Loosely speaking, one may say that this is because recombination is greatly influenced by the proximity of one gene to another. If two genes are located close together on a chromosome, the likelihood that a recombination event will separate these two genes is less than if they were farther apart. Genetic linkage describes the tendency of genes to be inherited together as a result of their location on the same chromosome. Linkage disequilibrium describes a situation in which some combinations of genes or genetic markers occur more or less frequently in a population than would be expected from their distances apart. This concept is applied when searching for a gene that may cause a particular disease. This is done by comparing the occurrence of a specific DNA sequence with the appearance of a disease. When a high correlation between the two is found, it is likely that the appropriate gene sequence is really closer.[23]
Non-homologous crossover[edit]
Crossovers typically occur between homologous regions of matching chromosomes, but similarities in sequence and other factors can result in mismatched alignments. Most DNA is composed of base pair sequences repeated very large numbers of times.[24] These repetitious segments, often referred to as satellites, are fairly homogenous among a species.[24] During DNA replication, each strand of DNA is used as a template for the creation of new strands using a partially-conserved mechanism; proper functioning of this process results in two identical, paired chromosomes, often called sisters. Sister chromatid crossover events are known to occur at a rate of several crossover events per cell per division in eukaryotes. [24] Most of these events involve an exchange of equal amounts of genetic information, but unequal exchanges may occur due to sequence mismatch. These are referred to by a variety of names, including non-homologous crossover, unequal crossover, and unbalanced recombination, and result in an insertion or deletion of genetic information into the chromosome. While rare compared to homologous crossover events, these mutations are drastic, affecting many loci at the same time. They are considered the main driver behind the generation of gene duplications and are a general source of mutation within the genome.[25]
Many proteins are implicated in DSB repair; this figure shows a probable, simplified mechanism.
The specific causes of non-homologous crossover events are unknown, but several influential factors are known to increase the likelihood of an unequal crossover. One common vector leading to unbalanced recombination is the repair of double-strand breaks (DSBs).[26] DSBs are often repaired using non-homologous end joining, a process which involves invasion of a template strand by the DSB strand (see figure below). Nearby homologous regions of the template strand are often used for repair, which can give rise to either insertions or deletions in the genome if a non-homologous but complementary part of the template strand is used.[26] Sequence similarity is a major player in crossover – crossover events are more likely to occur in long regions of close identity on a gene.[27] This means that any section of the genome with long sections of repetitive DNA is prone to crossover events.
The presence of transposable elements is another influential element of non-homologous crossover. Repetitive regions of code characterize transposable elements; complementary but non-homologous regions are ubiquitous within transposons. Because chromosomal regions composed of transposons have large quantities of identical, repetitious code in a condensed space, it is thought that transposon regions undergoing a crossover event are more prone to erroneous complementary match-up;[28] that is to say, a section of a chromosome containing a lot of identical sequences, should it undergo a crossover event, is less certain to match up with a perfectly homologous section of complementary code and more prone to binding with a section of code on a slightly different part of the chromosome. This results in unbalanced recombination, as genetic information may be either inserted or deleted into the new chromosome, depending on where the recombination occurred.
While the motivating factors behind unequal recombination remain obscure, elements of the physical mechanism have been elucidated. Mismatch repair (MMR) proteins, for instance, are a well-known regulatory family of proteins, responsible for regulating mismatched sequences of DNA during replication and escape regulation.[29] The operative goal of MMRs is the restoration of the parental genotype. One class of MMR in particular, MutSβ, is known to initiate the correction of insertion-deletion mismatches of up to 16 nucleotides.[29] Little is known about the excision process in eukaryotes, but E. coli excisions involve the cleaving of a nick on either the 5’ or 3’ strand, after which DNA helicase and DNA polymerase III bind and generate single-stranded proteins, which are digested by exonucleases and attached to the strand by ligase.[29] Multiple MMR pathways have been implicated in the maintenance of complex organism genome stability, and any of many possible malfunctions in the MMR pathway result in DNA editing and correction errors.[30] Therefore, while it is not certain precisely what mechanisms lead to errors of non-homologous crossover, it is extremely likely that the MMR pathway is involved.
See also[edit]
Unequal crossing over
Coefficient of coincidence
Genetic distance
Independent assortment
Mitotic crossover
Recombinant frequency
References[edit]
Jump up ^ Griffiths, AJF; Gelbart, WM; Miller, JH; et al. (1999). "Modern Genetic Analysis: Mitotic Crossing-Over". New York: W. H. Freeman.
Jump up ^ Creighton H, McClintock B (1931). "A Correlation of Cytological and Genetical Crossing-Over in Zea Mays". Proc Natl Acad Sci USA. 17 (8): 492–7. PMC 1076098  . PMID 16587654. doi:10.1073/pnas.17.8.492. (Original paper)
Jump up ^ Rieger R. Michaelis A., Green M. M. (1976). Glossary of genetics and cytogenetics: Classical and molecular. Heidelberg - New York: Springer-Verlag. ISBN 3-540-07668-9. CS1 maint: Uses authors parameter (link)
Jump up ^ King R. C., Stransfield W. D. (1998): Dictionary of genetics. Oxford University Press, New York, Oxford, ISBN 0-19-50944-1-7; ISBN 0-19-509442-5.
^ Jump up to: a b c d e Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 in DNA Repair. Inna Kruman, editor. InTech Open Publisher. doi:10.5772/25117 http://www.intechopen.com/books/dna-repair/meiosis-as-an-evolutionary-adaptation-for-dna-repair
^ Jump up to: a b Bernstein, H; Bernstein, C (2010). "Evolutionary origin of recombination during meiosis". BioScience. 60 (7): 498–505. doi:10.1525/bio.2010.60.7.5.
Jump up ^ Dangel, NJ; Knoll, A; Puchta, H (2014). "MHF1 plays Fanconi anaemia complementation group M protein (FANCM)-dependent and FANCM-independent roles in DNA repair and homologous recombination in plants.". Plant J. 78 (5): 822–33. PMID 24635147. doi:10.1111/tpj.12507.
Jump up ^ Saponaro, M; Callahan, D; Zheng, X; Liberi, G (2010). "Cdk1 Targets Srs2 to Complete Synthesis-Dependent Strand Annealing and to Promote Recombinational Repair". PLoS Genet. 6 (2): e1000858. doi:10.1371/journal.pgen.1000858. Retrieved 20 March 2015.
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Jump up ^ Steinboeck, F (2010). "The relevance of oxidative stress and cytotoxic DNA lesions for spontaneous mutagenesis in non-replicating yeast cells.". Mutat Res. 688 (1–2): 47–52. PMID 20223252. doi:10.1016/j.mrfmmm.2010.03.006.
Jump up ^ Nedelcu, M; Marcu, O; Michod, RE (2004). "Sex as a response to oxidative stress: a twofold increase in cellular reactive oxygen species activates sex genes". Proc. R. Soc. B. 271: 1591–1596. PMC 1691771  . PMID 15306305. doi:10.1098/rspb.2004.2747. Retrieved 10 March 2015.
Jump up ^ Charpentier, X (2010). "Antibiotics and UV Radiation Induce Competence for Natural Transformation in Legionella pneumophila". Journal of Bacteriology. 193 (5): 1114–1121. PMC 3067580  . PMID 21169481. doi:10.1128/JB.01146-10. Retrieved 14 March 2015.
Jump up ^ Keeney, S; Giroux, CN; Kleckner, N (1997). "Meiosis-Specific DNA Double-Strand Breaks Are Catalyzed by Spo11, a Member of a Widely Conserved Protein Family". Cell. 88 (3): 375–84. PMID 9039264. doi:10.1016/S0092-8674(00)81876-0.
Jump up ^ Sauvageau, S; Stasiak, Az; Banville, I; Ploquin, M; Stasiak, A; Masson, Jy (June 2005). "Fission Yeast Rad51 and Dmc1, Two Efficient DNA Recombinases Forming Helical Nucleoprotein Filaments" (Free full text). Molecular and Cellular Biology. 25 (11): 4377–87. ISSN 0270-7306. PMC 1140613  . PMID 15899844. doi:10.1128/MCB.25.11.4377-4387.2005.
^ Jump up to: a b Pochart P, Woltering D, Hollingsworth NM (1997). "Conserved properties between functionally distinct MutS homologs in yeast". J. Biol. Chem. 272 (48): 30345–9. PMID 9374523. doi:10.1074/jbc.272.48.30345.
Jump up ^ Winand NJ, Panzer JA, Kolodner RD (1998). "Cloning and characterization of the human and Caenorhabditis elegans homologs of the Saccharomyces cerevisiae MSH5 gene". Genomics. 53 (1): 69–80. PMID 9787078. doi:10.1006/geno.1998.5447.
Jump up ^ Bocker T, Barusevicius A, Snowden T, Rasio D, Guerrette S, Robbins D, Schmidt C, Burczak J, Croce CM, Copeland T, Kovatich AJ, Fishel R (1999). "hMSH5: a human MutS homologue that forms a novel heterodimer with hMSH4 and is expressed during spermatogenesis". Cancer Res. 59 (4): 816–22. PMID 10029069.
Jump up ^ Krishnaprasad GN, Anand MT, Lin G, Tekkedil MM, Steinmetz LM, Nishant KT (2015). "Variation in crossover frequencies perturb crossover assurance without affecting meiotic chromosome segregation in Saccharomyces cerevisiae". Genetics. 199 (2): 399–412. PMC 4317650  . PMID 25467183. doi:10.1534/genetics.114.172320.
Jump up ^ Church K, Wimber DE (1969). "Meiosis in the grasshopper: chiasma frequency after elevated temperature and x-rays". Can. J. Genet. Cytol. 11 (1): 209–16. PMID 5797806. doi:10.1139/g69-025.
Jump up ^ Westerman M (1971). "The effect of x-irradiation on chiasma frequency in Chorthippus brunneus". Heredity. 27 (1): 83–91. PMID 5289295. doi:10.1038/hdy.1971.73.
Jump up ^ "genetic recombination".
Jump up ^ Genetic Recombination
^ Jump up to: a b c Smith, George P. (1976-01-01). "Evolution of Repeated DNA Sequences by Unequal Crossover". Science. 191 (4227): 528–535. JSTOR 1741301. doi:10.1126/science.1251186.
Jump up ^ Graur, Dan; Li, Wen-Hsiung (2000-01-01). Fundamentals of Molecular Evolution. Sinauer. ISBN 9780878932665.
^ Jump up to: a b Puchta, Holger (2005-01-01). "The repair of double-strand breaks in plants: mechanisms and consequences for genome evolution". Journal of Experimental Botany. 56 (409): 1–14. ISSN 0022-0957. PMID 15557293. doi:10.1093/jxb/eri025.
Jump up ^ Metzenberg, Ab; et al. (May 1991). "Homology Requirements for Unequal Crossing Over in Humans.". Genetics.
Jump up ^ Robberecht, Caroline; Voet, Thierry; Esteki, Masoud Zamani; Nowakowska, Beata A.; Vermeesch, Joris R. (2012-12-03). "Non-allelic homologous recombination between retrotransposable elements is a driver of de novo unbalanced translocations". Genome Research. 23: gr.145631.112. ISSN 1088-9051. PMC 3589530  . PMID 23212949. doi:10.1101/gr.145631.112.
^ Jump up to: a b c Kunkel, Thomas A.; Erie, Dorothy A. (2005-01-01). "Dna Mismatch Repair". Annual Review of Biochemistry. 74 (1): 681–710. PMID 15952900. doi:10.1146/annurev.biochem.74.082803.133243.
Jump up ^ Surtees, J. A., J.A.; Argueso, J.L.; Alani, E. (2004). "Mismatch Repair Proteins: Key Regulators of Genetic Recombination". Cytogenetic and Genome Research. 107: 146–59. doi:10.1159/000080593.
[hide]
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Genetics: homologous recombination / mobile genetic elements
Primarily prokaryotic
Conjugation
Transduction
Transformation
Occurs in eukaryotes
Transfection
Chromosomal crossover
Gene conversion
Fusion gene
Horizontal gene transfer
Sister chromatid exchange
Transposon
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