document_title: The Wizard of Oz on television

question_text: when was the wizard of oz first shown in color

question_tokens: when was the wizard of oz first shown in color

root

text: The Wizard of Oz on television

text: First telecast[edit]

text: The Wizard of Oz, which had been a critical but only modest financial success during its theatrical run, was chosen to be the first Hollywood film to be shown uncut in prime time on a coast to coast television network. [5]  The first telecast took place on Saturday November 3, 1956 as part of the final program in the canceled CBS anthology series Ford Star Jubilee . A rotating potpourri of highly budgeted but low rated specials including a well publicized debut hour hosted by Judy Garland.[6] The original asking price of $250,000 was negotiated by MGM attorney, later company president, Frank Rosenfelt. The network paid MGM $225,000 for the rights to televise the film twice with an option to re-broadcast it if the telecast was a success.[7]

tag: p

text: This 1956 broadcast shown as CBS's response to the successful color telecast of the Broadway musical Peter Pan with Mary Martin, which had been restaged especially for TV at NBC Studios as part of the anthology series Producers' Showcase. Peter Pan had first been shown live on TV by NBC in 1955, and been repeated (again live) by public demand in 1956. Its enormous success on television ushered in a temporary "fad" of mostly live family-oriented specials based on fantasy tales, such as Aladdin (1958, and no relation to the Disney film), Alice in Wonderland (1955) (a live-action version), Rodgers and Hammerstein's Cinderella (1957), The Pied Piper of Hamelin (1957), and Pinocchio (1957, no relation to the Disney film). As part of this trend, CBS bought the rights from MGM to telecast The Wizard of Oz.

text: For the first TV broadcast of The Wizard of Oz, the normally 90-minute Ford Star Jubilee was expanded to a full two hours to accommodate the entire film, which, in addition to having commercial breaks, was celebrity hosted. The main reason that CBS arranged for a host for the film was that at 101-minutes, the film was not considered long enough to run in the allotted 120-minute slot at that time, even with commercial breaks, without additional content to pad the entire telecast out to two hours.[8]

text: The 1956 television debut of the film marked the only time any actor who had appeared in the movie was selected to host it: Bert Lahr, who had played both the Cowardly Lion and farmhand Zeke in the film, appeared alongside then 10-year-old Liza Minnelli, and young Oz expert Justin G. Schiller. Lorna Luft, Minnelli's half-sister, did not appear as she was only four years old at the time, although she did have her picture taken with Minnelli in a promotional photo. Unlike several of the other Oz telecasts, apparently no stills were taken during the hosting sequences in 1956.[9] The practice of a show business celebrity regularly "hosting" The Wizard of Oz lasted from the film's first television showing until 1968, when the film went to NBC after being shown on CBS nine times.

text: The film was always presented uncut in a two-hour time slot between 1956 and 1968, despite having commercials and hosted segments. The Professor Marvel sequence has never been omitted (at least in American telecasts of the film),[citation needed] and the tracking shot of Munchkinland was not edited until the film went to NBC in 1968.[citation needed]

text: The film's first broadcast received positive critical notices in newspapers the following day. Variety magazine prophetically suggested that the film could be telecast annually and at an earlier time.[10]

text: Telecasts in the Pre-Cable Era[edit]

text: The film was not rebroadcast in 1957 or 1958. For telecasts from 1959 (the year of its second telecast) up until 1998, the film was always shown as a stand-alone TV special instead of as part of an anthology or movie series. Between 1959 and 1968, and again in 1990 when Angela Lansbury, star of CBS' Murder She Wrote, hosted the 50th-anniversary telecast, CBS chose its hosts from its then-current prime time lineup. In 1959, the host was Red Skelton (The Red Skelton Show); in 1960 it was Richard Boone (Have Gun, Will Travel), in 1961 and 1962 it was Dick Van Dyke (The Dick Van Dyke Show), and from 1964 through 1967, it was Danny Kaye (The Danny Kaye Show). Skelton, Boone and Van Dyke brought their then-young children along to appear with them.

text: Although the hosting segments for the 1956 telecast of the film had to be done live, Skelton's, Boone's, Van Dyke's and Kaye's segments were recorded on video tape in advance of the telecasts.[11] Lansbury's hosting segment was the first one recorded on film.

text: During these years, the hosting sequences were not staged in a run-of-the-mill manner, with the host merely standing in a studio set, but in what could be considered imaginative ways. Richard Boone was taped on the set of his television series Have Gun, Will Travel, in a "living room" with his real son about to watch the film on a TV set. Dick Van Dyke was shown in a living room set with his children,[11] and Danny Kaye's hosting segment featured him sitting on a prop toadstool against a painted backdrop of the Yellow Brick Road and the Emerald City.[12] The Kaye segment did not include a television set as part of its design.

text: Some humor as sometimes incorporated into these segments. Red Skelton was seen as two characters: Before the film began, he was seen in a studio set of an early 20th-century bookstore, in costume as a Victorian-era storyteller who introduced L. Frank Baum's original 1900 novel The Wonderful Wizard of Oz to a young girl played by Skelton's real daughter, and at film's end, he appeared in a studio recreation of a modern living room as himself. Danny Kaye reassured viewers not to panic when the film began in black-and-white rather than in color, and encouraged young viewers not to be afraid of the roaring MGM lion at the beginning of the film. No directors or writers were credited for the hosting sequences, just as none are credited for hosting segments prior to films on Turner Classic Movies.

text: The Wizard of Oz did not become an annual television tradition immediately — only after the 1959 showing, when, because of the earlier hour at which it was shown (6:00 P.M., E.S.T.), more children tuned into the broadcast, and it gained an even larger television audience than before. The 1959 telecast was especially welcomed by media critic John Crosby, who commented in the New York Herald Tribune, "Television — any television — looks awfully ordinary after The Wizard of Oz".[13] From 1959 until 1991, the film was telecast once every year, the one exception being 1963, when it was not telecast at all.

text: WISN-TV in Milwaukee, Wisconsin, did not carry the network's yearly Oz telecast in 1961, the year WISN began its affiliation with CBS, running Green Bay Packers football instead. However, due to viewer outcry, WISN was able to get permission to run the film locally at 2:00 p.m. C.S.T., on Christmas Eve.[14]

text: In 1978, after the film had returned to CBS, a computer malfunction at the CBS owned-and-operated WBBM-TV in Chicago accidentally cut off most of the ending to that year's Oz telecast, interrupting the final minute with a commercial block that was not supposed to air until after the movie had ended.[citation needed] Because the break was only 42 seconds long, no attempt was made to override the computer, for fear of making the problem worse.[citation needed] For several hours thereafter, the WBBM received angry calls from viewers, while those unable to get through chose to voice their displeasure wrote to the Chicago Tribune and Chicago Sun-Times newspapers.[citation needed]

text: "Wraparound" opening and closing credits[edit]

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tag: table

text: The film, as telecast on U.S. television between 1959 and 1968, was arguably given a much more elaborate TV presentation than it received afterwards. During those years, it always had videotaped wraparound opening and closing credits segments devised by CBS, accompanied by the network's own specially recorded opening and closing music based on the film's score. For the opening "wraparound" credits, the title The Wizard of Oz and the names of its five leading actors, Judy Garland, Frank Morgan, Ray Bolger, Bert Lahr, and Jack Haley, were first shown in CBS's own format and font, while an anonymous announcer read them off and then followed this with an announcement of the film's sponsor(s): "This portion of The Wizard of Oz is brought to you by...[name of sponsor mentioned]". These specially-devised opening credits never mentioned that the film was made by Metro-Goldwyn-Mayer. From 1959 to 1964, CBS created different wraparound credits for each showing, but because the same hosting segment — the Danny Kaye one — was shown between 1964 and 1968, audiences saw the same "wraparound" credits from 1964 until the film went to NBC.

text: The special CBS introduction was followed by the host speaking about the movie for about three minutes or so. Those remarks lead into the actual film, beginning with all of its original 1939 opening credits (shown against a background of moving clouds), including the MGM Leo the Lion logo, the name of the film, the cast list, and the film's principal technical staff, exactly as MGM had created them, with the film's main title music heard.

text: The host reappeared just before the film's second half began, to say a few more words about it, before the telecast proceeded with the rest of the film, commercials included.

text: However, at the end of the movie, the film's closing cast list, as created by MGM, was not be shown, and the title card with "The End" right after Dorothy's closing line was never seen on television during these early CBS showings. Instead, immediately after Dorothy spoke her last line ("Oh, Auntie Em, there's no place like home!"), and the camera faded out on her, television viewers once again saw CBS's own title card The Wizard of Oz, this time accompanied by some of the film's end title music, exactly as heard on the soundtrack, and the announcer's voice saying that the host for that year would return in a moment. After a final commercial, the host was then seen once again, to bid farewell to the TV audience, and CBS showed its own version of the cast list that appears during the film's end credits. At the end, referring to both the film and the hosting segments, the same announcer declared that "This has been a film and videotape presentation."

text: Shown in color[edit]

text: From the beginning The Wizard of Oz was telecast in color, although few people owned color television sets in 1956.  Except for 1961, all U.S. telecasts have been in color, an effect that seemed much more striking in the early 1960s, when there were still relatively few color programs on television. It was not televised in color in 1961 because color telecasts had to be paid for by their sponsors, who declined to do so that year.[15][16] Between 1956 and 1965, the Wizard of Oz showings were rare exceptions to the black and white program schedule at CBS. During this period, CBS had the ability to broadcast programs in color, but generally chose not to do so unless a sponsor paid for a film or program to be shown in color. During this period, the competing network NBC was owned by RCA, which by 1960 manufactured 95% of the color sets sold in the U.S. Hence, CBS perceived that increased use of color broadcasting would primarily benefit its rival by promoting sales of RCA color television sets.

text: Not until the fall of 1965 did color broadcasts play a major role at CBS and ABC, at which time half the network shows from ABC and CBS were being made and shown in color. Meanwhile, all but two NBC prime time shows were in color and most of NBC’s daytime shows were. By the fall of 1966, all three networks produced all of their prime time shows in color. By the time the movie went to NBC in 1968, all network shows (except for reruns of black and white movies) were in color.

text: Later hosts[edit]

text: This section, except for one footnote, needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (June 2013) (Learn how and when to remove this template message)

text: Partly because commercial time during programs increased beginning in the late 1960s, the idea of regularly having hosts to introduce the film was dropped when The Wizard of Oz went to NBC in 1968, where no "wraparound" sequence was shown. The presentation consisted only of the film itself, with its original opening and closing credits, and no special NBC-created credits or hosting segments. The famous NBC peacock was shown immediately prior to the beginning of the film, with announcer Mel Brandt saying that "the first 22 minutes of this program [i.e. the Kansas and tornado sequences] will be shown in black-and-white", a not quite accurate statement, since the final three minutes of the film also took place in Kansas, and were at that time also shown in black-and-white, rather than in the sepia tone in which they originally had been made (the sepia was not restored to the Kansas and tornado scenes until 1989, the film's 50th anniversary). However, one NBC telecast featured an on-screen host: the 1970 showing, which opened with veteran actor Gregory Peck paying tribute to the recently deceased Judy Garland (a segment directed by Oz producer Mervyn LeRoy, marking his first TV work), although this segment consisted of only a few brief remarks, while the opening hosting segments on CBS had gone on for about three minutes or so. The NBC Oz telecasts began the tradition of the film's annual showings during the Easter and Passover seasons of the year as opposed to the winter showings on CBS.

text: The switch in networks resulted because CBS was unwilling to meet MGM's increased price — fostered by the film's ever increasing popularity — for renewal of the rights to telecast it.[17]  The film stayed on NBC until 1976. When CBS, realizing its error in allowing it to go to another network, bought back the rights at MGM's asking price, their viewer ratings shot up, and one executive was heard to remark, "That picture is better [for the network] than a gushing oil well".[18]

text: After its 1976 return to CBS, the film was hosted on that network only once more, in a filmed segment featuring Angela Lansbury in 1990, but the CBS "wraparound" opening and closing credits were not - and have never been - revived, although, during those years, a blue card featuring a painting of a rainbow and the title The Wizard of Oz was shown on the screen while the night's pre-empted programs or programs to be shown at regular time and the sponsors were being announced, and immediately before and after commercial breaks. In the 1980s clips from the film shown on a red background with title was shown at the start, while a still of Emerald City with title was used during commercials. Angela Lansbury also narrated a documentary about the making of the film, originally entitled The Wonderful Wizard of Oz: 50 Years of Magic and years later retitled The Wonderful Wizard of Oz: The Making of a Movie Classic. It was first shown immediately after the movie's 1990 telecast, and is included as a supplement on all the DVD releases beginning with the 1999 DVD release. Jack Haley, Jr., the documentary's director, was nominated for an Emmy Award for his work.

text: When shown on Turner Classic Movies, The Wizard of Oz was usually hosted by TCM host Robert Osborne.

text: On June 3, 2007, Tom Kenny, the voice of SpongeBob SquarePants, hosted a telecast of the film on Turner Classic Movies, as part of Essentials, Jr., a special summer series of family movies.

text: On July 27, 2008, the film was shown twice in a row on Turner Network Television without a host, but with commercials, and with "pop-up" animated ads for other TNT programs at the bottom of the screen just before and after commercial breaks.

text: On March 24 and 25, 2012, the film was hosted on the Cartoon Network by television actor Robert Wu, who provides the voice of Mr. Washee Washee in the Family Guy episode, "Tiegs for Two". This telecast ran two-and-a-half hours, longer than any CBS or NBC telecasts.

text: On June 10, 2012, which would have been Judy Garland's 90th birthday, the film was telecast on Turner Classic Movies without commercials, and hosted by comedian Bill Hader, again as part of the Essentials, Jr. summer series of family films. Unlike the CBS hosting segments which were a part of the 1960s telecasts of the film, Hader's segment showed clips from the film before it actually began.

text: Sponsorship[edit]

text: Being a presentation of network TV, "The Wizard of Oz" was subject to sponsorship from a variety of advertisers.

text: The first telecast of "Oz" was a presentation of Ford Star Jubilee and thus sponsored by Ford Motor Company and its dealers. In the ensuing years of 1959 thru 1967, the yearly telecast was sponsored by one primary sponsor with a co-sponsor. Perhaps the best remembered examples were the telecasts sponsored by Procter and Gamble and promoted when the company gave free premiums of hand puppets with packages of some of its most popular products. Many also remember the 1970 telecast presented on NBC by Singer Manufacturing Company as a tribute to Judy Garland, who had died in June 1969.

text: In later years, several different advertisers shared sponsorship; no one advertiser predominated.

text: Television ratings[edit]

text: The showing in 1983 was the 25th network prime-time showing, a record then for any film or television special.  In the first nine showings, all on CBS, The Wizard of Oz gained at least 49% of the television audience.[19] In 1966, it ranked No. 1 in the ratings for the week that it was shown.[20] Between 1960 and 1968, the film even beat out episodes of ABC-TV's Walt Disney Presents (in 1960) and NBC's Walt Disney's Wonderful World of Color (from 1961 to 1968), which aired opposite the film.[21]  When the film moved temporarily from CBS to NBC, it always preempted the Disney program altogether, except for once, when NBC showed Oz on a Saturday in 1968. When CBS bought the film back from NBC in 1976, it again began to beat episodes of Disney in the ratings. It preempted Disney on one more occasion, after the series moved to that network in the early 1980s.[22]

text: In 2006, the first year it was shown on TV in high-definition, the film placed No. 11 in the Nielsen ratings among cable television programs for the week of November 11.[23]

text: Changes made in running time[edit]

text: From 1968 to 1984, minor cuts were made to the film to make room for added commercial time and in order to have the film "clock in" at two hours. No dialogue or singing was removed, only moments such as camera pans and establishing shots, as well as MGM's written foreword to the film.[11]

text: On a few occasions beginning in 1985, again because of the increased time spent on commercial breaks, the film was time-compressed to fit it into a two-hour running time without cutting it.[9] However, The Wizard of Oz is now always shown complete and at its regular speed on television, both with and without commercials. When shown with ads, the film now runs about two hours and fifteen minutes, simply because of the increase in commercial time.

text: March 1991 showing[edit]

text: The March 1991 showing was the first after the film gained protected status from the Library of Congress and the National Film Preservation Board. Networks opted to no longer shorten the film by "microcutting" a few individual moments throughout the movie as had been done from the late 1960s to the early 1980s to make room for commercials and keep it in a two-hour broadcast. This extended the running time of the film from 8 P.M to 10:07 p.m. and sometimes even longer, depending on the amount of time spent on commercials. It was one of the first 50 films selected for this protection.[24]

text: Move to cable[edit]

text: In the 1980s, Ted Turner purchased the film from MGM, alongside Gone with the Wind. Because both films were still licensed to CBS, Turner and the network negotiated a deal that extended CBS' license for Oz in exchange for relinquishing its rights to Gone with the Wind.[25] In 1991, the film was shown twice during the year for the first time.[citation needed] 1991 also marked the first time since 1956 that the film was shown in November. This also happened in 1993, when the film was telecast in both February and November of that year. The film was not shown on television at all in 1992, 1995 and 1997, marking the first time since 1963 that a year was skipped in showings of the film. Turner, which owned most of the pre-May 1986 MGM film and television library at the time (later owned by Warner Bros.), began moving to make its properties exclusive to Turner-owned outlets in the late 1990s; as such, in 1998, The Wizard of Oz made its last appearance on CBS, moving exclusively to Turner-owned properties the next year.[26]

text: 2000 marked the first time that the film was shown on U.S. television during the summer. In 2002, it was shown five times.[citation needed]

text: On November 6, 2011, TBS became the American television channel on which The Wizard of Oz has been shown most often, when the film had its 32nd showing on that channel, finally breaking CBS' record 31 showings. As of April 29, 2013, Turner Classic Movies has shown the film 23 times.

text: Differences between network and cable showings[edit]

text: In addition to the frequent cable showings, another difference between showings on NBC, CBS, the WB network, and cable channels is that when the film was shown on CBS and NBC, it was always presented as a special instead of just a televised film. From 1959 until it went to cable, the film was never shown on NBC Saturday Night at the Movies or any other movie anthology series, and telecasts of the film regularly preempted two hours or more of regular television programming.[11]

text: Promos for the CBS and NBC showings during the 1960s began airing on television as much as two weeks in advance of that year's telecast and were still airing as late as 1989.[citation needed]  On the major commercial networks, it was never termed a "CBS Movie Special" or an "NBC Movie Special", as movie specials shown on those networks are frequently termed, but as simply The Wizard of Oz.[citation needed]

text: Aspect ratio[edit]

text: The 1955 and 1998 theatrical re-releases were matted in movie theaters to produce a widescreen effect for the Academy-standard aspect-ratio film.

text: When shown in HDTV, the film is pillarboxed so that its aspect ratio is preserved.[27] On DVD, the film has always been issued in its original 1939 aspect ratio.[28][28][29]

text: Outside the United States[edit]

text: The movie has also been shown on television successfully in the United Kingdom, Canada and is shown every year in Australia, but it has not become the television phenomenon there that it has in the U.S.[30] In Indonesia, the film was aired for the first time on the newly-inaugurated television network RCTI in November 1989, with reruns on other four networks in the 1990s. In Japan, the film has aired on the Japanese version of The Magical World of Disney, although Disney had nothing to do with its production or release.[31]

text: Broadcast Television Airdates[edit]

text: Eastern Time (taken from TV Guide and from The Wizard of Oz: The Official 50th Anniversary Pictorial History).

text: 1956 - CBS - Saturday, November 3 - 9:00 p.m.

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text: 1959 - CBS - Sunday, December 13 - 6:00 p.m.

text:
1960 - CBS - Sunday, December 11 - 6:00 p.m.
1961 - CBS - Sunday, December 10 - 6:00 p.m. Only time fully shown in Black & White
1962 - CBS - Sunday, December 9 - 6:00 p.m.
1964 - CBS - Sunday, January 26 - 6:00 p.m.

text: The reason that the film did not air in December 1963 has never been stated, but some say that it was because of John F. Kennedy's assassination, others say room needed to be made for more Christmas themed specials and that the movie fit in better during January. For whatever reason, the movie continued to air early in the year rather than late in the year from 1964 until the 1990s.

text:
1965 - CBS - Sunday, January 17 - 6:00 p.m.
1966 - CBS - Sunday, January 9 - 6:00 p.m.
1967 - CBS - Sunday, February 12 - 6:00 p.m.

text: Last airing on CBS until 1976. Moved to NBC in 1968, at this time, the showings were moved from midwinter to late winter/early spring.

text: 1968 - NBC - Saturday, April 20 - 7:00 p.m.

text: First showing by NBC, in Spring rather than Winter. The first time that the film was shown at 7:00 P.M instead of 6:00 P.M.

text:
1969 - NBC - Sunday, March 9 - 7:00 p.m.
1970 - NBC - Sunday, March 15 - 7:00 p.m.
1971 - NBC - Sunday, April 18 - 7:00 p.m.
1972 - NBC - Tuesday, March 7 - 7:00 p.m.

text: The first time that the film aired in the middle of the week, rather than on a weekend.

text:
1973 - NBC - Sunday, April 8 - 7:00
1974 - NBC - Sunday, February 24 7:00 p.m. Due to a breaking news story, NBC decided to show the film two weeks later, on March 10, rather than show it too late or in progress.
1975 - NBC - Easter Sunday, March 30 - 7:00 p.m.
1976 - CBS - Sunday, March 14 - 7 p.m.

text: The year that The Wizard Of Oz returned to CBS. It remained there for twenty-two more years, a likely record in those years for a film's consecutive showings on one network. Initially the movie aired late winter/early spring, but beginning in 1991, the movie moved back and forth from spring to late fall. Another change was that CBS no longer began the movie before normal prime time. Before 1976, when run on a day other than Sunday, NBC took back the 7 p.m. Eastern / 6 p.m. Central time slots from affiliates to run the movie early enough so children could see it before bedtime. Also, prior to 1968, CBS always took the 6 p.m. hour to run the movie early. Logic was that times have changed and children now stay up a little later than they did in the past. From then on, if shown in a Sunday 6 to 8 or 7 to 9 pm time slot on CBS, the film preempted 60 Minutes, so beginning in 1978, CBS executives moved the showings to other nights of the week. If the film was shown on a Sunday, CBS usually scheduled it at 8 pm, so that it did not conflict with 60 Minutes. (As of 2017, 60 Minutes continues to air in the same Sunday 7 p.m. time slot on CBS.)

text:
1977 - CBS - Sunday, March 20 - 7 p.m.
1978 - CBS - Easter Sunday, March 26 - 7 p.m.
1979 - CBS - Friday, March 23 - 8 p.m.
1980 - CBS - Friday, March 7 - 8 p.m.
1981 - CBS - Friday, February 27 - 8 p.m.
1982 - CBS - Saturday, March 27 - 8 p.m.
1983 - CBS - Friday, March 18 - 8 p.m.
1984 - CBS - Friday, March 30 - 8 p.m.
1985 - CBS - Friday, March 1 - 8 p.m.
1986 - CBS - Saturday, February 15 - 8 p.m.
1987 - CBS - Friday, March 6 - 8 p.m.
1988 - CBS - Wednesday, February 24 - 8:30 p.m.

text: Delayed due to a CBS News Special report.

text: 1989 - CBS - Sunday, March 18 - 7 p.m.

text: The last time that the film pre-empted 60 Minutes.

text:
1990 - CBS - Tuesday, February 20 - 8 p.m.
1991 - CBS - Tuesday, March 19 - 8 p.m.

text: Soundtrack remastered in stereo using CBS StereoSound system.

text: 1991 - CBS - Wednesday, November 27 - 8 p.m.

text: The first time the film aired twice in the same year. This was done to switch it to a November date.

text:
1993 - CBS - Friday, February 26 - 8 p.m.
1993 - CBS - Wednesday, November 11 - 8 p.m.
1994 - CBS - Thursday, November 23 - 8 p.m.
1996 - CBS - Friday, May 10 - 8 p.m.

text: This was the same day that the film Twister, which contains plenty of references to The Wizard of Oz, was released to theatres. The first time that the film aired close to summertime.

text:
1998 - CBS - Friday, May 8 - 8 p.m. - This was the last airing on CBS. From then on, Turner Broadcasting was be in charge of telecasting the film.
2002 - WB - Sunday, November 24 - 7 p.m. First broadcast TV airing since 1998.
2003 - WB - Sunday, December 7 - 7 p.m.
2004 - WB - Sunday, December 19 - 7 p.m.
2005 - WB - Sunday, December 18 - 7 p.m.

text:
Cable showings
November 21, 1999, TBS
July 3, 2000, TCM

text:
November 19 and 25, 2000, TNT
July 4, 2001, TCM
December 1 and 2, 2001, TNT
July 4, 2002, TCM
November 28, 2002, TBS
December 8, 13, and 25, 2002, TNT
July 5 and 6, 2003, TCM
November 16 and 21, 2003, TBS
December 13 and 14, 2003, TNT
July 2 and 3, 2004, TCM
December 8 and 12, 2004, TNT
November 19, 20, and 24, 2004, TBS
July 3 and 4, 2005, TCM
November 11, 12, and 13, 2005, TBS
July 3 and 4, 2006, TCM
November 10, 11, and 12, 2006, TBS
December 11 and 17, 2006, TNT
June 3, 2007, TCM
November 9, 10, 11, 2007, TBS
November 9 and 25, 2007, TNT
December 22 and 23, 2007, TNT
March 21 and 22, 2008, TCM
July 27, 2008, TNT
November 14, 2008, TNT
November 14-16, 2008, TBS
December 20 and 21, 2008, TNT


document_title: The Good Dinosaur

question_text: what kind of dinosaurs are in the good dinosaur

text: The Good Dinosaur

text: Plot[edit]

text: In an alternate history, the asteroid that would have caused the extinction of the dinosaurs 65 million years ago passes safely over Earth.

text: Sixty-five million years later, Apatosaurus farmers Henry and Ida have children Libby, Buck, and the runt Arlo, who has trouble adjusting to farm life. While his successful siblings are allowed to "make their mark" (a mud-print on the family's corn silo), Arlo's timid nature makes tasks difficult for him. Henry attempts to give Arlo a sense of purpose by putting him in charge of guarding their silo, and helps him set a trap. It captures a feral caveboy, but Arlo doesn't have the heart to kill him, and sets him free. Disappointed, Henry takes Arlo to track the caveboy, leading them into a ravine. Henry saves Arlo from a flash flood before being swept away and killed.

text: Without his father, Arlo shoulders more of the workload. He spots the same caveboy inside the silo and, blaming him for his father's death, chases him until both fall into a river. Arlo cannot swim, and is swept downstream where he hits his head on a rock and is knocked unconscious. Awakening, he finds himself far from home and tries to survive on his own, but becomes trapped when a boulder pins his leg. The next day Arlo wakes to find his leg has been freed, and the caveboy appears with food for him. The caveboy then leads Arlo to a berry tree, where the caveboy fends off a large snake, amazing Arlo and impressing a nearby eccentric Styracosaurus who wants to keep the boy. He forces Arlo to compete with him to give the boy a name he will respond to, which Arlo finally wins when he calls him "Spot". Arlo and Spot bond as Arlo laments his lost family, and Spot reveals that his own parents are dead. Later, when a storm strikes, Arlo runs away in fear and loses the riverbank he has been following home.

text: The next morning, Arlo wakes to find Spot at his side. They are noticed by a band of pterodactyls, led by Thunderclap, who appear to be conducting a rescue operation but turn out to be savagely carnivorous. When the pterodactyls try to take Spot, Arlo and Spot flee, happening upon a pair of Tyrannosaurus named Nash and Ramsey, who ward off the pterodactyls. Nash, Ramsey, and their father Butch have lost their herd of longhorns, so Arlo offers Spot's help in sniffing them out. They locate the herd, but Butch recognizes the work of cattle rustlers, and uses Arlo as a lure. Arlo and Spot attract the attention of rustler Velociraptor, allowing Butch and his family to attack. During the fight, Arlo musters his courage and fends off two raptors who have overwhelmed Butch, helping to turn the fight in their favor. Having gained their respect, Arlo joins the Tyrannosaurus in driving the cattle south when he sees the familiar mountain peaks of his homeland in the distance, and leaves with Spot to return home. Along the way, they encounter an adult feral caveman in the distance, and though Spot shows interest, Arlo dissuades him and they continue on.

text: As another storm approaches, Thunderclap and the pterodactyls return and attack and carry Spot away. Arlo becomes entangled in vines, where he has a vision of Henry leading him home. Arlo instead resolves to save Spot, making the vision of his father proud before it fades away. Arlo finds and attacks the pterodactyls, who have cornered Spot at the river. Arlo and Spot together plunge Thunderclap and the pterodactyls into the water, where they are swept helplessly downstream. When another flash flood occurs, Arlo leaps into the water to rescue Spot as the two are swept away toward a waterfall. Arlo protects Spot as the two plummet down the fall, and carries him to shore.

text: As they approach Arlo's home, the two again hear the unknown caveman call, and are approached by an entire cave family. With great reluctance, Arlo pushes Spot to join his adoptive family, and the two of them share a tearful goodbye.

text: Arlo finally arrives home to his mother and siblings, and makes his mark on the silo between those of his mother and father.

text: Voice cast[edit]

text:
Raymond Ochoa as Arlo, the youngest of the Apatosaurus family[10][11]

Jack McGraw as Young Arlo

Jack Bright as Spot, a 6-7-year-old human caveboy who befriends Arlo.[10][11]
Sam Elliott as Butch, a Tyrannosaurus who runs a "ranch" filled with prehistoric "longhorns".[10]
Anna Paquin as Ramsey, Butch's daughter[10][11]
A. J. Buckley as Nash, Butch's son[10][11]
Jeffrey Wright as Poppa Henry, the father of the Apatosaurus family.[10][11]
Frances McDormand as Momma Ida, the mother of the Apatosaurus family.[10][11]
Marcus Scribner as Buck, Arlo's brother[10]

Ryan Teeple as Young Buck[12]

Maleah Padilla as Libby, Arlo's sister[12]
Steve Zahn as Thunderclap, a Nyctosaurus and the leader of a gang of pterodactyls.[10]
Mandy Freund as Downpour, a pterodactyl who is a member of Thunderclap's flock[12]
Steven Clay Hunter as Coldfront, a pterodactyl who is a member of Thunderclap's flock[12]
Peter Sohn as Forrest Woodbush, a Styracosaurus who has various animals perched on his horns.[12][13]
Dave Boat as Bubbha, the leader of a gang of Velociraptor rustlers[12]
Carrie Paff as Lurleane, a Velociraptor who is a member of Bubbha's pack.[12]
John Ratzenberger as Earl, a Velociraptor who is a member of Bubbha's pack.[12]
Calum Mackenzie Grant as Pervis, a Velociraptor who is a member of Bubbha's pack.[12]

text: Production[edit]

text: Bob Peterson and Peter Sohn started working on the film in 2009, when Peterson came up with the idea.[14] The film's first release date of November 27, 2013 was first announced in June 2011.[15] The plot, director and co-director, producer, and other small details were announced at the D23 Expo on August 20, 2011.[16] Peterson and John Walker announced the film as The Untitled Pixar Movie About Dinosaurs.[16] On April 24, 2012, Pixar announced that it would be titled The Good Dinosaur.[17]

text: The filmmakers wanted to explore what dinosaurs represent today, and how they are represented in stereotypes. Peterson stated: "It's time to do a movie where you get to know the dinosaur, what it's really like to be a dinosaur and to be with a dinosaur."[18] Peterson said the inspiration for the project came from a childhood visit to the 1964 New York World's Fair where he was impressed by some dinosaur Audio-Animatronics.[18]

text: On the film's title, Sohn stated, "The title is deceptively simple. It has more meaning than it seems."[18] He additionally explained: "Arlo has a lot of issues when he's born. He's fearful and he's weak and he's disconnected from the family because of these issues and he feels like he's not worthy, and so he finds a way to become worthy."[19]

text: In April 2012, Pixar announced that the film's release date had been shifted from November 27, 2013 to May 30, 2014.[17][20] On August 9, 2013, it was announced at the D23 Expo that Lucas Neff, John Lithgow, Frances McDormand, Neil Patrick Harris, Judy Greer, and Bill Hader had joined the cast of the film.[21]

text: Revisions[edit]

text: By summer 2013, Peterson and Walker had been removed from the film due to story problems.[1] Peterson, who could not crack the film's third act,[22] was absent from the D23 Expo, where Sohn and producer Denise Ream presented footage from the film.[23][24] Ed Catmull, Pixar's president, explained the departure: "All directors get really deep in their film. Sometimes you just need a different perspective to get the idea out. Sometimes directors… are so deeply embedded in their ideas it actually takes someone else to finish it up."[1] Peterson moved on to another project he is developing at Pixar, while Ream replaced Walker, who left to work on Disney's Tomorrowland. On the subject of changing directors, Ream stated, "Sadly, it happens more often than any of us would like… It's given a lot of thought and care. Bob still works at Pixar [and] he's a beloved member of the studio. He helps out on a lot of the movies. We care about him, but he just needed help getting the movie done."[25] John Lasseter, Lee Unkrich, Mark Andrews, and Sohn stepped in temporarily to work on various sections of the film.[1] In September 2013, The Good Dinosaur was pushed back from May 30, 2014 to November 25, 2015 (the scheduled release date for Pixar's Finding Dory). According to Ream, the primary reason for the rescheduling was "the story was not working, period, full stop, it just was not where it needed to be."[26] In November 2013, due to the delay, Pixar laid off 67 employees of its 1,200-person workforce,[22] following the closure of Pixar Canada a month before, when about 80 employees had been laid off, officially to refocus Pixar's efforts at its main headquarters.[27]

text: In the footage shown at the 2013 D23 Expo, Arlo was a huge dinosaur, Sohn recalled: "In the original version of the film, Spot was so tiny, we really wanted to make him feel like a bug. But people had a tough time connecting that these two had a relationship, because Arlo was so large. It was fun, but at the same time we never believed the growth that they could have together. It was really like, 'I’m best friends with a bee!' Bob Peterson, the original director, his first pitch was just a boy and his dog, and so when I took it I said, 'I want to honor that as much as I can.'"[28]

text: In August 2014, Lithgow revealed in an interview that the film had been dismantled and "completely reimagined" and that he was expected to re-record his role in the next month, while mentioning that McDormand was still part of the film.[29] In November 2014, it was reported that new elements had been added to the story, such as treating nature as the film's antagonist.[30] Pixar's general manager and president Jim Morris stated, "We all liked the movie from the beginning, but… the story had some kind of fatal flaws in it that none of us really saw. It was kind of a Billy Elliot story. It was a very different kind of movie—there was a whole dinosaur culture and community and so forth. It was one where the Arlo character was kind of repressed by the community. It never quite felt right. It always felt like that we were vilifying this group [of dinosaur characters] that was otherwise a very likeable group. It just didn't feel like it resonated truly."[31] Morris additionally explained: "We give people the chance to work on a story, and we give them our support, but it doesn't always work out… there were story issues that were just not solvable. So we changed the story to make it better, and we're very proud of the film we have now."[32] In October 2014, Sohn was announced as the new director of the film.[33]

text: In June 2015, it was announced that the majority of the cast had been revised.[10] Of the original cast, only McDormand retained her role in the film. It was revealed that Neff had been replaced by Raymond Ochoa, and Lithgow had been replaced by Jeffrey Wright.[10] Arlo's three siblings, to be voiced by Harris, Hader, and Greer had been cut down to a single brother named Buck, voiced by Marcus Scribner,[10] and later, a sister named Libby, voiced by Maleah Padilla. On the cast revisions, Sohn explained: "It was all about finding a younger Arlo, it was really about finding a boy [to play Arlo], so that we could push into that idea of him growing up and becoming a man, so the actor previous—who is a great actor—he was already a man, and so I needed to push that arc and find that compassionate kid, so that was the major kind of change. Then everything else, all the other characters that supported that story came in and out and changed and evolved and through that evolution, some of those performers changed out of it."[19] It was also confirmed that the farmer aspect was still part of the film.[34]

text: Design and setting[edit]

text: The filmmakers wanted nature to be the antagonist for the main character Arlo. Ream noted "Nature can overcome anything, including a massive dinosaur." In order to achieve the needed realism, the film's team traveled to the American Northwest, spending time in Jackson Hole, Wyoming, Juntura, Oregon, and southern Montana. Production designer Harley Jessup stated: "The area has a fantastic variety of landscapes, ranging from the Jackson Valley and the Tetons to the amazing geysers and waterfalls in Yellowstone, we studied the grasslands of Montana and the Red Desert, then incorporated all of it in Arlo’s journey." To use the landscapes that they had experienced, the filmmakers used data from the U.S. Geological Survey, and satellite images from Google Earth. The geographical data provided a foundation that the team then built on. According to supervising technical director Sanjay Bakshi, this gave Sohn "the freedom to shoot in any direction he wanted to make the world feel big and real." In addition, The Good Dinosaur features three-dimensional, volumetric clouds. In previous Pixar films, clouds have been "painted" onto the sets. Light and photography director Sharon Calahan explained: "Storm clouds are almost like a villain in the film. Plus, it’s very labor-intensive to paint clouds, and they’re in almost every scene." Calahan also noted "These particular clouds can be rendered and we can light them, which we’ve never been able to properly do before."[35][36]

text: According to the filmmakers, the environments and landscapes in the film are not photo-realistic, they are just detailed in a way that advanced technology and style decisions allow. In terms of animating Arlo, animators Rob Thompson and Kevin O’Hara went to a zoo and shot video of elephants in motion. Thompson stated: "One of the most intimidating things to animate is a quadruped, because there’s so much to them and there’s so much to manage. It’s very easy to make it not work if you don’t know what you’re managing. Locomotion is all about efficiency, a lot of times you think, 'We’re animating a big, heavy character. We should slam those feet. That’ll make it feel heavy.' The truth is, that’s not efficient. That wouldn’t be great for the legs."[35][36][37] A system where Arlo's head goes up and his chest goes down when his hips goes up, was therefore created. To get an idea of the scale of Arlo, a complete full-size model was built out of card and foam core. In total The Good Dinosaur took up 300TB of server space, ten times as much space as Monsters University (2013).[38]

text: Arlo was designed so that the audience could identify with him and be able to see the "boy" inside the dinosaur. Sohn explained "When Arlo gets lost in the wilderness, you need to worry that he's out of his element and in danger of getting killed, rather than just thinking 'You're an animal. Why don’t you just turn around and eat some leaves?'" He additionally stated "If you put like a realistic dinosaur in the woods, eat some leaves, you’re fine… But we really wanted to play with that twist of this is a boy out there. A dinosaur boy for sure, but trying to infuse those qualities into him… And also keeping him in a world that had an arc to itself. That it would start off being beautiful and then become dangerous. When we had these designs of the trees where they were a little bit more blocky and a little bit more graphic. We put Arlo in front of it, it’s like 'ooh that kind of is cool, but boy, it doesn’t feel like it's that dangerous.' Like when we get a rock stuck on him, it didn’t feel like anything ’cause when we went on our research trips, boy that idea of like you can get, you could die by a small thing like having a rock on your foot. Or you could die by a huge thing like an avalanche, which we tried to kind of explore in the film. So that was one of our main intents."[25][39]

text: In terms of the film's setting, Sohn stated "Pixar is all about the 'what if' question… 'What if the asteroid missed the earth?' Yeah, we went down the road of: they could be cooking in a dino diner; they could be in space; they could be in anything. But then there was a drawing of a dinosaur farming that kind of sparked people of the idea of 'he’s working hard and this character needs to survive out there.' All of the American dinosaur bones are found in the northwest—from Montana, to Oregon, to Wyoming, and the Dakotas, so the combination of hardworking farm families and the west really brought us into that frontier world immediately… [S]o we started doing research. We started finding all of these people who were surviving out there on the land. They were either cultivating the cattle, or were ranchers in Oregon, or were farmers in the Midwest. It just kept teaching us how much we want to honor this world, and how hard it is to survive out there. It started dictating the world itself."[28]

text: Since The Good Dinosaur is set in a world in which dinosaurs never became extinct, they were instead able to evolve. Herbivores like Arlo and his family become farmers, and carnivores like the T. rex become ranchers. Because they are meant to be reminiscent of cowboys, when the T. rex run, their lower bodies mimic a galloping horse, while their upper bodies have the feel of a riding cowboy. To help inspire Butch's physical look and performance, the filmmakers looked at classic film cowboys such as characters portrayed by Clint Eastwood and Jack Palance.[37]

text: Soundtrack[edit]

text:
The Good Dinosaur

Film score by Mychael Danna and Jeff Danna

Released
November 20, 2015

Recorded
2015

Studio
Eastwood Scoring Stage, Warner Bros., Los Angeles

Genre
Soundtrack, film score

Length
63:22

Label
Walt Disney

Producer
Chris Montan

Pixar film soundtrack chronology

Inside Out: Original Soundtrack
(2015)Inside Out: Original Soundtrack2015
The Good Dinosaur
(2015)
Finding Dory
(2016)Finding Dory2016

Mychael Danna chronology

The Captive
(2014) The Captive2014
The Good Dinosaur
(2015) The Good Dinosaur2015
Storks
(2016) Storks2016

Jeff Danna chronology

Montage of Heck
(2015) Montage of Heck2015
The Good Dinosaur
(2015) The Good Dinosaur2015
Storks
(2016) Storks2016

text: The film's score was composed by Mychael Danna and his brother, Jeff Danna.[3] It marks the first Pixar film to be scored by two composers.[3] Danna was approached by Sohn and Ream due to his score for Life of Pi, which won an Academy Award.[3] Having a lot of work, he invited his brother Jeff Danna as a co-writer.[3] Walt Disney Records released the soundtrack on November 20, 2015.[40]

text: All music composed by Mychael Danna and Jeff Danna.

text:
No.
Title
Length

1.
"Homestead"
2:11

2.
"Hello Arlo"
2:49

3.
"Chores"
0:55

4.
"Make Your Mark"
2:07

5.
"Fireflies"
2:16

6.
"Critter Problem"
1:04

7.
"You're Me and More"
3:05

8.
"Family Struggle"
1:23

9.
"Swept Away"
1:33

10.
"Mountain Top"
0:51

11.
"Lost in the World"
3:35

12.
"Offerings"
1:32

13.
"Unexpected Friend"
2:56

14.
"Pet Collector"
2:24

15.
"Swimming Lessons"
2:29

16.
"Orphans"
4:39

17.
"The Storm"
1:17

18.
"I'm Never Getting Home"
0:44

19.
"Storm Chasers"
1:22

20.
"Bloodhound"
1:37

21.
"Fight Them Rustlers"
1:46

22.
"Run With the Herd"
3:51

23.
"Returned Call"
1:25

24.
"Sky Sharks"
1:46

25.
"Arlo's Vision"
1:35

26.
"Rescue"
2:31

27.
"Over the Falls"
2:41

28.
"Goodbye Spot"
4:12

29.
"Homecoming"
1:24

30.
"Arlo Makes His Mark"
1:22

Total length:
63:22

text: Video game[edit]

text: A figure of Spot and Power Discs of Arlo, Ramsey, Nash, and Butch have been released for Disney Infinity 3.0.[41]

text: Release[edit]

text: The Good Dinosaur was theatrically released on November 25, 2015.[42] Before the rescheduling from 2014 to 2015, a Monsters University short film titled Party Central was set to accompany the film, but was instead shown with the theatrical release of Disney's Muppets Most Wanted.[43] In April 2015, it was announced that a new Pixar short, Sanjay's Super Team, directed by Sanjay Patel, would be shown in front of The Good Dinosaur instead.[44]

text: The film's teaser trailer was released on June 2, 2015,[45][46] and the first official trailer was released on July 21, 2015.[47] The film received an exclusive run at The Grand Rex in Paris a week before its U.S. and European premiere.[48]

text: Home media[edit]

text: The Good Dinosaur was released by Walt Disney Studios Home Entertainment on Blu-ray (2D and 3D), DVD, and digital download in the United States on February 23, 2016. Blu-ray bonus features include Sanjay's Super Team, audio commentary, behind-the-scenes featurettes, deleted scenes, and the "Hide and Seek" short promotional clip.[49][50]

text: Reception[edit]

text: Box office[edit]

text: The Good Dinosaur grossed $123.1 million in the United States and Canada and $209.1 million in other countries, for a worldwide total of $332.2 million[7] against a production budget of $175–200 million,[5][6] making it Pixar's lowest-grossing film. Adding in the marketing budget, the film had a cost of $350 million,[51] which it was not able to recoup.[52][53][54][55] However, the film proved to be much more financially successful in terms of home video sales.[56][57]

text: In North America, The Good Dinosaur opened on Wednesday, November 25, 2015 alongside Creed and Victor Frankenstein, as well as the wide releases of Brooklyn, Spotlight, and Trumbo. The film was projected to gross $60–65 million from 3,749 theaters in its first five days, including $45 million in its opening weekend.[58] It made $1.3 million from its Tuesday night previews and $9.8 million on its first day. The film ended up grossing $55.6 million in its first five days, including $39.2 million in its opening weekend, finishing second at the box office behind the second weekend of Mockingjay – Part 2, the final installment in the Hunger Games film series.[59]

text: Outside North America, it earned $28.7 million from 39 markets, including 8 significant markets. Unlike the USA, where it had the benefit of the Thanksgiving weekend, it did not have the same benefit internationally. Rather, its release date was designed to set it up to run through the December holiday.[60] The top openings were in the United Kingdom ($4.3 million), Mexico ($3.6 million), France ($3.2 million), Argentina ($2.2 million), and Russia ($2.1 million). Of those, it opened at No. 1 in Mexico, Argentina and Russia.[61] In terms of total earnings, its leading markets are the United Kingdom ($10.1 million), Mexico ($8.7 million), and France ($8.1 million).[62]

text: Critical response[edit]

text: The review aggregator website Rotten Tomatoes gives the film a rating of 76%, based on 198 reviews, with a rating average of 6.5/10. The site's critical consensus reads, "The Good Dinosaur delivers thrillingly beautiful animation in service of a worthy story that, even if it doesn't quite live up to the lofty standards set by Pixar, still adds up to charming, family-friendly entertainment."[63] On Metacritic, the film has a score of 66 out of 100, based on 37 critics, indicating "generally favorable reviews."[64] Audiences polled by CinemaScore gave the film an average grade of "A" on an A+ to F scale.[59]

text: Kenneth Turan of the Los Angeles Times described it as "antic and unexpected as well as homiletic, rife with subversive elements, wacky critters and some of the most beautiful landscapes ever seen in a computer animated film."[65] Manohla Dargis of The New York Times felt the film "has a few things on its mind, but its tone is overwhelmingly playful, not hectoring."[66] Joe Morgenstern of The Wall Street Journal opined that "As Pixar productions go, this one isn't a groundbreaker, but it's heartfelt and endearing, as well as visually splendiferous, and kids will love it for sure."[67] "Clever and cloying by turns, it's a movie that always seems to be trying to evolve beyond its conventional trappings, and not succeeding as often as Pixar devotees have come to expect," wrote Justin Chang of Variety.[68] Many of the reviewers praised the sophistication of its nearly photorealistic backgrounds.[66][69][70][71]

text: Christopher Orr of The Atlantic felt it to be the studio's first film explicitly targeted towards children, though it is "by no means a bad movie […] It's a simple story, well-told."[69] Mark Feeney, writing for The Boston Globe, felt similarly, deeming it a "very middling movie […] The Good Dinosaur generally features a sort of sentimentality and emotional reductiveness that make it seem meant for small children as no previous Pixar movie has."[70] The Washington Post's Stephanie Merry dubbed it "a nice, conventional story, but it’s not Pixar-level imaginative."[71] Richard Roeper, for the Chicago Sun-Times, felt it "one strange, aggressively gross and dark adventure […] Inconsistent and weird, The Good Dinosaur is second-level Pixar all the way."[72] Michael Rechtshaffen of The Hollywood Reporter derided its "disappointingly derivative" screenplay.[73]

text: Accolades[edit]

text:
List of awards and nominations

Award
Date of ceremony
Category
Recipient
Outcome

American Cinema Editors Awards
January 29, 2016
Best Edited Animated Feature Film
Stephen Schaffer
Nominated

Annie Awards[74]
February 6, 2016
Best Animated Feature
The Good Dinosaur

Outstanding Achievement in Animated Effects in an Animated Production
John Reisch and Stephen Marshall
Won

Outstanding Achievement in Character Animation in a Feature Production
Mark C. Harris
Nominated

K.C. Roeyer

Outstanding Achievement in Character Design in an Animated Feature Production
Matt Nolte

Outstanding Achievement in Music in an Animated Feature Production
Mychael Danna and Jeff Danna

Outstanding Achievement in Production Design in an Animated Feature Production
Harley Jessup, Sharon Calahan, Bryn Imagire, Noah Klocek and Huy Nguyen

Outstanding Achievement in Storyboarding in an Animated Feature Production
Bill Presing

Rosana Sullivan

J.P. Vine, Tony Rosenast and Enrico Casarosa

Austin Film Critics Association Awards[75]
December 29, 2015
Best Animated Film
The Good Dinosaur

Black Reel Awards
February 18, 2016
Best Voice Performance
Jeffrey Wright (Poppa Henry)

British Academy Children's Awards[76][77]
November 20, 2016
Feature Film
Peter Sohn and Meg LeFauve

BAFTA Kids' Vote
The Good Dinosaur

Chicago Film Critics Association Awards
December 16, 2015
Best Animated Film

Critics' Choice Awards
January 17, 2016
Best Animated Feature

Florida Film Critics Circle Awards
December 23, 2015
Best Animated Film

Golden Globe Awards[78]
January 10, 2016
Best Animated Feature Film

Golden Reel Awards[79]
February 27, 2016
Best Sound Editing: Sound Effects, Foley, Dialogue & ADR in an Animation Feature Film
Shannon Mills

Houston Film Critics Society Awards
January 9, 2016
Best Animated Feature Film
The Good Dinosaur

Online Film Critics Society Awards[80]
December 14, 2015
Best Animated Film

Producers Guild of America Awards
January 23, 2016
Outstanding Producer of Animated Theatrical Motion Pictures
Denise Ream

San Diego Film Critics Society Awards[81][82]
December 14, 2015
Best Animated Film
The Good Dinosaur

Satellite Awards[83]
February 21, 2016
Best Motion Picture, Animated or Mixed Media

Saturn Awards[84]
June 22, 2016
Best Animated Film

St. Louis Gateway Film Critics Association Awards[85]
December 20, 2015
Best Animated Film

Village Voice Film Poll[86]
January 7, 2016
Best Animated Film
10th Place

Visual Effects Society Awards
February 2, 2016
Outstanding Visual Effects in an Animated Feature
Sanjay Bakshi, Denise Ream, Michael Venturini, and Jon Reisch
Won

Outstanding Animated Performance in an Animated Feature
Ana Gabriela Lacaze, Jacob Brooks, Lou Hamou-Lhadj, and Mark C. Harris (Spot)
Nominated

Outstanding Created Environment in an Animated Feature
David Munier, Matthew Webb, Matt Kuruc, and Tom Miller (The Farm)
Won

Outstanding Effects Simulations in an Animated Feature
Stephen Marshall, Magnus Wrenninge, Michael Hall, and Hemagiri Arumugam


document_title: Cell nucleus

question_text: which types of cells have genetic material that is contained in the nucleus

text: Cell nucleus

text: History

text: The nucleus was the first organelle to be discovered. What is most likely the oldest preserved drawing dates back to the early microscopist Antonie van Leeuwenhoek (1632–1723). He observed a "lumen", the nucleus, in the red blood cells of salmon.[2] Unlike mammalian red blood cells, those of other vertebrates still contain nuclei.

text: The nucleus was also described by Franz Bauer in 1804[3] and in more detail in 1831 by Scottish botanist Robert Brown in a talk at the Linnean Society of London. Brown was studying orchids under microscope when he observed an opaque area, which he called the "areola" or "nucleus", in the cells of the flower's outer layer.[4]

text: He did not suggest a potential function. In 1838, Matthias Schleiden proposed that the nucleus plays a role in generating cells, thus he introduced the name "cytoblast" (cell builder). He believed that he had observed new cells assembling around "cytoblasts". Franz Meyen was a strong opponent of this view, having already described cells multiplying by division and believing that many cells would have no nuclei. The idea that cells can be generated de novo, by the "cytoblast" or otherwise, contradicted work by Robert Remak (1852) and Rudolf Virchow (1855) who decisively propagated the new paradigm that cells are generated solely by cells ("Omnis cellula e cellula"). The function of the nucleus remained unclear.[5]

text: Between 1877 and 1878, Oscar Hertwig published several studies on the fertilization of sea urchin eggs, showing that the nucleus of the sperm enters the oocyte and fuses with its nucleus. This was the first time it was suggested that an individual develops from a (single) nucleated cell. This was in contradiction to Ernst Haeckel's theory that the complete phylogeny of a species would be repeated during embryonic development, including generation of the first nucleated cell from a "monerula", a structureless mass of primordial mucus ("Urschleim"). Therefore, the necessity of the sperm nucleus for fertilization was discussed for quite some time. However, Hertwig confirmed his observation in other animal groups, including amphibians and molluscs. Eduard Strasburger produced the same results for plants in 1884. This paved the way to assign the nucleus an important role in heredity. In 1873, August Weismann postulated the equivalence of the maternal and paternal germ cells for heredity. The function of the nucleus as carrier of genetic information became clear only later, after mitosis was discovered and the Mendelian rules were rediscovered at the beginning of the 20th century; the chromosome theory of heredity was therefore developed.[5]

text: Structures

text: The nucleus is the largest cellular organelle in animal cells.[6] In mammalian cells, the average diameter of the nucleus is approximately 6 micrometres (µm), which occupies about 10% of the total cell volume.[7] The viscous liquid within it is called nucleoplasm (or karyolymph), and is similar in composition to the cytosol found outside the nucleus.[8] It appears as a dense, roughly spherical or irregular organelle. The composition by dry weight of the nucleus is approximately: DNA 9%, RNA 1%, Histone Protein 11%, Residual Protein 14%, Acidic Proteins 65%.

text: Nuclear envelope and pores

text:
The eukaryotic cell nucleus. Visible in this diagram are the ribosome-studded double membranes of the nuclear envelope, the DNA (complexed as chromatin), and the nucleolus. Within the cell nucleus is a viscous liquid called nucleoplasm, similar to the cytoplasm found outside the nucleus.

A cross section of a nuclear pore on the surface of the nuclear envelope (1). Other diagram labels show (2) the outer ring, (3) spokes, (4) basket, and (5) filaments.

text: The nuclear envelope, otherwise known as nuclear membrane, consists of two cellular membranes, an inner and an outer membrane, arranged parallel to one another and separated by 10 to 50 nanometres (nm). The nuclear envelope completely encloses the nucleus and separates the cell's genetic material from the surrounding cytoplasm, serving as a barrier to prevent macromolecules from diffusing freely between the nucleoplasm and the cytoplasm.[9] The outer nuclear membrane is continuous with the membrane of the rough endoplasmic reticulum (RER), and is similarly studded with ribosomes.[9] The space between the membranes is called the perinuclear space and is continuous with the RER lumen.

text: Nuclear pores, which provide aqueous channels through the envelope, are composed of multiple proteins, collectively referred to as nucleoporins. The pores are about 125 million daltons in molecular weight and consist of around 50 (in yeast) to several hundred proteins (in vertebrates).[6] The pores are 100 nm in total diameter; however, the gap through which molecules freely diffuse is only about 9 nm wide, due to the presence of regulatory systems within the center of the pore. This size selectively allows the passage of small water-soluble molecules while preventing larger molecules, such as nucleic acids and larger proteins, from inappropriately entering or exiting the nucleus. These large molecules must be actively transported into the nucleus instead. The nucleus of a typical mammalian cell will have about 3000 to 4000 pores throughout its envelope,[10] each of which contains an eightfold-symmetric ring-shaped structure at a position where the inner and outer membranes fuse.[11] Attached to the ring is a structure called the nuclear basket that extends into the nucleoplasm, and a series of filamentous extensions that reach into the cytoplasm. Both structures serve to mediate binding to nuclear transport proteins.[6]

text: Most proteins, ribosomal subunits, and some DNAs are transported through the pore complexes in a process mediated by a family of transport factors known as karyopherins. Those karyopherins that mediate movement into the nucleus are also called importins, whereas those that mediate movement out of the nucleus are called exportins. Most karyopherins interact directly with their cargo, although some use adaptor proteins.[12] Steroid hormones such as cortisol and aldosterone, as well as other small lipid-soluble molecules involved in intercellular signaling, can diffuse through the cell membrane and into the cytoplasm, where they bind nuclear receptor proteins that are trafficked into the nucleus. There they serve as transcription factors when bound to their ligand; in the absence of a ligand, many such receptors function as histone deacetylases that repress gene expression.[6]

text: Nuclear lamina

text: In animal cells, two networks of intermediate filaments provide the nucleus with mechanical support: The nuclear lamina forms an organized meshwork on the internal face of the envelope, while less organized support is provided on the cytosolic face of the envelope. Both systems provide structural support for the nuclear envelope and anchoring sites for chromosomes and nuclear pores.[7]

text: The nuclear lamina is composed mostly of lamin proteins. Like all proteins, lamins are synthesized in the cytoplasm and later transported to the nucleus interior, where they are assembled before being incorporated into the existing network of nuclear lamina.[13][14] Lamins found on the cytosolic face of the membrane, such as emerin and nesprin, bind to the cytoskeleton to provide structural support. Lamins are also found inside the nucleoplasm where they form another regular structure, known as the nucleoplasmic veil,[15] that is visible using fluorescence microscopy. The actual function of the veil is not clear, although it is excluded from the nucleolus and is present during interphase.[16] Lamin structures that make up the veil, such as LEM3, bind chromatin and disrupting their structure inhibits transcription of protein-coding genes.[17]

text: Like the components of other intermediate filaments, the lamin monomer contains an alpha-helical domain used by two monomers to coil around each other, forming a dimer structure called a coiled coil. Two of these dimer structures then join side by side, in an antiparallel arrangement, to form a tetramer called a protofilament. Eight of these protofilaments form a lateral arrangement that is twisted to form a ropelike filament. These filaments can be assembled or disassembled in a dynamic manner, meaning that changes in the length of the filament depend on the competing rates of filament addition and removal.[7]

text: Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as laminopathies. The most notable laminopathy is the family of diseases known as progeria, which causes the appearance of premature aging in its sufferers. The exact mechanism by which the associated biochemical changes give rise to the aged phenotype is not well understood.[18]

text: Chromosomes

text: The cell nucleus contains the majority of the cell's genetic material in the form of multiple linear DNA molecules organized into structures called chromosomes. Each human cell contains roughly two meters of DNA. During most of the cell cycle these are organized in a DNA-protein complex known as chromatin, and during cell division the chromatin can be seen to form the well-defined chromosomes familiar from a karyotype. A small fraction of the cell's genes are located instead in the mitochondria.

text: There are two types of chromatin. Euchromatin is the less compact DNA form, and contains genes that are frequently expressed by the cell.[19] The other type, heterochromatin, is the more compact form, and contains DNA that is infrequently transcribed. This structure is further categorized into facultative heterochromatin, consisting of genes that are organized as heterochromatin only in certain cell types or at certain stages of development, and constitutive heterochromatin that consists of chromosome structural components such as telomeres and centromeres.[20] During interphase the chromatin organizes itself into discrete individual patches,[21] called chromosome territories.[22] Active genes, which are generally found in the euchromatic region of the chromosome, tend to be located towards the chromosome's territory boundary.[23]

text: Antibodies to certain types of chromatin organization, in particular, nucleosomes, have been associated with a number of autoimmune diseases, such as systemic lupus erythematosus.[24] These are known as anti-nuclear antibodies (ANA) and have also been observed in concert with multiple sclerosis as part of general immune system dysfunction.[25] As in the case of progeria, the role played by the antibodies in inducing the symptoms of autoimmune diseases is not obvious.

text: Nucleolus

text: The nucleolus is a discrete densely stained structure found in the nucleus. It is not surrounded by a membrane, and is sometimes called a suborganelle. It forms around tandem repeats of rDNA, DNA coding for ribosomal RNA (rRNA). These regions are called nucleolar organizer regions (NOR). The main roles of the nucleolus are to synthesize rRNA and assemble ribosomes. The structural cohesion of the nucleolus depends on its activity, as ribosomal assembly in the nucleolus results in the transient association of nucleolar components, facilitating further ribosomal assembly, and hence further association. This model is supported by observations that inactivation of rDNA results in intermingling of nucleolar structures.[26]

text: In the first step of ribosome assembly, a protein called RNA polymerase I transcribes rDNA, which forms a large pre-rRNA precursor. This is cleaved into the subunits 5.8S, 18S, and 28S rRNA.[27] The transcription, post-transcriptional processing, and assembly of rRNA occurs in the nucleolus, aided by small nucleolar RNA (snoRNA) molecules, some of which are derived from spliced introns from messenger RNAs encoding genes related to ribosomal function. The assembled ribosomal subunits are the largest structures passed through the nuclear pores.[6]

text: When observed under the electron microscope, the nucleolus can be seen to consist of three distinguishable regions: the innermost fibrillar centers (FCs), surrounded by the dense fibrillar component (DFC), which in turn is bordered by the granular component (GC). Transcription of the rDNA occurs either in the FC or at the FC-DFC boundary, and, therefore, when rDNA transcription in the cell is increased, more FCs are detected. Most of the cleavage and modification of rRNAs occurs in the DFC, while the latter steps involving protein assembly onto the ribosomal subunits occur in the GC.[27]

text: Other subnuclear bodies

text:
Subnuclear structure sizes

Structure name
Structure diameter

Cajal bodies
0.2–2.0 µm
[28]

Clastosomes
0.2-0.5 µm
[29]

PIKA
5 µm
[30]

PML bodies
0.2–1.0 µm
[31]

Paraspeckles
0.2–1.0 µm
[32]

Speckles
20–25 nm
[30]

text: Besides the nucleolus, the nucleus contains a number of other non-membrane-delineated bodies. These include Cajal bodies, Gemini of coiled bodies, polymorphic interphase karyosomal association (PIKA), promyelocytic leukaemia (PML) bodies, paraspeckles, and splicing speckles. Although little is known about a number of these domains, they are significant in that they show that the nucleoplasm is not a uniform mixture, but rather contains organized functional subdomains.[31]

text: Other subnuclear structures appear as part of abnormal disease processes. For example, the presence of small intranuclear rods has been reported in some cases of nemaline myopathy. This condition typically results from mutations in actin, and the rods themselves consist of mutant actin as well as other cytoskeletal proteins.[33]

text: Cajal bodies and gems

text: A nucleus typically contains between 1 and 10 compact structures called Cajal bodies or coiled bodies (CB), whose diameter measures between 0.2 µm and 2.0 µm depending on the cell type and species.[28] When seen under an electron microscope, they resemble balls of tangled thread[30] and are dense foci of distribution for the protein coilin.[34] CBs are involved in a number of different roles relating to RNA processing, specifically small nucleolar RNA (snoRNA) and small nuclear RNA (snRNA) maturation, and histone mRNA modification.[28]

text: Similar to Cajal bodies are Gemini of Cajal bodies, or gems, whose name is derived from the Gemini constellation in reference to their close "twin" relationship with CBs. Gems are similar in size and shape to CBs, and in fact are virtually indistinguishable under the microscope.[34] Unlike CBs, gems do not contain small nuclear ribonucleoproteins (snRNPs), but do contain a protein called survival of motor neuron (SMN) whose function relates to snRNP biogenesis. Gems are believed to assist CBs in snRNP biogenesis,[35] though it has also been suggested from microscopy evidence that CBs and gems are different manifestations of the same structure.[34] Later ultrastructural studies have shown gems to be twins of Cajal bodies with the difference being in the coilin component; Cajal bodies are SMN positive and coilin positive, and gems are SMN positive and coilin negative.[36]

text: RAFA and PTF domains

text: RAFA domains, or polymorphic interphase karyosomal associations, were first described in microscopy studies in 1991. Their function remains unclear, though they were not thought to be associated with active DNA replication, transcription, or RNA processing.[37] They have been found to often associate with discrete domains defined by dense localization of the transcription factor PTF, which promotes transcription of small nuclear RNA (snRNA).[38]

text: PML bodies

text: Promyelocytic leukaemia bodies (PML bodies) are spherical bodies found scattered throughout the nucleoplasm, measuring around 0.1–1.0 µm. They are known by a number of other names, including nuclear domain 10 (ND10), Kremer bodies, and PML oncogenic domains. PML bodies are named after one of their major components, the promyelocytic leukemia protein (PML). They are often seen in the nucleus in association with Cajal bodies and cleavage bodies.[31] PML bodies belong to the nuclear matrix, an ill-defined super-structure of the nucleus proposed to anchor and regulate many nuclear functions, including DNA replication, transcription, or epigenetic silencing.[39] The PML protein is the key organizer of these domains that recruits an ever-growing number of proteins, whose only common known feature to date is their ability to be SUMOylated. Yet, pml-/- mice (which have their PML gene deleted) cannot assemble nuclear bodies, develop normally and live well, demonstrating that PML bodies are dispensable for most basic biological functions.[39]

text: Splicing speckles

text: Speckles are subnuclear structures that are enriched in pre-messenger RNA splicing factors and are located in the interchromatin regions of the nucleoplasm of mammalian cells. At the fluorescence-microscope level they appear as irregular, punctate structures, which vary in size and shape, and when examined by electron microscopy they are seen as clusters of interchromatin granules. Speckles are dynamic structures, and both their protein and RNA-protein components can cycle continuously between speckles and other nuclear locations, including active transcription sites. Studies on the composition, structure and behaviour of speckles have provided a model for understanding the functional compartmentalization of the nucleus and the organization of the gene-expression machinery[40] splicing snRNPs[41][42] and other splicing proteins necessary for pre-mRNA processing.[43] Because of a cell's changing requirements, the composition and location of these bodies changes according to mRNA transcription and regulation via phosphorylation of specific proteins.[44] The splicing speckles are also known as nuclear speckles (nuclear specks), splicing factor compartments (SF compartments), interchromatin granule clusters (IGCs), B snurposomes.[45] B snurposomes are found in the amphibian oocyte nuclei and in Drosophila melanogaster embryos. B snurposomes appear alone or attached to the Cajal bodies in the electron micrographs of the amphibian nuclei.[46] IGCs function as storage sites for the splicing factors.[47]

text: Paraspeckles

text: Discovered by Fox et al. in 2002, paraspeckles are irregularly shaped compartments in the nucleus' interchromatin space.[48] First documented in HeLa cells, where there are generally 10–30 per nucleus,[49] paraspeckles are now known to also exist in all human primary cells, transformed cell lines, and tissue sections.[50] Their name is derived from their distribution in the nucleus; the "para" is short for parallel and the "speckles" refers to the splicing speckles to which they are always in close proximity.[49]

text: Paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity. They are transcription dependent[48] and in the absence of RNA Pol II transcription, the paraspeckle disappears and all of its associated protein components (PSP1, p54nrb, PSP2, CFI(m)68, and PSF) form a crescent shaped perinucleolar cap in the nucleolus. This phenomenon is demonstrated during the cell cycle. In the cell cycle, paraspeckles are present during interphase and during all of mitosis except for telophase. During telophase, when the two daughter nuclei are formed, there is no RNA Pol II transcription so the protein components instead form a perinucleolar cap.[50]

text: Perichromatin fibrils

text: Perichromatin fibrils are visible only under electron microscope. They are located next to the transcriptionally active chromatin and are hypothesized to be the sites of active pre-mRNA processing.[47]

text: Clastosome

text: Clastosomes are small nuclear bodies (0.2-0.5 µm) described as donut-shaped due to the peripheral capsule around these bodies.[29] This name is derived from the Greek klastos, broken and soma, body.[29] Clastosomes are not typically present in normal cells, making them hard to detect. They form under high proteolysis conditions within the nucleus and degrade once there is a decrease in activity or if cells are treated with proteasome inhibitors.[29][51] The scarcity of clastosomes in cells indicates that they are not required for proteasome function.[52] Osmotic stress has also been shown to cause the formation of clastosomes. [53] These nuclear bodies contain catalytic and regulatory sub-units of the proteasome and its substrates, indicating that clastosomes are sites for degrading proteins.[52]

text: The Clastosome would be an important addition to the ‘Cell Nucleus’ Wiki page because it plays a role in cell function. With further research, we may discover that this nuclear body has a more influential job in cell function than what is already known. [29]

text: Function

text: The nucleus provides a site for genetic transcription that is segregated from the location of translation in the cytoplasm, allowing levels of gene regulation that are not available to prokaryotes. The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell cycle.

text: The nucleus is an organelle found in eukaryotic cells. Inside its fully enclosed nuclear membrane, it contains the majority of the cell's genetic material. This material is organized as DNA molecules, along with a variety of proteins, to form chromosomes.

text: Cell compartmentalization

text: The nuclear envelope allows the nucleus to control its contents, and separate them from the rest of the cytoplasm where necessary. This is important for controlling processes on either side of the nuclear membrane. In most cases where a cytoplasmic process needs to be restricted, a key participant is removed to the nucleus, where it interacts with transcription factors to downregulate the production of certain enzymes in the pathway. This regulatory mechanism occurs in the case of glycolysis, a cellular pathway for breaking down glucose to produce energy. Hexokinase is an enzyme responsible for the first the step of glycolysis, forming glucose-6-phosphate from glucose. At high concentrations of fructose-6-phosphate, a molecule made later from glucose-6-phosphate, a regulator protein removes hexokinase to the nucleus,[54] where it forms a transcriptional repressor complex with nuclear proteins to reduce the expression of genes involved in glycolysis.[55]

text: In order to control which genes are being transcribed, the cell separates some transcription factor proteins responsible for regulating gene expression from physical access to the DNA until they are activated by other signaling pathways. This prevents even low levels of inappropriate gene expression. For example, in the case of NF-κB-controlled genes, which are involved in most inflammatory responses, transcription is induced in response to a signal pathway such as that initiated by the signaling molecule TNF-α, binds to a cell membrane receptor, resulting in the recruitment of signalling proteins, and eventually activating the transcription factor NF-κB. A nuclear localisation signal on the NF-κB protein allows it to be transported through the nuclear pore and into the nucleus, where it stimulates the transcription of the target genes.[7]

text: The compartmentalization allows the cell to prevent translation of unspliced mRNA.[56] Eukaryotic mRNA contains introns that must be removed before being translated to produce functional proteins. The splicing is done inside the nucleus before the mRNA can be accessed by ribosomes for translation. Without the nucleus, ribosomes would translate newly transcribed (unprocessed) mRNA, resulting in malformed and nonfunctional proteins.

text: Gene expression

text: Gene expression first involves transcription, in which DNA is used as a template to produce RNA. In the case of genes encoding proteins, that RNA produced from this process is messenger RNA (mRNA), which then needs to be translated by ribosomes to form a protein. As ribosomes are located outside the nucleus, mRNA produced needs to be exported.[57]

text: Since the nucleus is the site of transcription, it also contains a variety of proteins that either directly mediate transcription or are involved in regulating the process. These proteins include helicases, which unwind the double-stranded DNA molecule to facilitate access to it, RNA polymerases, which bind to the DNA promoter to synthesize the growing RNA molecule, topoisomerases, which change the amount of supercoiling in DNA, helping it wind and unwind, as well as a large variety of transcription factors that regulate expression.[58]

text: Processing of pre-mRNA

text: Newly synthesized mRNA molecules are known as primary transcripts or pre-mRNA. They must undergo post-transcriptional modification in the nucleus before being exported to the cytoplasm; mRNA that appears in the cytoplasm without these modifications is degraded rather than used for protein translation. The three main modifications are 5' capping, 3' polyadenylation, and RNA splicing. While in the nucleus, pre-mRNA is associated with a variety of proteins in complexes known as heterogeneous ribonucleoprotein particles (hnRNPs). Addition of the 5' cap occurs co-transcriptionally and is the first step in post-transcriptional modification. The 3' poly-adenine tail is only added after transcription is complete.

text: RNA splicing, carried out by a complex called the spliceosome, is the process by which introns, or regions of DNA that do not code for protein, are removed from the pre-mRNA and the remaining exons connected to re-form a single continuous molecule. This process normally occurs after 5' capping and 3' polyadenylation but can begin before synthesis is complete in transcripts with many exons.[6] Many pre-mRNAs, including those encoding antibodies, can be spliced in multiple ways to produce different mature mRNAs that encode different protein sequences. This process is known as alternative splicing, and allows production of a large variety of proteins from a limited amount of DNA.

text: Dynamics and regulation

text: Nuclear transport

text: The entry and exit of large molecules from the nucleus is tightly controlled by the nuclear pore complexes. Although small molecules can enter the nucleus without regulation,[59] macromolecules such as RNA and proteins require association karyopherins called importins to enter the nucleus and exportins to exit. "Cargo" proteins that must be translocated from the cytoplasm to the nucleus contain short amino acid sequences known as nuclear localization signals, which are bound by importins, while those transported from the nucleus to the cytoplasm carry nuclear export signals bound by exportins. The ability of importins and exportins to transport their cargo is regulated by GTPases, enzymes that hydrolyze the molecule guanosine triphosphate to release energy. The key GTPase in nuclear transport is Ran, which can bind either GTP or GDP (guanosine diphosphate), depending on whether it is located in the nucleus or the cytoplasm. Whereas importins depend on RanGTP to dissociate from their cargo, exportins require RanGTP in order to bind to their cargo.[12]

text: Nuclear import depends on the importin binding its cargo in the cytoplasm and carrying it through the nuclear pore into the nucleus. Inside the nucleus, RanGTP acts to separate the cargo from the importin, allowing the importin to exit the nucleus and be reused. Nuclear export is similar, as the exportin binds the cargo inside the nucleus in a process facilitated by RanGTP, exits through the nuclear pore, and separates from its cargo in the cytoplasm.

text: Specialized export proteins exist for translocation of mature mRNA and tRNA to the cytoplasm after post-transcriptional modification is complete. This quality-control mechanism is important due to these molecules' central role in protein translation. Mis-expression of a protein due to incomplete excision of exons or mis-incorporation of amino acids could have negative consequences for the cell; thus, incompletely modified RNA that reaches the cytoplasm is degraded rather than used in translation.[6]

text: Assembly and disassembly

text: During its lifetime, a nucleus may be broken down or destroyed, either in the process of cell division or as a consequence of apoptosis (the process of programmed cell death). During these events, the structural components of the nucleus — the envelope and lamina — can be systematically degraded. In most cells, the disassembly of the nuclear envelope marks the end of the prophase of mitosis. However, this disassembly of the nucleus is not a universal feature of mitosis and does not occur in all cells. Some unicellular eukaryotes (e.g., yeasts) undergo so-called closed mitosis, in which the nuclear envelope remains intact. In closed mitosis, the daughter chromosomes migrate to opposite poles of the nucleus, which then divides in two. The cells of higher eukaryotes, however, usually undergo open mitosis, which is characterized by breakdown of the nuclear envelope. The daughter chromosomes then migrate to opposite poles of the mitotic spindle, and new nuclei reassemble around them.

text: At a certain point during the cell cycle in open mitosis, the cell divides to form two cells. In order for this process to be possible, each of the new daughter cells must have a full set of genes, a process requiring replication of the chromosomes as well as segregation of the separate sets. This occurs by the replicated chromosomes, the sister chromatids, attaching to microtubules, which in turn are attached to different centrosomes. The sister chromatids can then be pulled to separate locations in the cell. In many cells, the centrosome is located in the cytoplasm, outside the nucleus; the microtubules would be unable to attach to the chromatids in the presence of the nuclear envelope.[60] Therefore, the early stages in the cell cycle, beginning in prophase and until around prometaphase, the nuclear membrane is dismantled.[15] Likewise, during the same period, the nuclear lamina is also disassembled, a process regulated by phosphorylation of the lamins by protein kinases such as the CDC2 protein kinase.[61] Towards the end of the cell cycle, the nuclear membrane is reformed, and around the same time, the nuclear lamina are reassembled by dephosphorylating the lamins.[61]

text: However, in dinoflagellates, the nuclear envelope remains intact, the centrosomes are located in the cytoplasm, and the microtubules come in contact with chromosomes, whose centromeric regions are incorporated into the nuclear envelope (the so-called closed mitosis with extranuclear spindle). In many other protists (e.g., ciliates, sporozoans) and fungi, the centrosomes are intranuclear, and their nuclear envelope also does not disassemble during cell division.

text: Apoptosis is a controlled process in which the cell's structural components are destroyed, resulting in death of the cell. Changes associated with apoptosis directly affect the nucleus and its contents, for example, in the condensation of chromatin and the disintegration of the nuclear envelope and lamina. The destruction of the lamin networks is controlled by specialized apoptotic proteases called caspases, which cleave the lamin proteins and, thus, degrade the nucleus' structural integrity. Lamin cleavage is sometimes used as a laboratory indicator of caspase activity in assays for early apoptotic activity.[15] Cells that express mutant caspase-resistant lamins are deficient in nuclear changes related to apoptosis, suggesting that lamins play a role in initiating the events that lead to apoptotic degradation of the nucleus.[15] Inhibition of lamin assembly itself is an inducer of apoptosis.[62]

text: The nuclear envelope acts as a barrier that prevents both DNA and RNA viruses from entering the nucleus. Some viruses require access to proteins inside the nucleus in order to replicate and/or assemble. DNA viruses, such as herpesvirus replicate and assemble in the cell nucleus, and exit by budding through the inner nuclear membrane. This process is accompanied by disassembly of the lamina on the nuclear face of the inner membrane.[15]

text: Disease-related dynamics

text: Initially, it has been suspected that immunoglobulins in general and autoantibodies in particular do not enter the nucleus. Now there is a body of evidence that under pathological conditions (e.g. lupus erythematosus) IgG can enter the nucleus.[63]

text: Nuclei per cell

text: Most eukaryotic cell types usually have a single nucleus, but some have no nuclei, while others have several. This can result from normal development, as in the maturation of mammalian red blood cells, or from faulty cell division.

text: Anucleated cells

text: An anucleated cell contains no nucleus and is, therefore, incapable of dividing to produce daughter cells. The best-known anucleated cell is the mammalian red blood cell, or erythrocyte, which also lacks other organelles such as mitochondria, and serves primarily as a transport vessel to ferry oxygen from the lungs to the body's tissues. Erythrocytes mature through erythropoiesis in the bone marrow, where they lose their nuclei, organelles, and ribosomes. The nucleus is expelled during the process of differentiation from an erythroblast to a reticulocyte, which is the immediate precursor of the mature erythrocyte.[64] The presence of mutagens may induce the release of some immature "micronucleated" erythrocytes into the bloodstream.[65][66] Anucleated cells can also arise from flawed cell division in which one daughter lacks a nucleus and the other has two nuclei.

text: In flowering plants, this condition occurs in sieve tube elements.

text: Multinucleated cells

text: Multinucleated cells contain multiple nuclei. Most acantharean species of protozoa[67] and some fungi in mycorrhizae[68] have naturally multinucleated cells. Other examples include the intestinal parasites in the genus Giardia, which have two nuclei per cell.[69] In humans, skeletal muscle cells, called myocytes and syncytium, become multinucleated during development; the resulting arrangement of nuclei near the periphery of the cells allows maximal intracellular space for myofibrils.[6] Multinucleated and binucleated cells can also be abnormal in humans; for example, cells arising from the fusion of monocytes and macrophages, known as giant multinucleated cells, sometimes accompany inflammation[70] and are also implicated in tumor formation.[71]

text: A number of dinoflagellates are known to have two nuclei.[72] Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic diatom. The mitochondria and the plastids of the diatom somehow remain functional.

text: Evolution

text: As the major defining characteristic of the eukaryotic cell, the nucleus' evolutionary origin has been the subject of much speculation. Four major hypotheses have been proposed to explain the existence of the nucleus, although none have yet earned widespread support.[73]

text: The first model known as the "syntrophic model" proposes that a symbiotic relationship between the archaea and bacteria created the nucleus-containing eukaryotic cell. (Organisms of the Archaea and Bacteria domain have no cell nucleus.[74]) It is hypothesized that the symbiosis originated when ancient archaea, similar to modern methanogenic archaea, invaded and lived within bacteria similar to modern myxobacteria, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and chloroplasts, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria.[75] The archaeal origin of the nucleus is supported by observations that archaea and eukarya have similar genes for certain proteins, including histones. Observations that myxobacteria are motile, can form multicellular complexes, and possess kinases and G proteins similar to eukarya, support a bacterial origin for the eukaryotic cell.[76]

text: A second model proposes that proto-eukaryotic cells evolved from bacteria without an endosymbiotic stage. This model is based on the existence of modern planctomycetes bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures.[77] A similar proposal states that a eukaryote-like cell, the chronocyte, evolved first and phagocytosed archaea and bacteria to generate the nucleus and the eukaryotic cell.[78]

text: The most controversial model, known as viral eukaryogenesis, posits that the membrane-bound nucleus, along with other eukaryotic features, originated from the infection of a prokaryote by a virus. The suggestion is based on similarities between eukaryotes and viruses such as linear DNA strands, mRNA capping, and tight binding to proteins (analogizing histones to viral envelopes). One version of the proposal suggests that the nucleus evolved in concert with phagocytosis to form an early cellular "predator".[79] Another variant proposes that eukaryotes originated from early archaea infected by poxviruses, on the basis of observed similarity between the DNA polymerases in modern poxviruses and eukaryotes.[80][81] It has been suggested that the unresolved question of the evolution of sex could be related to the viral eukaryogenesis hypothesis.[82]

text: A more recent proposal, the exomembrane hypothesis, suggests that the nucleus instead originated from a single ancestral cell that evolved a second exterior cell membrane; the interior membrane enclosing the original cell then became the nuclear membrane and evolved increasingly elaborate pore structures for passage of internally synthesized cellular components such as ribosomal subunits.[83]