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Step by Step (TV series)
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Step by Step
Genre
Family sitcom
Created by
William Bickley
Michael Warren
Developed by
Thomas L. Miller
Robert L. Boyett
Starring
Patrick Duffy
Suzanne Somers
Staci Keanan
Brandon Call
Angela Watson
Christine Lakin
Patrika Darbo
Christopher Castile
Josh Byrne
Peggy Rea
Sasha Mitchell
Kristina Meyering
Lauren Meyering
Emily Mae Young
Jason Marsden
Bronson Pinchot
Theme music composer
Jesse Frederick
& Bennett Salvay
Opening theme
"Second Time Around",
performed by Jesse Frederick and Teresa James
Ending theme
"Second Time Around" (instrumental)
(season 1, used sporadically afterwards)
Composer(s)
Jesse Frederick &
Bennett Salvay (both; seasons 1–2)
Gary Boren (seasons 3–5)
Steven Chesne (seasons 3–7)
Country of origin
United States
Original language(s)
English
No. of seasons
7
No. of episodes
160 (list of episodes)
Production
Executive producer(s)
Thomas L. Miller
Robert L. Boyett
William Bickley
Michael Warren
Alan Eisenstock & Larry Mintz (season 1)
Ross Brown (seasons 2–7)
Bob Rosenfarb (seasons 6–7)
Production location(s)
Sony Pictures Studios
Culver City, California (1991–1993)
Warner Bros. Studios,
Burbank, California (1993–1998)
Camera setup
Film; Multi-camera
Running time
Approx. 23 minutes (per episode)
Production company(s)
Bickley-Warren Productions
Miller-Boyett Productions
Lorimar Television (1991–1993)
Warner Bros. Television
(1993–1998)
Distributor
Warner Bros. Television Distribution
Release
Original network
ABC (1991–1997)
CBS (1997–1998)
Picture format
480i (SDTV)
Original release
September 20, 1991 – June 26, 1998
Step by Step is an American television sitcom that aired for seven seasons. It ran on ABC as part of its TGIF Friday night lineup from September 20, 1991 to August 15, 1997, then moved to CBS, where it aired from September 19, 1997 to June 26, 1998. Patrick Duffy and Suzanne Somers starred as single parents, each with three children, who spontaneously marry after meeting on vacation, resulting in their becoming the heads of a large blended family.
Contents
1 Premise
2 Characters
2.1 Foster family
2.2 Lambert family
2.3 Others
3 Episodes
4 Production
4.1 Theme song and title sequence
5 Syndication
6 Other media
6.1 Streaming
6.2 DVD releases
7 Reception
8 References
9 External links
Premise[edit]
Frank Lambert, a divorced contractor, has three children: John Thomas (J.T.), tomboyish Alicia (Al), and Brendan. Carol Foster, a widowed beautician, also has three children: Dana, Karen, and Mark. Both families live in a substantially fictionalized version of Port Washington, Wisconsin.
Frank and Carol meet while vacationing in Jamaica and impulsively get married after a whirlwind courtship. They had planned to keep their marriage a secret, but Frank accidentally blurts it out to J.T. at a barbecue he and Carol hold to introduce all the children. The children are surprised and angry at first.
Each episode depicts typical situations for a new blended family. Family members’ differences cause arguments and resentments, but over time they grow to tolerate and become loyal to one another.
During the series' seven-season run, the main cast changed as the storyline progressed. Frank's nephew, Cody Lambert, was introduced early in the first season as a recurring character (becoming a series regular in the second season) and lived in his van in the driveway; he then left the series in 1996. In the 1994–95 season finale, Carol gave birth to a daughter named Lily. Like many other sitcom babies, she was suddenly aged five years in the sixth season, after having spent the entire fifth season depicted as a baby. In the sixth season, J.T.'s friend, Rich Halke, moved in with him and the family after becoming Dana's boyfriend. Meanwhile J.T. meets Samantha, the woman of his dreams, and they date off and on for the sixth and seventh seasons. Also in the sixth season Carol got a partner, Jean-Luc Rieupeyroux, at her hair salon. In the seventh season Brendan is written out of the show as is Jean-Luc.
Characters[edit]
Cast of Step by Step (seasons 2–5)
Foster family[edit]
Carol Foster-Lambert (née Williams) (Suzanne Somers) – the matriarch of the Foster family, who works as a beautician and runs a hair salon out of a room in her house (which is located next to the kitchen), originally with the cooperation of her mother, Ivy, and sister, Penny. Eventually, with Frank's help, she opens a bigger studio at a stand-alone location in the sixth season.
Dana Foster (Staci Keanan) – Dana is the eldest child in the Foster family. She is depicted as a smart, wisecracking feminist. She is often seen being openly hostile to the Lamberts, particularly J.T., (with whom she frequently trades barbs regarding his hormonal tendencies with girls, his lack of intelligence and his poor hygiene), Frank (due partly to his perceived "caveman" behavior), and Cody (who has an unrequited crush on her, despite being technically related to her as an in-law). She and J.T. generally do not get along.
Karen Foster (Angela Watson) – Karen is the middle child in the Foster family. She is an aspiring model and sometimes a country singer, and is portrayed as vain and not very bright, but with frequent moments of level-headedness.
Mark Foster (Christopher Castile) – Mark is the youngest child in the Foster family, until Lily is born. He is into computers and academics and has a tendency to be timid.
Lambert family[edit]
Frank Lambert (Patrick Duffy) – Frank is the patriarch of the Lambert family, and works as a contractor with his own company. He is laidback and an avid sports fan, particularly of the Green Bay Packers.
John Thomas "J.T." Lambert (Brandon Call) – J.T. is the oldest child in the Lambert family. He is depicted as a slacker, is into sports (like his father, he is a Packers fan), and is portrayed as academically challenged, which is later attributed to dyslexia. He resents his new stepfamily, particularly Dana, whom he often calls "Barky" and also trades other various insults at her expense.
Alicia "Al" Lambert (Christine Lakin) – Al is the middle child in the Lambert family. She is depicted as a tomboyish all-American girl, who later matured and softened. She is typically addressed by her nickname, the more masculine name "Al", and is rarely referred to as "Alicia". Several episodes during the seventh season centered on her newfound interest in acting. She is openly hostile towards her stepfamily.
Brendan Lambert (Josh Byrne) – Brendan is the youngest child in the Lambert family, until Lily is born. He is portrayed as a shy, carefree youngster, and one of the most accepting of his new stepfamily. He appeared less and less as the show progressed, especially after Lily was introduced in the season four episode "A Foster/Lambert Production". When the show moved from ABC to CBS, he was written out of the series; the producers later admitted in a TV Guide interview that despite his absence, the Lamberts would still refer to their "seven children", making him an unseen character for the final season. Actor Josh Byrne appeared on the sitcoms Who's the Boss? and The Family Man.[1] He portrays the younger version of Billy Crystal's character in Mr. Saturday Night, which also features Jason Marsden, who appears in later run of the series.
Cody Lambert (Sasha Mitchell) – Frank's nephew, who appears as a recurring cast member in the first season, then is upgraded to a regular character in the second season. Sporting a crew cut and a Valley teen accent and vocabulary (such Valley-isms as "dude" and "cheesed off"), he often shows maturity and intelligence belying his dimwitted nature. He lived in his van in the driveway. He was written out of the series after the fifth season after legal problems that began with Mitchell's 1995 arrest on accusations of domestic violence by his first wife, Jeannette Robbins,[2][3][4] charges which were later dropped and he was cleared of wrongdoing. He returned as a guest star for one episode in the seventh season. Mitchell previously played James Beaumont, the nephew of Duffy's character Bobby Ewing in Dallas.
Others[edit]
Penny Baker (Patrika Darbo; season 1) – Carol's man-hungry sister; she was written out of the series after the first season.
Ivy Baker (Peggy Rea; season 1) – Carol's outspoken mother; she was written out of the series after the first season.
Lily Foster-Lambert (Lauren Meyering and Kristina Meyering; seasons 4–5), (Emily Mae Young; seasons 6–7) – Lily is Frank and Carol's biological child, who is introduced in the season four episode "A Foster/Lambert Production". Originally depicted as a baby, her age is retconned to five in the sixth season after she is SORASed. For her age she is very smart and is always asking questions of everyone.
Rich Halke (Jason Marsden; 1 appearance in season 3, seasons 5–7) – Rich is J.T.'s best friend. He is depicted as being both a slacker and a seriously devoted person. He later began dating Dana (to the others' dismay) in the sixth season. He was named after Richard P. Halke, who served a member of the series' writing staff from seasons one through three, and served as a story editor during the third one. In his one appearance in the latter he plays a guy named Doug.
Jake "Flash" Gordon (Jeff Juday; season 5) – Flash (only appears in 4 episodes) is a goofy, but well-meaning handyman hired by Frank towards the end of the fifth season. He joined the family on their trip to Walt Disney World, where he attempted to visit every attraction. At the very end of season, Frank and Carol invited him to move in with them, which he accepted. Afterwards, he was never mentioned again. According to Jeff Juday, he was written in as a replacement for Cody.[5] The following season, Flash was replaced by Jean-Luc.
Jean-Luc Rieupeyroux (Bronson Pinchot; season 6) – Jean-Luc is a male beautician, and serves as Carol's business partner. He was brought in to take the place of Cody on the series, but disappeared when Bronson Pinchot took the title role in the short-lived CBS sitcom Meego (which aired alongside Step by Step when the latter moved to CBS for its seventh season in 1997).
Samantha Milano (Alexandra Adi; seasons 6–7) – Samantha is J.T.'s one-time girlfriend, who was introduced at the very end of the season six episode "The "L" Word". They date off and on for 2 seasons. She worked as a mechanic in a garage.
Episodes[edit]
Main article: List of Step by Step episodes
Season
Episodes
Originally aired
First aired
Last aired
Network
1 22 September 20, 1991 (1991-09-20) April 24, 1992 (1992-04-24) ABC
2 24 September 18, 1992 (1992-09-18) May 21, 1993 (1993-05-21)
3 23 September 24, 1993 (1993-09-24) May 20, 1994 (1994-05-20)
4 24 September 23, 1994 (1994-09-23) May 19, 1995 (1995-05-19)
5 24 September 22, 1995 (1995-09-22) May 17, 1996 (1996-05-17)
6 24 March 7, 1997 (1997-03-07) August 15, 1997 (1997-08-15)
7 19 September 19, 1997 (1997-09-19) June 26, 1998 (1998-06-26) CBS
Production[edit]
This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (January 2017) (Learn how and when to remove this template message)
The series was created and executive produced by William Bickley and Michael Warren, and developed and executive produced by Thomas L. Miller and Robert L. Boyett.[6] It was produced by Bickley-Warren Productions, Miller-Boyett Productions and Lorimar Television (the latter doing so from 1991 to 1993, when Warner Bros. Television assumed production responsibilities for it and Lorimar's other television series after shared corporate parent Time Warner consolidated the two production companies). The casting of Patrick Duffy fulfilled a contractual obligation that Lorimar made to give him a new show after his previous series, Dallas (which was also produced by Lorimar), had ended its run. It was created off of the idea of combining two of the most popular television stars from the 1970s known for their good looks (Duffy and Somers) to star as parents to attract adult viewers, with current teen celebrities (Staci Keanan from My Two Dads and Going Places, Brandon Call from Baywatch and Sasha Mitchell from Dallas) to star as their children to attract children and teen viewers.
Staci Keanan and Christopher Castile had previously appeared on the Miller-Boyett-produced ABC sitcom Going Places, which debuted the season prior to Step By Step, playing characters with no familial relation (Keanan – who played Lindsay Bowen, the teenage neighbor of the show's adult characters – as a series regular, Castile as a recurring character). Keanan was the first of the two Going Places stars to be cast on Step By Step in the spring of 1991. Castile, who had played gawky child Sam Roberts on Going Places, brought the same character traits to the Mark Foster role, which was speculated to be Miller-Boyett's continued attempt to give the Family Matters character Steve Urkel a white counterpart. In a similar instance of hiring actors over from their other TV series, the producers also cast Josh Byrne as Brendan Lambert on Step By Step, after he had just finished a supporting role as Patrick Kozak on Miller-Boyett's single-season CBS sitcom The Family Man.
When the series was casting its characters, child actor Jarrett Lennon originally landed the role of Mark Foster. Lennon had been chosen by the producers after guest starring in the last original episode of the Miller-Boyett series The Hogan Family, which was produced in late 1990. After shooting the original (unaired) pilot for Step By Step, Lennon was dismissed from the role of Mark, and the producers later replaced him with Castile (who had blonde hair like on-screen mother Somers, as opposed to Lennon having brown hair). Most of Lennon's pilot scenes were reshot with Castile, but during the first season, footage of Lennon remained in the show's opening title sequence. Lennon only appeared in wide shots with the Lambert/Foster family or, in the case of Suzanne Somers' credit scene, fleetingly appearing at the bottom of camera view as the kids huddled around Somers. All traces of Lennon were edited out by the second season. In 1996, these two actors had the tables turned on each other; Castile served as the original voice of Eugene Horowitz on the Nickelodeon animated series Hey Arnold!, but after a few episodes was replaced by Lennon, who voiced Eugene for the remainder of the show's first season.
Going Places creators Robert Griffard and Howard Adler would end up employed with Step By Step as co-executive producers and members of its writing staff until the show's fifth season; Adler and Griffard later wrote an additional episode of the show as freelancers in the seventh season. Patrick Duffy directed several episodes, starting with the second season. The house shown in establishing shots for scenes set at the Lambert-Foster home is located at 2011 Fletcher Avenue in South Pasadena, California, although the series was actually filmed at Warner Bros. Studios in Burbank, California.
ABC chose to delay the series' sixth season to the 1996–97 mid-season (premiering in March 1997), in order to make room on that season's fall schedule for freshman sitcoms Sabrina the Teenage Witch and Clueless, which joined established series Family Matters and Boy Meets World on the TGIF lineup; the network canceled it after six seasons in May 1997, due to declining ratings. CBS concurrently reached a deal with Miller-Boyett Productions to acquire the rights to it and Family Matters from ABC, as that network attempted to build its own Friday night lineup of family-friendly situation comedies for the fall of 1997, called the "CBS Block Party".[7] The series' ratings, which had been declining for several seasons, continued to erode, and the show ended its run in June 1998. It ended without an official series finale, although the last episode centered on Frank and Carol considering selling the house.
Theme song and title sequence[edit]
The series' theme song "Second Time Around" was written and composed by Jesse Frederick and Bennett Salvay (both wrote the themes for other sitcoms produced by Miller-Boyett Productions such as Full House, Perfect Strangers, and Family Matters), and was performed by Frederick and Teresa James. The full 87-second version of it was only used during the first season. It was routinely edited over the following three seasons to allow additional time for scenes: the fourth verse was removed and the chorus was truncated in the second season edit, the kid chorus accompanying Jesse Frederick during the chorus' lyrics was removed in the version used during the third season, and the edit heard during most of the fourth season and the entire fifth season (which lasted for only 65 seconds) eliminated the electric guitar/drum/symphonic instrumental at the beginning.
The theme song and title sequence were dropped entirely for the sixth season, relegating the credits for the show's main cast and principal producers to appearing over each episode's cold open (which was preceded for the final two seasons by an abbreviated opening bumper featuring the family applauding as Lily blows out the candles on her birthday cake). The theme song was brought back for the seventh season, upon the show's move from ABC to CBS, although it was edited down to include only the chorus and the closing instrumental flourish (restoring the original longer symphonic instrumental heard in the original long version, and the edits used for the second and third seasons).
The amusement park seen in the opening credits is depicted as being located along the shores of Lake Michigan in Wisconsin (no amusement park exists there). The one used is actually Six Flags Magic Mountain in Valencia, California, located miles inland from the Pacific Ocean. The body of water depicted in the opening and closing credit sequences (the latter being seen on in season one) located next to the roller coaster – which is digitally inserted into that particular excerpt – is placed over what in actuality is the parking lot of Six Flags Magic Mountain.
Syndication[edit]
In September 1995, Warner Bros. Domestic Television Distribution began distributing the series for broadcast in off-network syndication.
ABC Family was the first to acquire cable television rights to the series, and it became one of the cable channel's longest-running off-network syndicated programs in its history. Reruns began airing on there in 2001 (on what was then known as Fox Family), airing in various timeslots during its run ranging from late afternoon to the morning hours. On March 26, 2010, ABC Family's contract expired after less than nine years.[8]
The series returned to U.S. syndication on October 7, 2013, when the Hub Network began airing reruns;[9] the network dropped it on October 12, 2014, as part of a change in the focus.
In Australia, Step by Step aired on the Seven Network from 1991 to 1995 and on the Nine Network from 1996 to 2000. In 2011, Step by Step was acquired by 7TWO. In 2015, 111 Greats started airing the whole series.[citation needed]
On September 29, 2017, Hulu acquired the streaming rights to Step by Step along with fellow Warner Bros. TV properties Family Matters, Full House, Hangin' with Mr. Cooper and Perfect Strangers,[10] in addition to fellow ABC programs Boy Meets World, Dinosaurs and Home Improvement.[11]
Other media[edit]
Streaming[edit]
All episodes are available for online streaming on Hulu.[12] .
DVD releases[edit]
Warner Home Video released a six-episode Television Favorites collection on DVD on June 27, 2006.[13] Much like other television shows from it, the Step by Step edition has gone out of print. The first season was released on DVD June 12, 2018.[14]
Reception[edit]
Reviewing the pilot episode, Jean Rosenbluth of Variety wrote that, despite being an unoriginal clone of The Brady Bunch, it is a "modestly amusing, occasionally heartwarming show".[15]
References[edit]
Jump up ^ Dabnam, Betty (18 December 1993). "Meet Josh Byrne". The Bryan Times. Retrieved 7 December 2010.
Jump up ^ "Sitcom Star Pleads Not Guilty in Spousal Assault : Courts: Sasha Mitchell of 'Step by Step' is charged with three new misdemeanor counts in alleged beating. Attorney says police fabricated case". Los Angeles Times.
Jump up ^ Arrest Warrant Issued for Actor Sasha Mitchell
Jump up ^ Jake Jarmell. "Shocking! An Interview with Cody Lambert". oocities.org.
Jump up ^ Michael Portantiere (July 23, 2001). "Just Jeff – Theater News – Jul 23, 2001". theatermania.
Jump up ^ "Step by Step Review - TV Reviews and News - EW.com". EW.com.
Jump up ^ Hal Boedeker (July 18, 1997). "He's A Goober But CBS Has A Lot Riding On Urkel TV". Orlando Sentinel. Tribune Publishing. Retrieved April 12, 2017.
Jump up ^ Step by Step Leaving ABC Family After 9 Years; ABC Family March 2010, Sitcoms Online, February 10, 2010.
Jump up ^ "'Step by Step' to Premiere October 7 on the Hub" (Press release). Hub Network. September 6, 2013. Retrieved August 22, 2016 – via TV By The Numbers.
Jump up ^ Pedersen, Erik (July 27, 2017). "Hulu Gets SVOD Rights To 'Full House,' 'Family Matters' & Other 'TGIF' Comedies – TCA". Deadline. Retrieved October 3, 2017.
Jump up ^ Hatchett, Keisha (September 29, 2017). "This Is Not a Drill: Boy Meets World Is Now On Hulu". TVGuide.com. Retrieved October 28, 2017.
Jump up ^ "Step by Step (TV)". Hulu. Retrieved 2018-04-11.
Jump up ^ "Step by Step (Television Favorites Compilation) (1991)". amazon.com. Retrieved 2008-10-20.
Jump up ^ https://www.facebook.com/tvshowsondvd/posts/10156202608481063 accessed Jun. 7, 2018
Jump up ^ Rosenbluth, Jean (1994). Variety TV REV 1991–92 17. Taylor & Francis. September 20, 1991. ISBN 9780824037963.
External links[edit]
Television in the United States portal
Comedy portal
Wisconsin portal
1990s portal
Step by Step on IMDb
Step by Step at TV.com
Step by Step at epguides.com
show
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TGIF
1980
Perfect Strangers (1989–1993)
Just the Ten of Us (1989–1990)
Full House (1989–1991)
Family Matters (1989–1997)
T G I f
1990
New Attitude (1990)
Going Places (1990–1991)
Baby Talk (1991–1992)
Dinosaurs (1991–1993)
Hi Honey, I'm Home! (1991)
Step By Step (1991–1997)
Billy (1992)
Camp Wilder (1992–1993)
Where I Live (1993)
Getting By (1993)
Home Free (1993)
Hangin' with Mr. Cooper (1993–1996)
Boy Meets World (1993–2000)
Sister, Sister (1994–1995)
On Our Own (1994–1995)
Muppets Tonight (1996)
Aliens in the Family (1996)
Clueless (1996–1997)
Sabrina the Teenage Witch (1996–2000)
Teen Angel (1997–1998)
You Wish (1997–1998)
Two of a Kind (1998–1999)
Brother's Keeper (1998–1999)
The Hughleys (1999–2000)
Odd Man Out (1999–2000)
2000
George Lopez (2003–2004)
Life with Bonnie (2003–2004)
Hope & Faith (2003–2005)
Married to the Kellys (2003–2004)
The Big House (2004)
8 Simple Rules (2004–2005)
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show
v
t
e
Miller-Boyett Productions
Miller-Milkis
Productions
TV series
Petrocelli (1974–76)
Happy Days (1974–84)1
Laverne & Shirley (1976–83)1
Blansky's Beauties (1977)
Mork & Mindy (1978–82)1
Makin' It (1979)
Out of the Blue (1979)
$weepstake$ (1979)
Angie (1979–80)
Films
The Heist (1972)
The Weekend Nun (1972)
Silver Streak (1976)
Walkin' Walter (1977)
Foul Play (1978)
Miller-Milkis-Boyett
Productions
TV series
Goodtime Girls (1980)
Bosom Buddies (1980–82)
Foul Play (1981)
Joanie Loves Chachi (1982–83)
Films
The Best Little Whorehouse in Texas (1982)
Miller-Boyett
Productions
The Hogan Family (1986–91)
Perfect Strangers (1986–93)
Full House (1987–95)
Family Matters (1989–98)
The Family Man (1990–91)
Going Places (1990–91)
Step by Step (1991–98)
Getting By (1993–94)
On Our Own (1994–95)
Fuller House (2016–present)
Miller-Boyett-Warren
Productions
Meego (1997)
Two of a Kind (1998–99)
Robert L. Boyett
Productions
Partners (2014)
See also
Charles Fox
Jesse Frederick
Bickley-Warren Productions
Paramount Television
CBS Paramount Television
CBS Television Studios
Lorimar Television
Warner Bros. Television
1 This series began being produced under the Miller-Milkis-Boyett banner beginning with the 1980–81 television season.
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Michael Clarke Duncan
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Michael Clarke Duncan
Duncan in January 2009
Born
(1957-12-10)December 10, 1957
Chicago, Illinois, U.S.
Died
September 3, 2012(2012-09-03) (aged 54)
Los Angeles, California, U.S.
Cause of death
Complications of heart attack
Resting place
Forest Lawn Memorial Park (Hollywood Hills)
Occupation
Actor
Years active
1995–2012
Height
6 ft 5 in (1.96 m)
Partner(s)
Omarosa Manigault
(2010–2012; his death)
Michael Clarke Duncan (December 10, 1957 – September 3, 2012) was an American actor, best known for his breakout role as John Coffey in The Green Mile (1999), for which he was nominated for the Academy Award for Best Supporting Actor and various similar honors. He also appeared in motion pictures such as Armageddon (1998), The Whole Nine Yards (2000), The Scorpion King (2002), Daredevil (2003) and Talladega Nights: The Ballad of Ricky Bobby (2006). His voice can be heard in films such as Brother Bear (2003), Kung Fu Panda (2008) and Green Lantern (2011).
Contents
[hide]
1 Early life
2 Career
3 Personal life
3.1 Death
4 Filmography
4.1 Films
4.2 Television
4.3 Video games
4.4 Music videos
5 References
6 External links
Early life
Duncan was born in Chicago, Illinois, and grew up in a single-parent household with his sister, Judy, and mother, Jean Duncan (a house cleaner), after his father left.[1][2] He always wanted to act, but had to drop out of the Communications program at Alcorn State University to support his family when his mother became ill. Duncan's large frame—6 feet 5 inches (196 cm)[3] and 315 pounds (143 kg)—helped him in his jobs digging ditches for the People's Gas Company and being a bouncer at several Chicago clubs.[4] Duncan also played basketball at Kankakee Community College and for one season at Alcorn State.[5]
In 1979, he participated in the Disco Demolition Night at Comiskey Park, home of the Chicago White Sox, where he was among the first 100 people to run onto the field and he slid into third base. During the ensuing riot his silver belt buckle was stolen[6] while he was stealing a baseball bat from the dugout.[7]
Career
Duncan took other security jobs while in Los Angeles while trying to get some acting work in commercials. During this time, he worked as a bodyguard for celebrities like Will Smith, Martin Lawrence, Jamie Foxx, LL Cool J, and The Notorious B.I.G., all the while doing bit parts in television and films. When rapper Notorious B.I.G. was killed in 1997, Duncan quit the personal-protection business.[3]
After having begun his career with several bit parts playing bouncers in films such as Bulworth and A Night at the Roxbury, Duncan first came to prominence when he was cast as Bear in the blockbuster Michael Bay action film Armageddon (1998). During the production of the film, Duncan struck up a friendship with castmate Bruce Willis and it was Willis' influence that helped him to get his breakout role as gentle giant John Coffey in the Frank Darabont film The Green Mile (1999). Starring alongside Tom Hanks, Duncan's acclaimed performance netted him an Academy Award nomination for Best Actor in a Supporting Role and a Golden Globe nomination for Best Performance by an Actor in a Supporting Role in a Motion Picture.[8][9]
Following his iconic dramatic turn as Coffey, Duncan was then cast in a string of films that helped to establish him as a star adept at both action and comedy: The Whole Nine Yards (2000), See Spot Run (2001), Planet of the Apes (2001), The Scorpion King (2002) (where he starred alongside his friend, Dwayne Johnson), and Daredevil (2003) (reuniting him with Armageddon co-star Ben Affleck) as Wilson Fisk, aka The Kingpin from Marvel Comics.
When Duncan was cast as the Kingpin in 2002,[10] he faced the dual challenge of portraying a typically white character[11][12] and having to gain 40 pounds to fit the character's large physique. In July 2006, Duncan showed interest in returning for the role of the Kingpin, but stated that he would not be willing to regain the weight that he had lost.[13] In 2009, he stopped eating meat and later appeared in a PETA ad campaign, touting the health benefits and his increased strength from a vegetarian diet.[14][15]
In 2005, Duncan appeared in two prominent action films, The Island (his second Michael Bay film) and Sin City (again alongside Bruce Willis) where he played Manute, a powerful mobster. Critic Roger Ebert singled out Duncan for praise for his role in The Island, writing that "[Duncan] has only three or four scenes, but they're of central importance, and he brings true horror to them."[16] Duncan appeared in a supporting role in the 2006 comedy Talladega Nights: The Ballad of Ricky Bobby as Lucius Washington and, in 2009, Duncan played Balrog in Street Fighter: The Legend of Chun-Li and starred as the titular Cleon "Slammin'" Salmon in Broken Lizard's farce The Slammin' Salmon.
Famous for his deep, resonant voice, Duncan also did speaking-only roles for films such as Brother Bear (2003) and its sequel, Brother Bear 2 (2006), Kung Fu Panda (2008) and Green Lantern (2011) playing the voice of Kilowog from DC Comics alongside Ryan Reynolds.
His other voice acting roles include' TV series such as Loonatics Unleased and Operation: Z.E.R.O., Quiznos commercials, and a number of video games such as Demon Stone, SOCOM II: U.S. Navy SEALs, The Suffering: Ties That Bind, Saints Row, Soldier of Fortune, and God of War II, where he provided the voice of the Titan Atlas. He additionally reprised his role as the Kingpin in Spider-Man: The New Animated Series.
In addition to his film roles, Duncan also guest starred in numerous television shows. Among these, he appeared in an episode of The Suite Life of Zack & Cody and a first-season episode of CSI: NY.[17] In 2008, he appeared as "Mr. Colt" in the second-season premiere of Chuck, "Chuck Versus the First Date"[18] and as a guest star on two episodes of Two and a Half Men. Most notably, in April 2011, Duncan guest starred on an episode of TV series Bones as Leo Knox which, in 2012, led to Duncan receiving his first starring role as the same character in the spinoff series The Finder. The Bones Season 8 episode "The Partners in the Divorce", which aired three weeks after his death, was dedicated to him.
During the week of May 14, 2012, Duncan appeared as a guest on the late night talk show The Late Late Show with Craig Ferguson when the show was taping for a week in Scotland. Duncan was one of the show's most frequent guests, appearing a total of 18 times, and, the day after Duncan's death in September, Ferguson began his show with a tribute to him. In January 2013 during The Late Late Show's winter break, reruns of the Scotland episodes were broadcast with a tribute to Duncan at the beginning of each of the five episodes, featuring Duncan on a pink background and the text "In memory of our friend Michael Clarke Duncan."
Personal life
At the time of his death, Duncan was engaged to reality television personality Omarosa Manigault.[19]
In 2013, Manigault appeared in the cast of The All-Star Celebrity Apprentice and played in Duncan's honor for his favorite charity and one he had benefited from himself, the Sue Duncan Children's Center. In episode 2 of the season, Manigault won $40,000 for the charity.[20]
Death
Crypt of Michael Clarke Duncan at Forest Lawn Hollywood Hills.
On July 13, 2012, Duncan was taken to Cedars Sinai Medical Center after suffering a heart attack.[21] Media reports suggested that his girlfriend, Omarosa Manigault, had tried to save his life by performing CPR.[22] Duncan's publicist, Joy Fehily, issued a statement on August 6 that read he was moved from the intensive-care unit but remained hospitalized following his heart attack.[23] On September 3, Duncan died in Los Angeles. Celebrations of Duncan's life were scheduled for a later date.[24]
Filmography
Films
Year
Title
Role
Notes
1995 Friday Craps Player Uncredited
1997 Back in Business Huge Guard N/A
1998 Caught Up BB N/A
1998 The Players Club Bodyguard Credited as Big Mike Duncan
1998 Bulworth Bouncer Credited as Michael "Big Mike" Duncan
1998 Armageddon Bear N/A
1998 A Night at the Roxbury Roxbury Bouncer Credited as Michael "Big Mike" Duncan
1999 The Underground Comedy Movie Gay Virgin N/A
1999 Breakfast of Champions Eli Credited as Michael Duncan
1999 The Green Mile John Coffey Broadcast Film Critics Association Award for Best Supporting Actor
Saturn Award for Best Supporting Actor
Black Reel Award for Best Supporting Actor
Nominated—Academy Award for Best Supporting Actor
Nominated—Chicago Film Critics Association Award for Best Supporting Actor
Nominated—Chicago Film Critics Association Award for Most Promising Actor
Nominated—Golden Globe Award for Best Supporting Actor – Motion Picture
Nominated—NAACP Image Award for Outstanding Actor in a Motion Picture
Nominated—MTV Movie Award for Best Breakthrough Performance
Nominated—Online Film Critics Society Award for Best Supporting Actor
Nominated—Screen Actors Guild Award for Outstanding Performance by a Male Actor in a Supporting Role
Nominated—Screen Actors Guild Award for Outstanding Performance by a Cast in a Motion Picture
2000 The Whole Nine Yards Franklin 'Frankie Figs' Figueroa N/A
2001 See Spot Run Murdoch N/A
2001 Cats & Dogs Sam Voice only
2001 Planet of the Apes Attar N/A
2002 The Scorpion King Balthazar N/A
2003 Daredevil Wilson Fisk/The Kingpin N/A
2003 George of the Jungle 2 Mean Lion Voice only
Direct-to-video
2003 Brother Bear Tug Voice only
2003 Kim Possible: A Sitch in Time Future Wade Voice only
2004 D.E.B.S. Mr. Phipps N/A
2004 George and the Dragon Tarik N/A
2004 Pursued Franklin N/A
2005 Racing Stripes Clydesdale Voice only
2005 Sin City Manute N/A
2005 The Golden Blaze Thomas Tatum/Quake Voice only
2005 Dinotopia: Quest for the Ruby Sunstone Stinktooth Voice only
2005 The Island Starkweather Two Delta/Jamal Starkweather N/A
2005 Stewie Griffin: The Untold Story The Stork Voice only
Direct-to-video
2005 The Land Before Time XI: Invasion of the Tinysauruses Big Daddy Voice only
2006 Talladega Nights: The Ballad of Ricky Bobby Lucius Washington N/A
2006 Brother Bear 2 Tug Voice only
2006 School for Scoundrels Lesher N/A
2006 One Way The General N/A
2006 Air Buddies Wolf Voice only
2007 The Last Mimzy Nathanial Broadman N/A
2007 Slipstream Mort/Phil Henderson/Patrolman N/A
2008 Welcome Home Roscoe Jenkins Otis N/A
2008 Delgo Elder Marley Voice only
2008 Kung Fu Panda Commander Vachir Voice only
2008 American Crude Spinks N/A
2009 The Slammin' Salmon Cleon Salmon N/A
2009 Street Fighter: The Legend of Chun-Li Balrog/M. Bison (Japan) N/A
2010 Redemption Road Augy N/A
2010 Cats & Dogs: The Revenge of Kitty Galore Sam Voice only
2011 Cross Erlik Direct-to-video
2011 A Crush on You Big Jim Nelson N/A
2011 Legend of Kung Fu Rabbit Slash Voice only
2011 Green Lantern Kilowog Voice only
2012 From the Rough Roger Posthumous release in the United States on 2014
2012 In the Hive Mr. Hollis N/A
2013 A Resurrection Addison Posthumous release seven months after Duncan's death
2015 The Challenger Duane Posthumous release, Duncan's last performance in a film
Television
Year
Title
Role
Notes
1995 Renegade Shake Episode: "Living Legend"
1995 The Fresh Prince of Bel-Air Tiny Episode: "Bourgie Sings the Blues"
1995 Married... with Children Bouncer Episode: "Flight of the Bumblebee"
1996 Skwids Body Builder N/A
1996 Weird Science Cardinal Carnage Episode: "Men in Tights"
1997 The Jamie Foxx Show Inmate Episode: "Little Red Corvette"
1997 Sparks Frank Episode: "Self Defense"
1997 The Wayans Bros. Big Mike Episode: "I Do..."
1997 Living Single Security Guard Episode: "High Anxiety"
1997 Built to Last N/A Episode: "A Family Affair"
1998 Arliss Lucian Balboa Episode: "Fans First"
1999 Sister, Sister Big Earl Episode: "Before There Was Hip Hop..."
2001 They Call Me Sirr Coach Griffin Television film
2002 King of the Hill Coach Webb Episode: "The Son Also Roses"
2003, 2005 The Adventures of Jimmy Neutron: Boy Genius Commander Baker Voice only
Episode: "My Big Fat Spy Wedding" and "Operation: Rescue Jet Fusion"
2003 Spider-Man: The New Animated Series The Kingpin / Wilson Fisk Voice only
Episode: "Royal Scam" and "Keeping Secrets"
2003 The Proud Family Mongo Voice only
Episode: "Smackmania 6: Mongo vs. Mama's Boy"
2004 Static Shock Rashid 'The Rocket' Randall Voice only
Episode: "Linked"
2004 The Fairly OddParents Rockwell Voice only
Episode: "Crash Nebula"
2004 George Lopez Dr. Holland Episode: "George to the Third Power"
2005–2007 Loonatics Unleashed Massive 4 episodes
2005 Teen Titans Hayden/Krall Voice only
Episode: "Cyborg the Barbarian"
2005 CSI: NY Quinn Sullivan Episode: "The Closer"
2006–2007, 2012 Family Guy Black Co-Worker/Boy on Mushrooms/Fozzie Bear 4 episodes
2006 Minoriteam Balactus Episode: "Balactus: Part 1" and "Balaztus: Part 2"
2008–2009 Two and a Half Men Jerome Episode: "The Two Finger Rule" and "The Mooch at the Boo"
2008 The Suite Life of Zack & Cody Coach Little Episode: "Benchwarmers"
2008 Chuck Colt Episode: "Chuck Versus the First Date"
2011 Bones Leo Knox Episode: "The Finder"
2012 The Finder Leo Knox 13 episodes
2012 The High Fructose Adventures of Annoying Orange Marshmallow King Voice only
Episode: "Marshmalia"
2012 Ultimate Spider-Man Groot Voice only
Video games
Year
Title
Role
Notes
1995 Panic in the Park Security Guard N/A
2000 Star Trek: Klingon Academy Opening Movie Klingons N/A
2000 Soldier of Fortune Hawk N/A
2003 SOCOM II: U.S. Navy SEALs SEAL Operative WARDOG Credited as Michael Clark Duncan
2004 Forgotten Realms: Demon Stone Slaad Lord Ygorl N/A
2005 The Suffering: Ties That Bind Blackmore N/A
2006 Saints Row Benjamin King N/A
2007 God of War II Atlas N/A
2013 Saints Row IV N/A Credited as Benjamin King in the game's credits in memoriam to him, Duncan was set to reprise his role as King in IV, with some voice work already done for the game before his death. During the game's credits, the song "Just a Friend" by Biz Markie is played as all the cast members sing along, the song fades out with Duncan's track. He was replaced by Terry Crews.
Music videos
Year
Artist
Title
Role
Notes
1994 Quo "Huh What" featuring Redman (rapper)
1995 R. Kelly featuring The Isley Brothers "Down Low (Nobody Has to Know)" Security Guard Cameo appearance
References
Jump up ^ "CNN – 'Green Mile''s giant has taken massive strides – December 10, 1999". CNN.
Jump up ^ USA WEEKEND Magazine[dead link]
^ Jump up to: a b Biography for Michael Clarke Duncan on IMDb
Jump up ^ Beale, Lewis. "Michael Clarke Duncan gets funny in 'Slammin' Salmon'." Newsday, (Melville, NY) 16 Dec. 2009: Newspaper Source Plus. Web. 17 Sept. 2012.
Jump up ^ "Former Alcorn hoops coach remembers actor Michael Clarke Duncan". CBS Sports.
Jump up ^ Zwecker, Bill. "Love may have bloomed again on set for 'Garden State' star." Chicago Sun-Times September 28, 2006
Jump up ^ Adam Caldarelli 3:09 p.m. CDT, May 20, 2006 (2006-05-20). "Caldarelli, Adam. "From the Cubicle." ''Chicago Tribune'' May 20, 2006". Chicagotribune.com. Retrieved 2012-11-01. CS1 maint: Multiple names: authors list (link)
Jump up ^ "'Green Miles giant has taken massive strides". CNN.
Jump up ^ Awards for The Green Mile on IMDb
Jump up ^ McNary, Dave (January 9, 2002). "Duncan takes on Daredevil". Variety. Retrieved 2008-02-23.
Jump up ^ Daniel Robert Epstein. "Michael Clarke Duncan Interview". UGO. Archived from the original on 2008-12-01. Retrieved 2008-02-23.
Jump up ^ Ryan J. Downey (February 6, 2003). "Ben Affleck Dares to Dream Daredevil". MTV. Retrieved 2008-02-21.
Jump up ^ Chavez, Kellvin (July 6, 2006). "Duncan Not in Transformers But Talks Daredevil 2!". LatinoReview.com. Retrieved 2008-12-23.
Jump up ^ "Green Mile Star Joins Veggie Campaign," KSHB.com, 12 May 2012.
Jump up ^ 04 Sep 2012 (2012-09-04). "Michael Clarke Duncan". London: Telegraph. Retrieved 2012-11-01.
Jump up ^ Ebert, Roger (2005). "The Island", Chicago Sun Times, 22 July 2005, accessed 13 September 2012
Jump up ^ The Closer on IMDb
Jump up ^ "NBC's 'Chuck': Good-natured, not much adult appeal September 29, 2008". Chicagotribune.com. Archived from the original on February 8, 2009. Retrieved 2012-11-01.
Jump up ^ "Michael Clarke Duncan's fiancee 'devastated' by his death". Retrieved September 4, 2012.
Jump up ^ "'All-Star Celebrity Apprentice': Omarosa Gets Emotional Over Chance To Honor Late Fiance Michael Clarke Duncan (VIDEO)". The Huffington Post. March 11, 2013.
Jump up ^ Saad, Nardine (2012-07-13). "'Green Mile' actor Michael Clarke Duncan in ICU after heart attack". latimes.com.
Jump up ^ "CPR: How It Helped Save Michael Clarke Duncan From Cardiac Arrest". The Huffington Post. Huffingtonpost.com. 16 July 2012. Archived from the original on 17 July 2012. Retrieved 16 July 2012.
Jump up ^ "ACTOR MICHAEL CLARKE DUNCAN REMAINS HOSPITALIZED". Associated Press. Retrieved 7 August 2012.
Jump up ^ "Michael Clarke Duncan dead at 54: ‘Green Mile’ actor dies nearly two months after suffering heart attack". Daily News. New York. Associated Press. 3 September 2012. Retrieved 3 September 2012.
External links
Wikimedia Commons has media related to Michael Clarke Duncan.
Michael Clarke Duncan on IMDb
Centerstage article on Michael Clarke Duncan
2011 radio interview with Michael Clarke Duncan
Michael Clarke Duncan at Find a Grave
[show]
Awards for Michael Clarke Duncan
[show]
v
t
e
Critics' Choice Movie Award for Best Supporting Actor
Kevin Spacey / Ed Harris (1995)
Cuba Gooding Jr. (1996)
Anthony Hopkins (1997)
Billy Bob Thornton (1998)
Michael Clarke Duncan (1999)
Joaquin Phoenix (2000)
Ben Kingsley (2001)
Chris Cooper (2002)
Tim Robbins (2003)
Thomas Haden Church (2004)
Paul Giamatti (2005)
Eddie Murphy (2006)
Javier Bardem (2007)
Heath Ledger (2008)
Christoph Waltz (2009)
Christian Bale (2010)
Christopher Plummer (2011)
Philip Seymour Hoffman (2012)
Jared Leto (2013)
J. K. Simmons (2014)
Sylvester Stallone (2015)
Mahershala Ali (2016)
[show]
v
t
e
Saturn Award for Best Supporting Actor
Marty Feldman (1974/75)
Jay Robinson (1976)
Alec Guinness (1977)
Burgess Meredith (1978)
Arte Johnson (1979)
Scatman Crothers (1980)
Burgess Meredith (1981)
Richard Lynch (1982)
John Lithgow (1983)
Tracey Walter (1984)
Roddy McDowall (1985)
Bill Paxton (1986)
Richard Dawson (1987)
Robert Loggia (1988)
Thomas F. Wilson (1989/90)
William Sadler (1991)
Robin Williams (1992)
Lance Henriksen (1993)
Gary Sinise (1994)
Brad Pitt (1995)
Brent Spiner (1996)
Vincent D'Onofrio (1997)
Ian McKellen (1998)
Michael Clarke Duncan (1999)
Willem Dafoe (2000)
Ian McKellen (2001)
Andy Serkis (2002)
Sean Astin (2003)
David Carradine (2004)
Mickey Rourke (2005)
Ben Affleck (2006)
Javier Bardem (2007)
Heath Ledger (2008)
Stephen Lang (2009)
Andrew Garfield (2010)
Andy Serkis (2011)
Clark Gregg (2012)
Ben Kingsley (2013)
Richard Armitage (2014)
Adam Driver (2015)
John Goodman (2016)
Authority control
WorldCat Identities
VIAF: 117917494
LCCN: no2001001757
ISNI: 0000 0001 1480 5609
GND: 142222402
SUDOC: 140543848
BNF: cb145090133 (data)
IATH: w6h1351g
Retrieved from "https://en.wikipedia.org/w/index.php?title=Michael_Clarke_Duncan&oldid=803529300"
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Chromosomal crossover
From Wikipedia, the free encyclopedia
Jump to: navigation, search
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.
Jump up ^ Esposito, M (September 1978). "Evidence that Spontaneous Mitotic Recombination Occurs at the Two-Strand Stage". Proceedings of the National Academy of Sciences of the USA. 75 (9): 4436–4440. doi:10.1073/pnas.75.9.4436.
Jump up ^ Kumar, A; Bassi, F; Paux, E (2012). "DNA repair and crossing over favor similar chromosome regions as discovered in radiation hybrid of Triticum". BMC Genomics. 13 (339). PMC 3443642  . PMID 22827734. doi:10.1186/1471-2164-13-339. Retrieved 14 March 2015.
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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