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Life (1999 film)
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Life
Theatrical release poster
Directed by
Ted Demme
Produced by
Brian Grazer
Eddie Murphy
Screenplay by
Robert Ramsey
Matthew Sto
Story by
Eddie Murphy
Starring
Eddie Murphy
Martin Lawrence
Music by
Basil Poledouris
R. Kelly
Wyclef Jean
Cinematography
Geoffrey Simpson
Edited by
Jeffrey Wolf
Production
company
Imagine Entertainment
Distributed by
Universal Pictures
Release date
April 16, 1999 (1999-04-16)
Running time
109 minutes
Country
United States
Language
English
Budget
$80 million
Box office
$73,345,029
Life is a 1999 American comedy-drama film written by Robert Ramsey and Matthew Stone and directed by Ted Demme. The film stars Eddie Murphy and Martin Lawrence. It is the second film that Eddie Murphy and Martin Lawrence have worked on, the first being Boomerang. The supporting cast includes Obba Babatundé, Bernie Mac, Anthony Anderson, Miguel A. Núñez Jr., Bokeem Woodbine, Guy Torry and Barry Shabaka Henley. The film's format is a story being told by an elderly inmate about two of his friends, who are both wrongly convicted of murder and given a life sentence in prison. As of 2018[update], Life stands as the last R-rated feature to date for Eddie Murphy, whose subsequent works have earned either PG or PG-13 ratings.
Contents
1 Plot
2 Cast
3 Box office
4 Reception
5 Location
6 Soundtrack
7 Awards and nominations
8 References
9 External links
Plot[edit]
Elderly inmate Willie Long (Obba Babatundé) attends the burial of two friends who recently perished in an infirmary fire in a Mississippi prison. He begins telling the two young inmates digging the graves his friends' life story.
Ray Gibson (Eddie Murphy) and Claude Banks (Martin Lawrence) are two New Yorkers in 1932 from two different worlds. Ray is a small-time hustler and petty thief, and Claude, an honest, yet often selfish minded man, has just been accepted for a job as a bank teller at First Federal of Manhattan. They are both at a club called Spanky's when Ray picks Claude as his mark to pick-pocket. They both end up in the bad graces of the club's owner Spanky (Rick James), and Ray arranges for himself and Claude to do some boot-legging to pay off their debt, heading down south in order to buy a carload of Mississippi 'hooch'. Before they can get back to New York, a man named Winston Hancock (Clarence Williams III), who swindled Ray in a card game, is murdered outside of a juke joint by the town's sheriff, Warren Pike (Ned Vaughn), who frames Ray and Claude for the murder. A short time later, they go to trial, are convicted, and sentenced to life.
Ray and Claude are sent to an infamous prison camp called 'Camp 8' (now Mississippi State Penitentiary) to perform hard labor. They spend the next 65 years trying to escape from prison, while making new friends - Biscuit (Miguel A. Núñez Jr.), Jangle Leg (Bernie Mac), Radio (Guy Torry), Goldmouth (Michael Taliferro), Cookie (Anthony Anderson) and Pokerface (Barry Shabaka Henley) - and dodging guards Sergeant Dillard (Nick Cassavetes) and Hoppin' Bob (Brent Jennings) as their own friendship grows. Claude has his attorney cousin, Melvin, appeal against his conviction with his girlfriend Daisy's help, but it is denied, and shortly afterwards he finds out that Daisy has left him for Melvin. With any chance of being freed gone, Claude attempts to escape one night with Ray, getting as far as Tallahatchie before being captured and sentenced to a week in solitary confinement.
Around 1944, 12 years later, aged 37 years old, they meet a mute inmate nicknamed 'Can't-Get-Right' (Bokeem Woodbine), a talented baseball player and likely was the one to impregnate the Superintendent's daughter when he discovered the baby is biracial. He catches the eye of a Negro League scout who states that he can get him out of prison if he will play baseball. Ray and Claude, seeing an opportunity to freedom, try to convince the scout to help let them out as well, as they relate to 'Can't-Get-Right' and get him to play best. During a dance social, Biscuit confides to Ray that he is going to be released; however, Biscuit is afraid to return to his family, being gay, and committs suicide by deliberately running across the "firing line", getting shot and killed by the guard. After 'Can't-Get-Right' is released to play for the Pittsburgh Crawfords, Ray devises an escape but Claude refuses, upset with the fact that 'Can't-Get-Right' was released without them, leading to an argument that results in Ray and Claude going their separate ways.
Twenty-eight years later, in 1972, Ray and Claude are now 65 years old. All of the duo's friends are gone either by death or release from prison, except for Willie, now confined to a wheelchair. Dillard still runs the camp, who one day informs Ray and Claude that they will be transferred to live and work at Superintendent Dexter Wilkins' (Ned Beatty) mansion. Claude forms a friendship with Wilkins, and is entrusted to drive and pick up the new superintendent, Sheriff Warren Pike (R. Lee Ermey), the man who framed them forty years earlier. While on a pheasant hunt, Ray confronts Pike, where the sheriff admits to framing him and Claude with no remorse saying that "The state of Mississippi got forty years of cheap labor out of the deal." As Claude struggles to stop Ray from taking revenge, Pike attempts to kill them both, but he is shot and killed by Wilkins, realizing that Ray and Claude are indeed innocent, covering it up as a hunting accident. He intends to give them a pardon, but dies of a heart attack before he can do so.
In 1997 (present day), Ray and Claude are elderly, aged 90 years, living in the prison's infirmary. Claude tells Ray of yet another plan he has devised, but Ray has accepted his fate of dying in prison. That night, the infirmary catches fire, and they seemingly perish in the flames. Willie concludes the tale by outlining Claude's plan: Ray and Claude would steal two bodies from the morgue, start the blaze, plant the bodies, hide in the fire trucks and depart with them in the morning. When the workers ask why the plan didn't work, Willie tells them that he "never said it didn't work", as the inmates begin to realize that the bodies they buried are not Ray and Claude, who have escaped back to New York, watching a New York Yankees baseball game. The film concludes by revealing the two are again on good terms, living together in Harlem.
Cast[edit]
Eddie Murphy as Rayford "Ray" Gibson
Martin Lawrence as Claude Banks
Obba Babatundé as Willie Long
Ned Beatty as Dexter Wilkins
Nick Cassavetes as Sergeant Dillard
Bernie Mac as Jangle Leg
Miguel A. Núñez Jr. as Biscuit
Bokeem Woodbine as Can't Get Right
Brent Jennings as Hoppin' Bob
Anthony Anderson as Cookie
Barry Shabaka Henley as Pokerface
Michael Taliferro as Goldmouth
Guy Torry as Radio
Sanaa Lathan as Daisy
O'Neal Compton as Superintendent Abernathy
Noah Emmerich as Stan Blocker
Rick James as Spanky
R. Lee Ermey as Older Sheriff Pike
Ned Vaughn as Younger Sheriff Pike
Clarence Williams III as Winston Hancock
Heavy D as Jake
Kenn Whitaker as Issac
Bonz Malone as Leon
Lisa Nicole Carson as Sylvia
Poppy Montgomery as Older Mae Rose
Johnny Brown as Blind Reverend Clay
Don Harvey as Billy Bob
Box office[edit]
Life was released on April 16, 1999 in North America. The film grossed $73,345,029 worldwide against an $80 million budget, making it a financial disappointment.[1][2]
Reception[edit]
The film has received mixed to positive reviews and currently has a 50% rating on Rotten Tomatoes. The site's critic consensus reads, "Entertaining if not over-the-top humor from a solid comic duo provides plenty of laughs."[3]
Location[edit]
Even though Life was set in Jackson, Mississippi, Life was filmed in California;[4] filming locations include Brentwood, CA, Locke, CA, Los Angeles, Downey, CA, and Sacramento, CA. Parts of the film were shot at a Rockwell Defense Plant in California.[citation needed]
Soundtrack[edit]
Main article: Life (soundtrack)
A soundtrack containing hip hop and R&B music was released on March 16, 1999 on Rock Land/Interscope Records. It peaked at 10 on the Billboard 200 and 2 on the Top R&B/Hip-Hop Albums and was certified platinum with over 1 million copies sold on June 18, 1999.
Awards and nominations[edit]
Academy Award
nominated for Best Makeup (2000)
NAACP Image Award
nominated for Outstanding Motion Picture (2000)
BMI Film & TV Awards
(won) for Most Performed Song from a Film (2000)
Blockbuster Entertainment Awards
nominated with Eddie Murphy for Favorite Comedy Team (2000) for the movie
nominated for Favorite Song from a Movie (Fortunate)
References[edit]
Jump up ^ "Eddie Murphy's Charmed 'Life'". Los Angeles Times. Retrieved 2012-05-31.
Jump up ^ http://boxofficemojo.com/movies/?id=life.htm
Jump up ^ "Life (1999)". Rotten Tomatoes. Retrieved November 12, 2010.
Jump up ^ Cheseborough, Steve, Blues Traveling: The Holy Sites of Delta Blues. University Press of Mississippi, 2004. 96. Retrieved from Google Books on September 29, 2010. ISBN 1-57806-650-6, ISBN 978-1-57806-650-6.
External links[edit]
Life on IMDb
Life at Box Office Mojo
show
v
t
e
Films directed by Ted Demme
The Bet (1992)
Who's the Man? (1993)
The Ref (1994)
Beautiful Girls (1996)
Monument Ave. (1998)
Life (1999)
Blow (2001)
A Decade Under the Influence (2003)
show
v
t
e
Brian Grazer
Films produced
Night Shift (1982)
Splash (1984)
Real Genius (1985)
Spies Like Us (1985)
Armed and Dangerous (1986)
Like Father Like Son (1987)
Parenthood (1989)
Kindergarten Cop (1990)
Closet Land (1991)
My Girl (1991)
Far and Away (1992)
Housesitter (1992)
Boomerang (1992)
For Love or Money (1993)
My Girl 2 (1994)
Greedy (1994)
The Paper (1994)
The Cowboy Way (1994)
Apollo 13 (1995)
Sgt. Bilko (1996)
Fear (1996)
The Nutty Professor (1996)
The Chamber (1996)
Ransom (1996)
Liar Liar (1997)
Inventing the Abbotts (1997)
Mercury Rising (1998)
Psycho (1998)
Edtv (1999)
Life (1999)
Bowfinger (1999)
Beyond the Mat (2000)
Nutty Professor II: The Klumps (2000)
How the Grinch Stole Christmas (2000)
A Beautiful Mind (2001)
Undercover Brother (2002)
Blue Crush (2002)
8 Mile (2002)
Intolerable Cruelty (2003)
The Cat in the Hat (2003)
The Missing (2003)
Friday Night Lights (2004)
Inside Deep Throat (2005)
Cinderella Man (2005)
Flightplan (2005)
Fun with Dick and Jane (2005)
Inside Man (2006)
The Da Vinci Code (2006)
American Gangster (2007)
Changeling (2008)
Frost/Nixon (2008)
Angels & Demons (2009)
Robin Hood (2010)
The Dilemma (2011)
Restless (2011)
Cowboys & Aliens (2011)
J. Edgar (2011)
Tower Heist (2011)
Katy Perry: Part of Me (2012)
Rush (2013)
In the Heart of the Sea (2015)
Inferno (2016)
Lowriders (2016)
American Made (2017)
TV series created
Shadow Chasers
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text:
The Lord of the Rings: The Two Towers (soundtrack)
From Wikipedia, the free encyclopedia
Jump to: navigation, search
The Lord of the Rings: The Two Towers: Original Motion Picture Soundtrack
Soundtrack album by Howard Shore
Released
December 10, 2002 (2002-12-10)
Genre
Soundtrack
Length
72:46
77:09 (limited edition)
Label
Reprise
Producer
Howard Shore
Middle Earth film series soundtrack chronology
The Lord of the Rings: The Fellowship of the Ring
(2001)The Lord of the Rings: The Fellowship of the Ring2001 The Lord of the Rings: The Two Towers
(2002) The Lord of the Rings: The Return of the King
(2003)The Lord of the Rings: The Return of the King2003
The Lord of the Rings: The Two Towers: Original Motion Picture Soundtrack was released on December 10, 2002. It was composed, orchestrated, and conducted by Howard Shore, and performed by the London Philharmonic Orchestra, the London Voices, and the London Oratory School Schola.[1] The Two Towers comes in a regular and a limited edition, the latter containing the additional track "Farewell to Lórien".
Contents
[hide]
1 Reception
2 Track listing
3 Charts and certifications
3.1 Charts
3.2 Certifications
4 Complete Recording and additional music
4.1 Additional Music
5 See also
6 References
Reception[edit]
Professional ratings
Review scores
Source
Rating
AllMusic [2]
Filmtracks [3]
Movie Music UK [4]
Tracksounds [5]
The Two Towers won the Grammy Award for Best Score Soundtrack Album.
Track listing[edit]
All music composed by Howard Shore.
No.
Title
Length
1. "Foundations of Stone" 3:51
2. "The Taming of Sméagol" 2:48
3. "The Riders of Rohan" 4:05
4. "The Passage of the Marshes" 2:46
5. "The Uruk-hai" 2:58
6. "The King of the Golden Hall" 3:49
7. "The Black Gate Is Closed" 3:17
8. "Evenstar" (feat. Isabel Bayrakdarian) 3:15
9. "The White Rider" 2:28
10. "Treebeard" 2:43
11. "The Leave Taking" 3:41
12. "Helm's Deep" 3:53
13. "The Forbidden Pool" 5:27
14. "Breath of Life" (feat. Sheila Chandra) 5:07
15. "The Hornburg" 4:36
16. "Forth Eorlingas" (feat. Ben Del Maestro) 3:15
17. "Isengard Unleashed" (feat. Elizabeth Fraser and Ben Del Maestro) 5:01
18. "Samwise the Brave" 3:46
19. "Gollum's Song" (performed by Emilíana Torrini, lyrics by Fran Walsh) 5:51
Total length: 72:46
Limited edition bonus track
No.
Title
Length
20. "Farewell to Lórien" (feat. Hilary Summers) 4:37
Charts and certifications[edit]
Charts[edit]
Chart (2002-03)
Peak
position
Australian Albums (ARIA)[6]
31
Austrian Albums (Ö3 Austria)[7]
11
Belgian Albums (Ultratop Flanders)[8]
33
Belgian Albums (Ultratop Wallonia)[9]
19
Finnish Albums (Suomen virallinen lista)[10]
3
French Albums (SNEP)[11]
35
German Albums (Offizielle Top 100)[12]
8
Hungarian Albums (Mahasz)[13]
12
Irish Albums (IRMA)[14]
47
Irish Classical Albums (IRMA)[15]
1
Dutch Albums (MegaCharts)[16]
20
Norwegian Albums (VG-lista)[17]
18
Swedish Albums (Sverigetopplistan)[18]
16
Swiss Albums (Schweizer Hitparade)[19]
14
UK Albums (OCC)[20]
28
US Billboard 200[21]
43
US Soundtrack Albums (Billboard)[22]
2
Certifications[edit]
Region
Certification
Certified units/Sales
Canada (Music Canada)[23]
Gold 50,000^
United Kingdom (BPI)[24]
Gold 100,000^
United States (RIAA)[25]
Gold 500,000^
^shipments figures based on certification alone
Complete Recording and additional music[edit]
In 2006, Reprise Records released a multi-disc set for the film, titled The Complete Recordings. These contain the entire score for the extended versions of the film on CD, along with an additional DVD-Audio disc that offers 2.0 stereo and 5.1 surround mixes of the soundtrack. The album also featured extensive liner notes by music journalist Doug Adams which reviews all of the tracks and provides information about the process of composing and recording the score, as well as a detailed list of all musical instruments, people and organizations involved. The cover artwork uses the film series' logo and an inscription in Tolkien's tengwar letters, over a background that depicts Rohan and Fangorn in dark blue.
Disc one
No.
Title
Length
1. "Glamdring" 3:50
2. "Elven Rope" 2:19
3. "Lost in Emyn Muil" 4:15
4. "My Precious" 2:56
5. "Uglúk's Warriors" 1:41
6. "The Three Hunters" 6:12
7. "The Banishment of Éomer" 3:55
8. "Night Camp" 2:50
9. "The Plains of Rohan" 4:15
10. "Fangorn" 5:13
11. "The Dead Marshes" 5:08
12. ""Wraiths on Wings"" 2:08
13. "Gandalf the White" 6:48
14. "The Dreams of Trees" 1:55
15. "The Heir of Númenor" 6:51
16. "Ent-Draught" 2:54
Total length: 63:10
Disc two
No.
Title
Length
1. "Edoras" 4:34
2. "The Court of Meduseld" 3:11
3. "Théoden King" (feat. "The Funeral of Théodred", performed by Miranda Otto) 6:12
4. "The King's Decision" 2:08
5. "Exodus from Edoras" 5:43
6. "The Forests of Ithilien" 6:38
7. "One of the Dúnedain" (feat. "Evenstar", performed by Isabel Bayrakdarian) 7:13
8. "The Wolves of Isengard" 4:22
9. "Refuge at Helm's Deep" 4:00
10. "The Voice of Saruman" 1:12
11. "Arwen's Fate" (feat. "The Grace of the Valar", performed by Sheila Chandra) 3:59
12. "The Story Foretold" 3:39
13. "Sons of the Steward" 6:03
14. "Rock and Pool" 2:55
15. "Faramir's Good Council" 2:21
Total length: 63:59
Disc three
No.
Title
Length
1. "Aragorn's Return" 2:12
2. "War Is Upon Us" 3:36
3. ""Where Is the Horse and the Rider?"" 6:16
4. "The Host of the Eldar" 2:51
5. "The Battle of the Hornburg" 2:53
6. "The Breach of the Deeping Wall" 3:03
7. "The Entmoot Decides" 2:06
8. "Retreat" (feat. "Haldir's Lament", performed by Elizabeth Fraser) 4:41
9. "Master Peregrin's Plan" 2:32
10. "The Last March of the Ents" (feat. Ben Del Maestro) 2:31
11. "The Nazgûl Attack" 2:45
12. "Théoden Rides Forth" (feat. Ben Del Maestro) 5:48
13. "The Tales That Really Matter" 12:01
14. ""Long Ways to Go Yet"" (feat. "Gollum's Song", performed by Emilíana Torrini) 8:06
Total length: 61:12
Additional Music[edit]
Additional music for the film was featured in The Rarities Archive release, attached to Doug Adams' book on the three film scores:
Track listing
No.
Title
Length
1. ""Gwenwin in în ("Arwen's Song" Alternate/Mock-up)"" 2:02
2. ""Arwen's Song (Complete)"" 2:11
3. ""Emyn Muil (Alternate)"" 3:23
4. "Flight to the Ford (Alternate)" 4:04
5. ""The Rohan Fanfare (Mock-up)"" 3:09
6. ""The Eaves of Fangorn (Alternate)"" 5:25
7. ""The Ent Theme (Mock-up)"" 2:00
Total length: 22:14
Along with about 4 minutes of alternate material from the original release,[26] about 8 minutes of alternates in the fan credits,[27] and some additional alternates, there are about 3:40 minute of finalized music from The Two Towers.
See also[edit]
Music of The Lord of the Rings film series
References[edit]
Jump up ^ The Lord of the Rings: The Two Towers: Original Motion Picture Soundtrack (CD liner notes). Howard Shore. Reprise Records. 2002.
Jump up ^ Fennessy, Kathleen C.. The Lord of the Rings: The Two Towers [Original Motion Picture Soundtrack] at AllMusic. Retrieved 3 January 2014.
Jump up ^ Clemmensen, Christian (11 December 2002). "The Lord of the Rings: The Two Towers (Howard Shore)". Filmtracks. Retrieved 3 January 2014.
Jump up ^ Broxton, Jonathan (20 December 2002). "THE LORD OF THE RINGS: THE TWO TOWERS – Howard Shore". Movie Music UK. Retrieved 3 January 2014.
Jump up ^ Coleman, Christopher. "Score of the Rings: Part II". Tracksounds. Retrieved 3 January 2014.
Jump up ^ "Australiancharts.com – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". Hung Medien.
Jump up ^ "Austriancharts.at – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers" (in German). Hung Medien.
Jump up ^ "Ultratop.be – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers" (in Dutch). Hung Medien.
Jump up ^ "Ultratop.be – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers" (in French). Hung Medien.
Jump up ^ "Soundtrack / Howard Shore: The Lord of the Rings - The Two Towers" (in Finnish). Musiikkituottajat – IFPI Finland.
Jump up ^ "Lescharts.com – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". Hung Medien.
Jump up ^ "Officialcharts.de – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". GfK Entertainment Charts.
Jump up ^ "Archívum › Kereső - előadó/cím szerint" (in Hungarian). Association of Hungarian Record Companies.
Jump up ^ "GFK Chart-Track Albums: Week 4, 2003". Chart-Track. IRMA.
Jump up ^ "GFK Chart-Track – Classical Albums: Week 4, 2003". Chart-Track. IRMA.
Jump up ^ "Dutchcharts.nl – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers" (in Dutch). Hung Medien.
Jump up ^ "Norwegiancharts.com – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". Hung Medien.
Jump up ^ "Swedishcharts.com – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". Hung Medien.
Jump up ^ "Swisscharts.com – Soundtrack / Howard Shore – The Lord of the Rings - The Two Towers". Hung Medien.
Jump up ^ "Official Albums Chart Top 100". Official Charts Company.
Jump up ^ "Howard Shore Chart History (Billboard 200)". Billboard.
Jump up ^ "The Lord of the Rings: The Two Towers [Original Motion Picture Soundtrack] - Howard Shore – Chart History". Billboard. Prometheus Global Media.
Jump up ^ "Canadian album certifications – Various Artists – The Lord of the Rings: The Two Towers". Music Canada. Retrieved December 21, 2012.
Jump up ^ "British album certifications – Original Soundtrack – The Lord of the Rings - The Two Towers - OST". British Phonographic Industry. Retrieved December 21, 2012. Enter The Lord of the Rings - The Two Towers - OST in the search field and then press Enter.
Jump up ^ "American album certifications – Soundtrack – The Lord of the Rings: The Two Towers". Recording Industry Association of America. Retrieved December 21, 2012. If necessary, click Advanced, then click Format, then select Album, then click SEARCH
Jump up ^ http://www.lyndhursthouse.plus.com/ttt_ost_breakdown.pdf
Jump up ^ http://www.amagpiesnest.com/cbc_analysis/ttt/tcredit_music.htm
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Gravitropism
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A straight and vertical tree. The shape has been regulated by the gravitropic curvature at the base.
Gravitropism (also known as geotropism) is a turning or growth movement by a plant or fungus in response to gravity. It is a general feature of all higher and many lower plants as well as other organisms. Charles Darwin was one of the first to scientifically document that roots show positive gravitropism and stems show negative gravitropism. That is, roots grow in the direction of gravitational pull (i.e., downward) and stems grow in the opposite direction (i.e., upwards). This behavior can be easily demonstrated with any potted plant. When laid onto its side, the growing parts of the stem begin to display negative gravitropism, growing (biologists say, turning; see tropism) upwards. Herbaceous (non-woody) stems are capable of a small degree of actual bending, but most of the redirected movement occurs as a consequence of root or stem growth outside.
Contents
[hide]
1 Gravitropism in the root
2 Gravitropism in shoots
3 Gravity Sensing Mechanisms
3.1 Statoliths
3.2 Modulation by phytochrome
4 Compensation
5 Gravitropic mutants
6 See also
7 References
Gravitropism in the root[edit]
Root growth occurs by division of stem cells in the root meristem located in the tip of the root, and the subsequent expansion of cells in a region just primal to the tip known as the elongation zone. Differential growth during tropisms mainly involves changes in cell expansion versus changes in cell division, although a role for cell division in tropic growth has not been formally ruled out. Gravity is sensed in the root tip and this information must then be relayed to the elongation zone so as to maintain growth direction and mount effective growth responses to changes in orientation and continue to grow its roots in the same direction as gravity.[1]
Abundant evidence demonstrates that roots bend in response to gravity due to a regulated movement of the plant hormone auxin known as polar auxin transport.[2] This was described in the 1920s in the Cholodny-Went model. The model was independently proposed by the Russian scientist N. Cholodny of the University of Kiev in 1927 and by Frits Went of the California Institute of Technology in 1928, both based on work they had done in 1926.[3] Auxin exists in nearly every organ and tissue of a plant, but its has been reoriented in the gravity field, can initiate differential growth resulting in root curvature.
Experiments show that auxin distribution is characterized by a fast movement of auxin to the lower side of the root in response to a gravity stimulus at a 90° degree angle or more. However, once the root tip reaches a 40° angle to the horizontal of the stimulus, auxin distribution quickly shifts to a more symmetrical arrangement. This behavior is described as a "tipping point" mechanism for auxin transport in response to a gravitational stimulus.[4]
Gravitropism in shoots[edit]
Play media
Apex reorientation in Pinus pinaster during the first 24h after experimental inclination of the plant.
Gravitropism is an integral part of plant growth, orienting its position to maximize contact with sunlight, as well as ensuring that the roots are growing in the correct direction. Growth due to gravitropism is mediated by changes in concentration of the plant hormone auxin within plant cells.
As plants mature, gravitropism continues to guide growth and development along with phototropism. While amyloplasts continue to guide plants in the right direction, plant organs and function rely on phototropic responses to ensure that the leaves are receiving enough light to perform basic functions such as photosynthesis. In complete darkness, mature plants have little to no sense of gravity, unlike seedlings that can still orient themselves to have the shoots grow upward until light is reached when development can begin.[5]
Differential sensitivity to auxin helps explain Darwin's original observation that stems and roots respond in the opposite way to the forces of gravity. In both roots and stems, auxin accumulates towards the gravity vector on the lower side. In roots, this results in the inhibition of cell expansion on the lower side and the concomitant curvature of the roots towards gravity (positive gravitropism).[1][6] In stems, the auxin also accumulates on the lower side, however in this tissue it increases cell expansion and results in the shoot curving up (negative gravitropism).[7]
A recent study showed that for gravitropism to occur in shoots, only a fraction of an inclination, instead of a strong gravitational force, is necessary. This finding sets aside gravity sensing mechanisms that would rely on detecting the pressure of the weight of statoliths.[8]
Gravity Sensing Mechanisms[edit]
Statoliths[edit]
Plants possess the ability to sense gravity in several ways, one of which is through statoliths. Statoliths are dense amyloplasts, organelles that synthesize and store starch involved in the perception of gravity by the plant (gravitropism), that collect in specialized cells called statocytes. Statocytes are located in the starch parenchyma cells near vascular tissues in the shoots and in the columella in the caps of the roots.[9] These specialized amyloplasts are denser than the cytoplasm and can sediment according to the gravity vector. The statoliths are enmeshed in a web of actin and it is thought that their sedimentation transmits the gravitropic signal by activating mechanosensitive channels.[1] The gravitropic signal then leads to the reorientation of auxin efflux carriers and subsequent redistribution of auxin streams in the root cap and root as a whole.[10] The changed relations in concentration of auxin leads to differential growth of the root tissues. Taken together, the root is then turning to follow the gravity stimuli. Statoliths are also found in the endodermic layer of the hypocotyl, stem, and inflorescence stock. The redistribution of auxin causes increased growth on the lower side of the shoot so that it orients in a direction opposite that of the gravity stimuli.
Modulation by phytochrome[edit]
Phytochromes are red and far-red photoreceptors that help induce changes in certain aspects of plant development. Apart being itself the tropic factor (phototropism), light may also suppress the gravitropic reaction.[11] In seedlings, red and far-red light both inhibit negative gravitropism in seedling hypocotyls (the shoot area below the cotyledons) causing growth in random directions. However, the hypocotyls readily orient towards blue light. This process is may be caused by phytochrome disrupting the formation of starch-filled endodermal amyloplasts and stimulating their conversion to other plastid types, such as chloroplasts or etiolaplasts.[11]
Compensation[edit]
The compensation reaction of the bending Coprinus stem. C - the compensating part of the stem.
Bending mushroom stems follow some regularities that are not common in plants. After turning into horizontal the normal vertical orientation the apical part (region C in the figure below) starts to straighten. Finally this part gets straight again, and the curvature concentrates near the base of the mushroom.[12] This effect is called compensation (or sometimes, autotropism). The exact reason of such behavior is unclear, and at least two hypotheses exist.
The hypothesis of plagiogravitropic reaction supposes some mechanism that sets the optimal orientation angle other than 90 degrees (vertical). The actual optimal angle is a multi-parameter function, depending on time, the current reorientation angle and from the distance to the base of the fungi. The mathematical model, written following this suggestion, can simulate bending from the horizontal into vertical position but fails to imitate realistic behavior when bending from the arbitrary reorientation angle (with unchanged model parameters).[12]
The alternative model supposes some “straightening signal”, proportional to the local curvature. When the tip angle approaches 30° this signal overcomes the bending signal, caused by reorientation, straightening resulting.[13]
Both models fit the initial data well, but the latter was also able to predict bending from various reorientation angles. Compensation is less obvious in plants, but in some cases it can be observed combining exact measurements with mathematical models. The more sensitive roots are stimulated by lower levels of auxin; higher levels of auxin in lower halves stimulate less growth, resulting in downward curvature (positive gravitropism).
Gravitropic mutants[edit]
Mutants with altered responses to gravity have been isolated in several plant species including Arabidopsis thaliana (one of the genetic model systems used for plant research). These mutants have alterations in either negative gravitropism in hypocotyls and/or shoots, or positive gravitropism in roots, or both.[7] Mutants have been identified with varying effects on the gravitropic responses in each organ, including mutants which nearly eliminate gravitropic growth, and those whose effects are weak or conditional. Once a mutant has been identified, it can be studied to determine the nature of the defect (the particular difference(s) it has compared to the non-mutant 'wildtype'). This can provide information about the function of the altered gene, and often about the process under study. In addition the mutated gene can be identified, and thus something about its function inferred from the mutant phenotype.
Gravitropic mutants have been identified that affect starch accumulation, such as those affecting the PGM1 (which encodes the enzyme phosphoglucomutase) gene in Arabidopsis, causing plastids - the presumptive statoliths - to be less dense and, in support of the starch-statolith hypothesis, less sensitive to gravity.[14] Other examples of gravitropic mutants include those affecting the transport or response to the hormone auxin.[7] In addition to the information about gravitropism which such auxin-transport or auxin-response mutants provide, they have been instrumental in identifying the mechanisms governing the transport and cellular action of auxin as well as its effects on growth.
There are also several cultivated plants that display altered gravitropism compared to other species or to other varieties within their own species. Some are trees that have a weeping or pendulate growth habit; the branches still respond to gravity, but with a positive response, rather than the normal negative response. Others are the lazy (i.e. ageotropic or agravitropic) varieties of corn (Zea mays) and varieties of rice, barley and tomatoes, whose shoots grow along the ground.
See also[edit]
Plants portal
Gardening portal
Clinostat - a device used to negate the effects of gravitational pull.
Amyloplast - starch organelle involved in sensing gravitropism
References[edit]
^ Jump up to: a b c PERRIN, ROBYN M.; YOUNG, LI-SEN; NARAYANA MURTHY, U.M.; HARRISON, BENJAMIN R.; WANG, YAN; WILL, JESSICA L.; MASSON, PATRICK H. (2017-04-21). "Gravity Signal Transduction in Primary Roots". Annals of Botany. 96 (5): 737–743. ISSN 0305-7364. PMC 4247041  . PMID 16033778. doi:10.1093/aob/mci227.
Jump up ^ Swarup, Ranjan; Kramer, Eric M.; Perry, Paula; Knox, Kirsten; Leyser, H. M. Ottoline; Haseloff, Jim; Beemster, Gerrit T. S.; Bhalerao, Rishikesh; Bennett, Malcolm J. (2005-11-01). "Root gravitropism requires lateral root cap and epidermal cells for transport and response to a mobile auxin signal". Nature Cell Biology. 7 (11): 1057–1065. ISSN 1465-7392. PMID 16244669. doi:10.1038/ncb1316.
Jump up ^ Janick, Jules (2010). Horticultural Reviews. John Wiley & Sons. p. 235. ISBN 0470650532.
Jump up ^ Band, L. R.; Wells, D. M.; Larrieu, A.; Sun, J.; Middleton, A. M.; French, A. P.; Brunoud, G.; Sato, E. M.; Wilson, M. H.; Peret, B.; Oliva, M.; Swarup, R.; Sairanen, I.; Parry, G.; Ljung, K.; Beeckman, T.; Garibaldi, J. M.; Estelle, M.; Owen, M. R.; Vissenberg, K.; Hodgman, T. C.; Pridmore, T. P.; King, J. R.; Vernoux, T.; Bennett, M. J. (5 March 2012). "Root gravitropism is regulated by a transient lateral auxin gradient controlled by a tipping-point mechanism". Proceedings of the National Academy of Sciences. 109 (12): 4668–4673. doi:10.1073/pnas.1201498109.
Jump up ^ Hangarter, R.P. (1997). "Gravity, light, and plant form". Plant, Cell, and Environment. 20: 796–800.
Jump up ^ Hou, G., Kramer, V. L., Wang, Y.-S., Chen, R., Perbal, G., Gilroy, S. and Blancaflor, E. B. (2004). "The promotion of gravitropism in Arabidopsis roots upon actin disruption is coupled with the extended alkalinization of the columella cytoplasm and a persistent lateral auxin gradient". The Plant Journal. 39: 113–125. CS1 maint: Multiple names: authors list (link)
^ Jump up to: a b c Masson, Patrick H.; Tasaka, Masao; Morita, Miyo T.; Guan, Changhui; Chen, Rujin; Boonsirichai, Kanokporn (2002-01-01). "Arabidopsis thaliana: A Model for the Study of Root and Shoot Gravitropism". The Arabidopsis Book: e0043. PMC 3243349  . PMID 22303208. doi:10.1199/tab.0043.
Jump up ^ Chauvet, Hugo; Pouliquen, Olivier; Forterre, Yoël; Legué, Valérie; Moulia, Bruno (14 October 2016). "Inclination not force is sensed by plants during shoot gravitropism". Scientific Reports. 6 (1). doi:10.1038/srep35431.
Jump up ^ Chen, Rujin; Rosen, Elizabeth; Masson, Patrick H. (1 June 1999). "Gravitropism in Higher Plants". Plant Physiology. 120 (2): 343–350. doi:10.1104/pp.120.2.343.
Jump up ^ Sato, Ethel Mendocilla; Hijazi, Hussein; Bennett, Malcolm J.; Vissenberg, Kris; Swarup, Ranjan (2015-04-01). "New insights into root gravitropic signalling". Journal of Experimental Botany. 66 (8): 2155–2165. ISSN 0022-0957. doi:10.1093/jxb/eru515.
^ Jump up to: a b Kim, Keunhwa; Shin, Jieun; Lee, Sang-Hee; Kweon, Hee-Seok; Maloof, Julin N.; Choi, Giltsu (2011-01-25). "Phytochromes inhibit hypocotyl negative gravitropism by regulating the development of endodermal amyloplasts through phytochrome-interacting factors". Proceedings of the National Academy of Sciences. 108 (4): 1729–1734. ISSN 0027-8424. PMC 3029704  . PMID 21220341. doi:10.1073/pnas.1011066108.
^ Jump up to: a b Meškauskas, A., Novak Frazer, L. N., & Moore, D. (1999). "Mathematical modelling of morphogenesis in fungi: a key role for curvature compensation (‘autotropism’) in the local curvature distribution model". New Phytologist. 143: 387–399. CS1 maint: Multiple names: authors list (link)
Jump up ^ Meškauskas A., Jurkoniene S., Moore D (1999). "Spatial organization of the gravitropic response in plants: applicability of the revised local curvature distribution model to Triticum aestivum coleoptiles". New Phytologist. 143: 401–407. CS1 maint: Multiple names: authors list (link)
Jump up ^ Wolverton, Chris; Paya, Alex M.; Toska, Jonida (2011-04-01). "Root cap angle and gravitropic response rate are uncoupled in the Arabidopsis pgm-1 mutant". Physiologia Plantarum. 141 (4): 373–382. ISSN 1399-3054. doi:10.1111/j.1399-3054.2010.01439.x.
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Nastic movements (non-directional)
Nyctinasty
Thigmonasty
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