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Helter Skelter (song)
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For other songs of similar name, see Helter Skelter.
"Helter Skelter"
1976 US promotional single
Song by the Beatles
from the album The Beatles
Published
Northern Songs
Released
22 November 1968
Recorded
9–10 September 1968,
EMI Studios, London
Genre
Hard rock[1][2]
heavy metal[3]
proto-punk[4]
Length
4:29 (stereo LP)
3:40 (mono LP)
Label
Apple
Songwriter(s)
Lennon–McCartney
Producer(s)
George Martin
Audio sample
file
help
"Helter Skelter" is a song by the English rock band the Beatles that was released in 1968 on their self-titled double album, often known as "the White Album". It was written by Paul McCartney[5][6] and credited to Lennon–McCartney. The song was a product of McCartney's attempt to create a sound as loud and dirty as possible. The Beatles' recording has been noted for its "proto-metal roar" and is considered by music historians to be a key influence in the early development of heavy metal.[7] Rolling Stone magazine ranked "Helter Skelter" 52nd on its list of the "100 Greatest Beatles songs".[8]
Contents
1 Writing and inspiration
2 Recording
3 Critical reaction
4 Charles Manson
5 Cover versions
6 Paul McCartney live performances
7 Personnel
8 Notes
9 References
10 External links
Writing and inspiration[edit]
McCartney was inspired to write the song after reading a 1967 Guitar Player magazine interview with the Who's Pete Townshend where he described their latest single, "I Can See for Miles", as the loudest, rawest, dirtiest song the Who had ever recorded. McCartney then "wrote 'Helter Skelter' to be the most raucous vocal, the loudest drums, et cetera" and said he was "using the symbol of a helter skelter as a ride from the top to the bottom; the rise and fall of the Roman Empire – and this was the fall, the demise."[5] In British English, a helter skelter is an amusement park attraction which features a tall spiral slide winding round a tower.[9] McCartney has cited this song as a response to critics who accuse him of writing only ballads.[10]
On 20 November 1968, two days before the release of The Beatles, McCartney gave Radio Luxembourg an exclusive interview, in which he commented on several of the album's songs. Speaking of "Helter Skelter", he said: "Umm, that came about just 'cause I'd read a review of a record which said, 'and this group really got us wild, there's echo on everything, they're screaming their heads off.' And I just remember thinking, 'Oh, it'd be great to do one. Pity they've done it. Must be great – really screaming record.' And then I heard their record and it was quite straight, and it was very sort of sophisticated. It wasn't rough and screaming and tape echo at all. So I thought, 'Oh well, we'll do one like that, then.' And I had this song called 'Helter Skelter,' which is just a ridiculous song. So we did it like that, 'cuz I like noise."[11]
Recording[edit]
The song was recorded many times during sessions for The Beatles. During the 18 July 1968 sessions, the Beatles recorded a version of the song lasting 27 minutes and 11 seconds,[12] although this version is rather slow and hypnotic, differing greatly from the volume and rawness of the album version.[13] Another recording from the same day, originally 12 minutes long, was edited down to 4:37 for Anthology 3. On 9 September, 18 takes of approximately five minutes each were recorded, and the last one is featured on the original LP.[12] After the 18th take, Ringo Starr flung his drum sticks across the studio[14] and screamed, "I got blisters on my fingers!"[5][12] Starr's shout was included on the stereo mix of the song. At around 3:40, the song completely fades out, gradually fades back in, fades back out partially and finally fades back in quickly with three cymbal crashes and Ringo's scream (some sources erroneously credit the "blisters" line to Lennon; in fact, Lennon can be heard asking "How's that?" before Ringo's outburst).[15] The mono version (originally on LP only) ends on the first fadeout without Starr's outburst. The mono version was not initially available in the US as mono albums had already been phased out there. The mono version was later released in the American version of the Rarities album. In 2009, it was made available on the CD mono re-issue of The Beatles as part of the Beatles in Mono CD box set.
According to Chris Thomas, who was present,[12] the 9 September session was especially spirited: "While Paul was doing his vocal, George Harrison had set fire to an ashtray and was running around the studio with it above his head, doing an Arthur Brown."[12] Starr's recollection is less detailed, but agrees in spirit: "'Helter Skelter' was a track we did in total madness and hysterics in the studio. Sometimes you just had to shake out the jams."[10]
Critical reaction[edit]
Among music critics commenting on "Helter Skelter", Richie Unterberger of AllMusic views it as "one of [the] fiercest and most brutal rockers done by anyone" and "extraordinary".[16] Writing for MusicHound in 1999, Guitar World editor Christopher Scapelliti identified the track as one of three "fascinating standouts" on the White Album.[17] While admiring the diversity of McCartney's songwriting on the album, Mark Richardson of Pitchfork cites "Helter Skelter" as one of "the roughest, rawest tunes in his Beatles oeuvre".[18]
Ian MacDonald was highly critical of the song, however, calling it "ridiculous, McCartney shrieking weedily against a massively tape-echoed backdrop of out-of-tune thrashing".[19] Rob Sheffield was also unimpressed, writing in The Rolling Stone Album Guide (2004) that, following the double album's release on CD, "now you can program 'Sexy Sadie' and 'Long, Long, Long' without having to lift the needle to skip over 'Helter Skelter.'"[20] Alan W. Pollack said the song will "scare and unsettle" listeners, citing "Helter Skelter"'s "obsessive nature" and "undercurrent of violence", and noted McCartney's "savage vocal delivery" as reinforcing this theme.[21]
In a 1980 interview, Lennon said, "That's Paul completely ... It has nothing to do with anything, and least of all to do with me."[6]
In March 2005, Q magazine ranked "Helter Skelter" number 5 in its list of the 100 Greatest Guitar Tracks.
Charles Manson[edit]
Main article: Helter Skelter (Manson scenario)
Charles Manson told his followers that several White Album songs including "Helter Skelter" were a part of the Beatles' coded prophecy of an apocalyptic war in which racist and non-racist whites would be manoeuvred into virtually exterminating each other over the treatment of blacks.[22][23][24] Upon the war's conclusion, after black militants would kill off the few whites they would know to have survived, Manson and his companions would emerge from an underground city in which they would have escaped the conflict. As the only remaining whites, they would rule blacks, who, as the vision went, would be incapable of running America. Manson employed "helter skelter" as the term for this sequence of events.
Los Angeles Deputy District Attorney Vincent Bugliosi, who led the prosecution of Manson and four of his followers who acted on Manson's instruction in the Tate-LaBianca murders, named his best-selling book about the murders Helter Skelter.[22] The book was the basis for two television movies of the same title.
Cover versions[edit]
In 1975, Aerosmith recorded a cover of "Helter Skelter", but it was not released until 1991, on the Pandora's Box compilation.[25] The cover charted at #21 on the Mainstream Rock Tracks chart.
in 1978, Siouxsie and the Banshees released one of the first covers of the song on their debut album The Scream
in 1981, Pat Benatar included a cover of the song on her album Precious Time.
In 1983, the Bobs released an a cappella version on their eponymous album.[26] It earned them a 1984 Grammy nomination for Best Vocal Arrangement for Two or More Voices.[27]
In 1983, a remake of this song was also featured on the Mötley Crüe album Shout at the Devil.
In 1988, a U2 recording was used as the opening track on the Rattle and Hum album. The song was recorded live at the McNichols Sports Arena in Denver, Colorado on 8 November 1987.[28] Introducing the song, Bono controversially said, "This is a song Charles Manson stole from the Beatles. We’re stealing it back."[29]
In 1989, Bow Wow recorded "Helter Skelter", released it as a single and named their album after the song.
In 2000, Oasis recorded a cover of "Helter Skelter", originally as a B-side to the single release of "Who Feels Love?" They also performed the song on their world tour promoting their fourth album Standing on the Shoulder of Giants in the early 2000s; one live version was recorded for their 2001 live album Familiar to Millions.
In 2007, Dana Fuchs performs the song in Across the Universe.
In 2007, Stereophonics recorded a version of the song for their cd-single "It Means Nothing".
In 2007, Beatallica recorded a parody called "Helvester of Skelter", which also was a parody of the Metallica song "Harvester of Sorrow".
In 2018, Marilyn Manson and Rob Zombie together recorded a version of the song released on 11 July 2018.
Paul McCartney live performances[edit]
Since 2004 McCartney has performed the song with his band on every tour, starting on 24 May 2004, while on the '04 Summer Tour, through The 'US' Tour (2005), the Summer Live '09 (2009), the Good Evening Europe Tour (2009), the Up and Coming Tour (2010/2011), the On the Run Tour (2011/2012) and the Out There Tour, which started on 4 May 2013. In the last tours, the song has been generally inserted on the third encore, which is the last time the band enters the stage. It is usually the last but one song, performed after "Yesterday" and before the final medley including "The End". Paul played the song on his One on One Tour at Fenway Park on 17 July 2016 accompanied by the Grateful Dead's Bob Weir and New England Patriots football player Rob Gronkowski.
McCartney performed the song live at the 48th Annual Grammy Awards on 8 February 2006 at the Staples Center in Los Angeles. In 2009, McCartney performed the song live on top of the Ed Sullivan Theater during his appearance on the Late Show with David Letterman.
The version of the song from McCartney's live album Good Evening New York City, recorded during the Summer Live '09 tour, was nominated at the 53rd Grammy Awards in the category of Best Solo Rock Vocal Performance.[30] It won, becoming McCartney's first solo Grammy win since he won for arranging "Uncle Albert/Admiral Halsey" in 1972.[31]
McCartney opened his set at 12-12-12: The Concert for Sandy Relief with the song.[32]
Personnel[edit]
Paul McCartney – lead vocal, electric guitar, piano
John Lennon – backing vocal, six-string bass, electric guitar, sound effects (through brass instruments)
George Harrison – backing vocal, rhythm guitar, electric slide guitar, sound effects
Ringo Starr – drums, cowbell, vocal shout
Mal Evans – trumpet
Personnel per Mark Lewisohn[12] and Alan W. Pollack[21]
Notes[edit]
Jump up ^ McKinney, Devin (2003). Magic Circles: The Beatles in Dream and History. Harvard University Press. p. 231. ISBN 0-674-01202-X.
Jump up ^ Winn, John C (2009). That Magic Feeling: The Beatles' Recorded Legacy, Volume Two, 1966–1970. Three Rivers Press. p. 210. ISBN 0-307-45239-5.
Jump up ^ Rowley, David (2013). All Together Now. Troubador Publishing Ltd. p. 68.
Jump up ^ Athitakis, Mark (September–October 2013). "A Beatles Reflection". Humanities. National Endowment of the Humanities. Retrieved 24 April 2016.
^ Jump up to: a b c Miles 1997, pp. 487–488.
^ Jump up to: a b Sheff 2000, p. 200.
Jump up ^ Erlewine 2007.
Jump up ^ "100 Greatest Beatles Songs". 19 September 2011. Retrieved 15 October 2014.
Jump up ^ AskOxford 2008.
^ Jump up to: a b The Beatles 2000, p. 311.
Jump up ^ Beatles Interview Database 1968.
^ Jump up to: a b c d e f Lewisohn 1988, p. 154.
Jump up ^ Marck 2008.
Jump up ^ Spitz 2005, p. 794.
Jump up ^ Brown 2007.
Jump up ^ Allmusic 2007.
Jump up ^ Graff & Durchholz 1999, p. 88.
Jump up ^ Richardson, Mark (10 September 2009). "The Beatles: The Beatles". Pitchfork Media. Retrieved 24 November 2015.
Jump up ^ MacDonald 2005, p. 298.
Jump up ^ Brackett & Hoard, p. 53.
^ Jump up to: a b Pollack 1998.
^ Jump up to: a b Bugliosi 1997, pp. 240–247.
Jump up ^ Linder 2007a.
Jump up ^ Linder 2007b.
Jump up ^ "Pandora's Box - Aerosmith". AllMusic. Retrieved 7 September 2017.
Jump up ^ "The Bobs - The Bobs". AllMusic. Retrieved 7 September 2017.
Jump up ^ 1984 Grammy award nomination, Best Vocal Arrangement for Two or More Voices, Richard Greene, Gunnar Madsen - Helter Skelter (The Bobs) LA Times, "The Envelope" awards database, accessed 2010 Jan 13.
Jump up ^ "U2 - Helter Skelter". U2songs.com. Retrieved 20 November 2017.
Jump up ^ "Bono Bites Back". Mother Jones Magazine. Retrieved 20 November 2017.
Jump up ^ Final Nominations List, 53rd Grammy Awards Archived 14 December 2010 at the Wayback Machine., National Academy of Recording Arts and Sciences. Retrieved on 10 February 2011.
Jump up ^ [1][permanent dead link] Yahoo! Entertainment Story - Reuters. Retrieved on 13 February 2011.[dead link]
Jump up ^ '12-12-12': Paul McCartney fronts Nirvana 'reunion' and more highlights from Sandy benefit concert Archived 14 December 2012 at the Wayback Machine.. Retrieved 13 December 2012.
References[edit]
"Review of 'Helter Skelter'". Allmusic. 2007. Archived from the original on 28 December 2006. Retrieved 28 February 2007.
"Definition of helter-skelter". AskOxford. 2008. Retrieved 19 Sep 2010.
"Radio Luxembourg interview, Paul McCartney". Beatles Interview Database. 20 November 1968. Retrieved 6 December 2009.
The Beatles (2000). The Beatles Anthology. San Francisco: Chronicle Books. ISBN 0-8118-2684-8.
Brackett, Nathan; with Hoard, Christian (eds) (2004). The New Rolling Stone Album Guide (4th edn). New York, NY: Fireside/Simon & Schuster. ISBN 0-7432-0169-8.
Brown, Mike (2007). "Helter Skelter". What Goes On. Retrieved 12 June 2011.
Bugliosi, Vincent; Gentry, Burt (1994). Helter Skelter: The True Story of the Manson Murders (25th Anniversary ed.). W.W. Norton & Company. ISBN 0-393-08700-X.
"Dianne Heatherington - Helter Skelter". Discogs. 2009. Retrieved 15 April 2009.
Erlewine, Stephen Thomas (2007). "Review of The Beatles [White Album]". Allmusic. Retrieved 20 April 2007.
Graff, Gary; Durchholz, Daniel (eds) (1999). MusicHound Rock: The Essential Album Guide (2nd ed.). Farmington Hills, MI: Visible Ink Press. ISBN 1-57859-061-2. CS1 maint: Extra text: authors list (link)
Hoekstra, Ray (1978). "Will You Die For Me?". Archived from the original on 19 November 2010. Retrieved 17 April 2007.
Lewisohn, Mark (1988). The Beatles Recording Sessions. New York: Harmony Books. ISBN 0-517-57066-1.
Linder, Douglas (2007a). "Testimony of Paul Watkins in the Charles Manson Trial". The Trial of Charles Manson. University of Missouri-Kansas City School of Law. Archived from the original on 20 March 2007. Retrieved 28 February 2007.
Linder, Douglas (2007b). "The Influence of the Beatles on Charles Manson". The Trial of Charles Manson. University of Missouri-Kansas City School of Law. Archived from the original on 21 December 2002. Retrieved 28 February 2007.
MacDonald, Ian (2005). Revolution in the Head: The Beatles' Records and the Sixties (Second Revised ed.). London: Pimlico (Rand). ISBN 1-84413-828-3.
Marck, John T. (2008). "Helter Skelter - Music History". I Am The Beatles. Retrieved 22 June 2008.
Miles, Barry (1997). Paul McCartney: Many Years From Now. New York: Henry Holt & Company. ISBN 0-8050-5249-6.
Pollack, Alan W (7 June 1998). "Helter Skelter". Notes on ... series.
Sheff, David (2000). All We Are Saying: The Last Major Interview with John Lennon and Yoko Ono. New York: St. Martin's Press. ISBN 0-312-25464-4.
Spitz, Bob (2005). The Beatles: The Biography. Boston: Little, Brown. ISBN 0-316-80352-9.
External links[edit]
Wikiquote has quotations related to: The Beatles (album)
Alan W. Pollack's Notes on "Helter Skelter"
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"Back in the U.S.S.R."
"Dear Prudence"
"Glass Onion"
"Ob-La-Di, Ob-La-Da"
"Wild Honey Pie"
"The Continuing Story of Bungalow Bill"
"While My Guitar Gently Weeps"
"Happiness Is a Warm Gun"
Side two
"Martha My Dear"
"I'm So Tired"
"Blackbird"
"Piggies"
"Rocky Raccoon"
"Don't Pass Me By"
"Why Don't We Do It in the Road?"
"I Will"
"Julia"
Side three
"Birthday"
"Yer Blues"
"Mother Nature's Son"
"Everybody's Got Something to Hide Except Me and My Monkey"
"Sexy Sadie"
"Helter Skelter"
"Long, Long, Long"
Side four
"Revolution 1"
"Honey Pie"
"Savoy Truffle"
"Cry Baby Cry"
"Revolution 9"
"Good Night"
Non-album single
"Hey Jude
Revolution"
Outtakes
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"Child of Nature"
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"Junk"
"Not Guilty"
"Sour Milk Sea"
"What's the New Mary Jane"
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1963
"Please Please Me" / "Ask Me Why"
"From Me to You" / "Thank You Girl"
"She Loves You" / "I'll Get You"
1964
"Can't Buy Me Love" / "You Can't Do That"
"I Feel Fine" / "She's a Woman"
1965
"Ticket to Ride" / "Yes It Is"
"Help!" / "I'm Down"
"We Can Work It Out" / "Day Tripper"
1966
"Paperback Writer" / "Rain"
"Yellow Submarine" / "Eleanor Rigby"
1967
"Penny Lane" / "Strawberry Fields Forever"
"All You Need Is Love" / "Baby, You're a Rich Man"
"Hello, Goodbye" / "I Am the Walrus"
1968
"Lady Madonna" / "The Inner Light"
"Hey Jude" / "Revolution"
1969
"Get Back" / "Don't Let Me Down"
"The Ballad of John and Yoko" / "Old Brown Shoe"
"Something" / "Come Together"
1970
"Let It Be" / "You Know My Name (Look Up the Number)"
1978
"Sgt. Pepper's Lonely Hearts Club Band/With a Little Help from My Friends" / "A Day in the Life"
1982
"The Beatles Movie Medley" / "I'm Happy Just to Dance with You"
1995
"Free as a Bird" / "Christmas Time (Is Here Again)"
1996
"Real Love" / "Baby's in Black"
UK only
(Parlophone,
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1962
"My Bonnie" / "The Saints"
"Love Me Do" / "P.S. I Love You"
1963
"I Want to Hold Your Hand" / "This Boy"
1964
"Ain't She Sweet" / "If You Love Me, Baby"
"A Hard Day's Night" / "Things We Said Today"
1976
"Yesterday" / "I Should Have Known Better"
"Back in the U.S.S.R." / "Twist and Shout"
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(Vee-Jay,
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Apple)
1963
"I Want to Hold Your Hand" / "I Saw Her Standing There"
1964
"Twist and Shout" / "There's a Place"
"Do You Want to Know a Secret" / "Thank You Girl"
"Love Me Do" / "P.S. I Love You"
"Sie liebt dich" / "I'll Get You"
"I'll Cry Instead" / "I'm Happy Just to Dance with You"
"And I Love Her" / "If I Fell"
"A Hard Day's Night" / "I Should Have Known Better"
"Matchbox" / "Slow Down"
1965
"Eight Days a Week" / "I Don't Want to Spoil the Party"
"Yesterday" / "Act Naturally"
1966
"Nowhere Man" / "What Goes On"
1970
"The Long and Winding Road" / "For You Blue"
1976
"Got to Get You into My Life" / "Helter Skelter"
"Ob-La-Di, Ob-La-Da" / "Julia"
Other
countries
(Odeon,
Parlophone,
Apple)
1963
"All My Loving" / "This Boy" (Canada)
1964
"Komm, gib mir deine Hand / Sie liebt dich" (Germany)
1965
"Rock and Roll Music" / "I'm a Loser" (Europe, Australia)
1966
"Norwegian Wood (This Bird Has Flown)" / "Nowhere Man" (Australia)
"Michelle" / "Girl" (Europe)
1968
"Ob-La-Di, Ob-La-Da" / "While My Guitar Gently Weeps" (Europe, Japan, Australia)
"Ob-La-Di, Ob-La-Da" / "I Will" (Philippines)
"Back in the U.S.S.R." / "Don't Pass Me By" (Sweden)
1969
"You're Going to Lose That Girl" / "Tell Me What You See" (Japan)
1970
"Oh! Darling" / "Here Comes the Sun" (Japan)
1972
"All Together Now" / "Hey Bulldog" (Europe)
1978
"Sgt. Pepper's Lonely Hearts Club Band/With a Little Help from My Friends" / "Within You Without You" (Germany)
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Light-dependent reactions
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Light-dependent reactions of photosynthesis at the thylakoid membrane
In photosynthesis, the light-dependent reactions take place on the thylakoid membranes. The inside of the thylakoid membrane is called the lumen, and outside the thylakoid membrane is the stroma, where the light-independent reactions take place. The thylakoid membrane contains some integral membrane protein complexes that catalyze the light reactions. There are four major protein complexes in the thylakoid membrane: Photosystem II (PSII), Cytochrome b6f complex, Photosystem I (PSI), and ATP synthase. These four complexes work together to ultimately create the products ATP and NADPH.
The four photosystems absorb light energy through pigments—primarily the chlorophylls, which are responsible for the green color of leaves. The light-dependent reactions begin in photosystem II. When a chlorophyll a molecule within the reaction center of PSII absorbs a photon, an electron in this molecule attains a higher energy level. Because this state of an electron is very unstable, the electron is transferred from one to another molecule creating a chain of redox reactions, called an electron transport chain (ETC). The electron flow goes from PSII to cytochrome b6f to PSI. In PSI, the electron gets the energy from another photon. The final electron acceptor is NADP. In oxygenic photosynthesis, the first electron donor is water, creating oxygen as a waste product. In anoxygenic photosynthesis various electron donors are used.
Cytochrome b6f and ATP synthase work together to create ATP. This process is called photophosphorylation, which occurs in two different ways. In non-cyclic photophosphorylation, cytochrome b6f uses the energy of electrons from PSII to pump protons from the stroma to the lumen. The proton gradient across the thylakoid membrane creates a proton-motive force, used by ATP synthase to form ATP. In cyclic photophosphorylation, cytochrome b6f uses the energy of electrons from not only PSII but also PSI to create more ATP and to stop the production of NADPH. Cyclic phosphorylation is important to create ATP and maintain NADPH in the right proportion for the light-independent reactions.
The net-reaction of all light-dependent reactions in oxygenic photosynthesis is:
2H
2O + 2NADP+
+ 3ADP + 3Pi → O
2 + 2NADPH + 3ATP
The two photosystems are protein complexes that absorb photons and are able to use this energy to create an electron transport chain. Photosystem I and II are very similar in structure and function. They use special proteins, called light-harvesting complexes, to absorb the photons with very high effectiveness. If a special pigment molecule in a photosynthetic reaction center absorbs a photon, an electron in this pigment attains the excited state and then is transferred to another molecule in the reaction center. This reaction, called photoinduced charge separation, is the start of the electron flow and is unique because it transforms light energy into chemical forms.
Contents
1 The reaction center
2 Photosynthetic electron transport chains in chloroplasts
2.1 Photosystem II
2.1.1 The water-splitting complex
2.1.2 The reaction center
2.1.3 Link of water-splitting complex and chlorophyll excitation
2.1.4 Summary
2.2 Cytochrome b6
2.3 Photosystem I
3 Photosynthetic electron transport chains in bacteria
3.1 Cyanobacteria
3.2 Purple bacteria
3.3 Green sulfur bacteria
4 History
5 See also
6 References
The reaction center[edit]
Main article: Photosynthetic reaction center
The reaction center is in the thylakoid membrane. It transfers light energy to a dimer of chlorophyll pigment molecules near the periplasmic (or thylakoid lumen) side of the membrane. This dimer is called a special pair because of its fundamental role in photosynthesis. This special pair is slightly different in PSI and PSII reaction center. In PSII, it absorbs photons with a wavelength of 680 nm, and it is therefore called P680. In PSI, it absorbs photons at 700 nm, and it is called P700. In bacteria, the special pair is called P760, P840, P870, or P960. Where "P" means pigment, and the number following it is the wavelength of light absorbed.
If an electron of the special pair in the reaction center becomes excited, it cannot transfer this energy to another pigment using resonance energy transfer. In normal circumstances, the electron should return to the ground state, but, because the reaction center is arranged so that a suitable electron acceptor is nearby, the excited electron can move from the initial molecule to the acceptor. This process results in the formation of a positive charge on the special pair (due to the loss of an electron) and a negative charge on the acceptor and is, hence, referred to as photoinduced charge separation. In other words, electrons in pigment molecules can exist at specific energy levels. Under normal circumstances, they exist at the lowest possible energy level they can. However, if there is enough energy to move them into the next energy level, they can absorb that energy and occupy that higher energy level. The light they absorb contains the necessary amount of energy needed to push them into the next level. Any light that does not have enough or has too much energy cannot be absorbed and is reflected. The electron in the higher energy level, however, does not want to be there; the electron is unstable and must return to its normal lower energy level. To do this, it must release the energy that has put it into the higher energy state to begin with. This can happen various ways. The extra energy can be converted into molecular motion and lost as heat. Some of the extra energy can be lost as heat energy, while the rest is lost as light. (This re-emission of light energy is called fluorescence.) The energy, but not the e- itself, can be passed onto another molecule. (This is called resonance.) The energy and the e- can be transferred to another molecule. Plant pigments usually utilize the last two of these reactions to convert the sun's energy into their own.
This initial charge separation occurs in less than 10 picoseconds (10−11 seconds). In their high-energy states, the special pigment and the acceptor could undergo charge recombination; that is, the electron on the acceptor could move back to neutralize the positive charge on the special pair. Its return to the special pair would waste a valuable high-energy electron and simply convert the absorbed light energy into heat. In the case of PSII, this backflow of electrons can produce reactive oxygen species leading to photoinhibition.[1][2] Three factors in the structure of the reaction center work together to suppress charge recombination nearly completely.
Another electron acceptor is less than 10 Å away from the first acceptor, and so the electron is rapidly transferred farther away from the special pair.
An electron donor is less than 10 Å away from the special pair, and so the positive charge is neutralized by the transfer of another electron
The electron transfer back from the electron acceptor to the positively charged special pair is especially slow. The rate of an electron transfer reaction increases with its thermodynamic favorability up to a point and then decreases. The back transfer is so favourable that it takes place in the inverted region where electron-transfer rates become slower.[1]
Thus, electron transfer proceeds efficiently from the first electron acceptor to the next, creating an electron transport chain that ends if it has reached NADPH.
Photosynthetic electron transport chains in chloroplasts[edit]
The photosynthesis process in chloroplasts begins when an electron of P680 of PSII attains a higher-energy level. This energy is used to reduce a chain of electron acceptors that have subsequently lowered redox-potentials. This chain of electron acceptors is known as an electron transport chain. When this chain reaches PS I, an electron is again excited, creating a high redox-potential. The electron transport chain of photosynthesis is often put in a diagram called the z-scheme, because the redox diagram from P680 to P700 resembles the letter z.[3]
The final product of PSII is plastoquinol, a mobile electron carrier in the membrane. Plastoquinol transfers the electron from PSII to the proton pump, cytochrome b6f. The ultimate electron donor of PSII is water. Cytochrome b6f proceeds the electron chain to PSI through plastocyanin molecules. PSI is able to continue the electron transfer in two different ways. It can transfer the electrons either to plastoquinol again, creating a cyclic electron flow, or to an enzyme called FNR (Ferredoxin—NADP(+) reductase), creating a non-cyclic electron flow. PSI releases FNR into the stroma, where it reduces NADP+
to NADPH.
Activities of the electron transport chain, especially from cytochrome b6f, lead to pumping of protons from the stroma to the lumen. The resulting transmembrane proton gradient is used to make ATP via ATP synthase.
The overall process of the photosynthetic electron transport chain in chloroplasts is:
H
2O → PS II → plastoquinone → cytb
6 → plastocyanin → PS I → NADPH
Photosystem II[edit]
Main article: Photosystem II
PS II is extremely complex, a highly organized transmembrane structure that contains a water-splitting complex, chlorophylls and carotenoid pigments, a reaction center (P680), pheophytin (a pigment similar to chlorophyll), and two quinones. It uses the energy of sunlight to transfer electrons from water to a mobile electron carrier in the membrane called plastoquinone:
H
2O → P680 → P680* → plastoquinone
Plastoquinone, in turn, transfers electrons to cytb
6, which feeds them into PS I.
The water-splitting complex[edit]
The step H
2O → P680 is performed by a poorly understood structure embedded within PS II called the water-splitting complex or the oxygen-evolving complex. It catalyzes a reaction that splits water into electrons, protons and oxygen:
2H
2O → 4H+ + 4e− + O
2
The actual steps of the above reaction are running in the following way (Dolai's diagram of S-states): (I) 2H2O(monoxide) (II) OH. H2O (hydroxide) (III) H2O2(peroxide) (IV)HO2(super oxide)(V) O2(di-oxygen).[4] (Dolai's mechanism)
The electrons are transferred to special chlorophyll molecules (embedded in PS II) that are promoted to a higher-energy state by the energy of photons.
The reaction center[edit]
The excitation P680 → P680*of the reaction center pigment P680 occurs here. These special chlorophyll molecules embedded in PS II absorb the energy of photons, with maximal absorption at 680 nm. Electrons within these molecules are promoted to a higher-energy state. This is one of two core processes in photosynthesis, and it occurs with astonishing efficiency (greater than 90%) because, in addition to direct excitation by light at 680 nm, the energy of light first harvested by antenna proteins at other wavelengths in the light-harvesting system is also transferred to these special chlorophyll molecules.
This is followed by the step P680*→ pheophytin, and then on to plastoquinone, which occurs within the reaction center of PS II. High-energy electrons are transferred to plastoquinone before it subsequently picks up two protons to become plastoquinol. Plastoquinol is then released into the membrane as a mobile electron carrier.
This is the second core process in photosynthesis. The initial stages occur within picoseconds, with an efficiency of 100%. The seemingly impossible efficiency is due to the precise positioning of molecules within the reaction center. This is a solid-state process, not a chemical reaction. It occurs within an essentially crystalline environment created by the macromolecular structure of PS II. The usual rules of chemistry (which involve random collisions and random energy distributions) do not apply in solid-state environments.
Link of water-splitting complex and chlorophyll excitation[edit]
When the chlorophyll passes the electron to pheophytin, it obtains an electron from P680*. In turn, P680* can oxidize the Z (or YZ) molecule. Once oxidized, the Z molecule can derive electrons from the oxygen-evolving complex (OEC).[5] Dolai's S-state diagrams show the reactions of water splitting in the oxygen-evolving complex.
Summary[edit]
PS II is a transmembrane structure found in all chloroplasts. It splits water into electrons, protons and molecular oxygen. The electrons are transferred to plastoquinone, which carries them to a proton pump. Molecular oxygen is released into the atmosphere.
The emergence of such an incredibly complex structure, a macromolecule that converts the energy of sunlight into potentially useful work with efficiencies that are impossible in ordinary experience, seems almost magical at first glance. Thus, it is of considerable interest that, in essence, the same structure is found in purple bacteria.
Cytochrome b
6[edit]
PS II and PS I are connected by a transmembrane proton pump, cytochrome b
6 complex (plastoquinol—plastocyanin reductase; EC 1.10.99.1). Electrons from PS II are carried by plastoquinol to cytb
6, where they are removed in a stepwise fashion (reforming plastoquinone) and transferred to a water-soluble electron carrier called plastocyanin. This redox process is coupled to the pumping of four protons across the membrane. The resulting proton gradient (together with the proton gradient produced by the water-splitting complex in PS II) is used to make ATP via ATP synthase.
The similarity in structure and function between cytochrome b
6 (in chloroplasts) and cytochrome bc
1 (Complex III in mitochondria) is striking. Both are transmembrane structures that remove electrons from a mobile, lipid-soluble electron carrier (plastoquinone in chloroplasts; ubiquinone in mitochondria) and transfer them to a mobile, water-soluble electron carrier (plastocyanin in chloroplasts; cytochrome c in mitochondria). Both are proton pumps that produce a transmembrane proton gradient.
Photosystem I[edit]
The cyclic light-dependent reactions occur only when the sole photosystem being used is photosystem I. Photosystem I excites electrons which then cycle from the transport protein, ferredoxin (Fd), to the cytochrome complex, b6f, to another transport protein, plastocyanin (Pc), and back to photosystem I. A proton gradient is created across the thylakoid membrane (6) as protons (3) are transported from the chloroplast stroma (4) to the thylakoid lumen (5). Through chemiosmosis, ATP (9) is produced where ATP synthase (1) binds an inorganic phosphate group (8) to an ADP molecule (7).
Main article: Photosystem I
PS I accepts electrons from plastocyanin and transfers them either to NADPH (noncyclic electron transport) or back to cytochrome b
6 (cyclic electron transport):
                        plastocyanin → P700 → P700* → FNR → NADPH ↑ ↓ b
6 ← plastoquinone

PS I, like PS II, is a complex, highly organized transmembrane structure that contains antenna chlorophylls, a reaction center (P700), phylloquinine, and a number of iron-sulfur proteins that serve as intermediate redox carriers.
The light-harvesting system of PS I uses multiple copies of the same transmembrane proteins used by PS II. The energy of absorbed light (in the form of delocalized, high-energy electrons) is funneled into the reaction center, where it excites special chlorophyll molecules (P700, maximum light absorption at 700 nm) to a higher energy level. The process occurs with astonishingly high efficiency.
Electrons are removed from excited chlorophyll molecules and transferred through a series of intermediate carriers to ferredoxin, a water-soluble electron carrier. As in PS II, this is a solid-state process that operates with 100% efficiency.
There are two different pathways of electron transport in PS I. In noncyclic electron transport, ferredoxin carries the electron to the enzyme ferredoxin NADP+
reductase (FNR) that reduces NADP+
to NADPH. In cyclic electron transport, electrons from ferredoxin are transferred (via plastoquinone) to a proton pump, cytochrome b
6. They are then returned (via plastocyanin) to P700.
NADPH and ATP are used to synthesize organic molecules from CO
2. The ratio of NADPH to ATP production can be adjusted by adjusting the balance between cyclic and noncyclic electron transport.
It is noteworthy that PS I closely resembles photosynthetic structures found in green sulfur bacteria, just as PS II resembles structures found in purple bacteria.
Photosynthetic electron transport chains in bacteria[edit]
PS II, PS I, and cytochromeb
6 are found in chloroplasts. All plants and all photosynthetic algae contain chloroplasts, which produce NADPH and ATP by the mechanisms described above. In essence, the same transmembrane structures are also found in cyanobacteria.
Unlike plants and algae, cyanobacteria are prokaryotes. They do not contain chloroplasts. Rather, they bear a striking resemblance to chloroplasts themselves. This suggests that organisms resembling cyanobacteria were the evolutionary precursors of chloroplasts. One imagines primitive eukaryotic cells taking up cyanobacteria as intracellular symbionts in a process known as endosymbiosis.
Cyanobacteria[edit]
Cyanobacteria contain structures similar to PS II and PS I in chloroplasts. Their light-harvesting system is different from that found in plants (they use phycobilins, rather than chlorophylls, as antenna pigments), but their electron transport chain
         H
2O → PS II → plastoquinone → b
6 → cytochrome c
6 → PS I → ferredoxin → NADPH ↑ ↓ b
6 ← plastoquinone

is, in essence, the same as the electron transport chain in chloroplasts. The mobile water-soluble electron carrier is cytochrome c
6 in cyanobacteria, plastocyanin in plants.
Cyanobacteria can also synthesize ATP by oxidative phosphorylation, in the manner of other bacteria. The electron transport chain is
           NADH dehydrogenase → plastoquinone → b
6 → cytochrome c
6 → cytochrome aa
3 → O
2

where the mobile electron carriers are plastoquinone and cytochrome c
6, while the proton pumps are NADH dehydrogenase, b
6 and cytochrome aa
3.
Cyanobacteria are the only bacteria that produce oxygen during photosynthesis. Earth's primordial atmosphere was anoxic. Organisms like cyanobacteria produced our present-day oxygen-containing atmosphere.
The other two major groups of photosynthetic bacteria, purple bacteria and green sulfur bacteria, contain only a single photosystem and do not produce oxygen.
Purple bacteria[edit]
Purple bacteria contain a single photosystem that is structurally related to PS II in cyanobacteria and chloroplasts:
P870 → P870* → ubiquinone → bc
1 → cytochrome c
2 → P870
This is a cyclic process in which electrons are removed from an excited chlorophyll molecule (bacteriochlorophyll; P870), passed through an electron transport chain to a proton pump (cytochrome bc
1 complex, similar but not identical to cytochrome bc
1 in chloroplasts), and then returned to the chlorophyll molecule. The result is a proton gradient, which is used to make ATP via ATP synthase. As in cyanobacteria and chloroplasts, this is a solid-state process that depends on the precise orientation of various functional groups within a complex transmembrane macromolecular structure.
To make NADPH, purple bacteria use an external electron donor (hydrogen, hydrogen sulfide, sulfur, sulfite, or organic molecules such as succinate and lactate) to feed electrons into a reverse electron transport chain.
Green sulfur bacteria[edit]
Green sulfur bacteria contain a photosystem that is analogous to PS I in chloroplasts:
                           P840 → P840* → ferredoxin → NADH ↑ ↓ cyt c553 ← bc
1 ← menaquinone

There are two pathways of electron transfer. In cyclic electron transfer, electrons are removed from an excited chlorophyll molecule, passed through an electron transport chain to a proton pump, and then returned to the chlorophyll. The mobile electron carriers are, as usual, a lipid-soluble quinone and a water-soluble cytochrome. The resulting proton gradient is used to make ATP.
In noncyclic electron transfer, electrons are removed from an excited chlorophyll molecule and used to reduce NAD+ to NADH. The electrons removed from P840 must be replaced. This is accomplished by removing electrons from H
2S, which is oxidized to sulfur (hence the name "green sulfur bacteria").
Purple bacteria and green sulfur bacteria occupy relatively minor ecological niches in the present day biosphere. They are of interest because of their importance in precambrian ecologies, and because their methods of photosynthesis were the likely evolutionary precursors of those in modern plants.
History[edit]
The first ideas about light being used in photosynthesis were proposed by Colin Flannery in 1779[6] who recognized it was sunlight falling on plants that was required, although Joseph Priestley had noted the production of oxygen without the association with light in 1772.[7] Cornelis Van Niel proposed in 1931 that photosynthesis is a case of general mechanism where a photon of light is used to photo decompose a hydrogen donor and the hydrogen being used to reduce CO
2.[8] Then in 1939, Robin Hill showed that isolated chloroplasts would make oxygen, but not fix CO
2 showing the light and dark reactions occurred in different places. Although they are referred to as light and dark reactions, both of them take place only in the presence of light.[9] This led later to the discovery of photosystems I and II.
See also[edit]
Light-independent reaction
Photosynthetic reaction centre
Photosystem II
Compensation point
References[edit]
^ Jump up to: a b Vass, Imre; Cser, Krisztian (April 2009). "Janus-faced charge recombinations in photosystem II photoinhibition". Trends in Plant Science. 14 (4): 200–205. doi:10.1016/j.tplants.2009.01.009. ISSN 1360-1385. PMID 19303349. Retrieved 2012-10-30.
Jump up ^ Keren, Nir; Berg, Alexander; van Kan, Paul J. M.; Levanon, Haim; Ohad, Itzhak (1997-02-18). "Mechanism of photosystem II photoinactivation and D1 protein degradation at low light: The role of back electron flow". Proceedings of the National Academy of Sciences. 94 (4): 1579–1584. doi:10.1073/pnas.94.4.1579. ISSN 0027-8424. PMC 19834. PMID 11038602. Retrieved 2012-10-30.
Jump up ^ Rajni Govindjee. "The Z-Scheme Diagram of Photosynthesis". Retrieved March 2, 2006.
Jump up ^ Dolai, U. (2017). "Chemical Scheme of Water -Splitting Process during Photosynthesis by the way of Experimental Analysis ". IOSR Journal of Pharmacy and Biological Sciences. 12(6):65-67. doi: 10.9790/3008-1206026567. ISSN 2319-7676.
Jump up ^ "Photosynthesis". McGraw Hill Encyclopedia of Science and Technology. 2007. p. 472.
Jump up ^ Ingenhousz, J (1779). Experiments Upon Vegetables. London: Elmsly and Payne.
Jump up ^ Priestley, J (1772). "Observations on Different Kinds of Air". 62. London: Phil. Trans. Roy. Soc.: 147–264. doi:10.1098/rstl.1772.0021.
Jump up ^ van Niel, C. B. (1931). "On the morphology and physiology of the purple and green sulfur bacteria". Arch. Microbiol. 3: 1–114. doi:10.1007/BF00454965.
Jump up ^ Hill, R. (May 1939). "Oxygen Produced by Isolated Chloroplasts". Proceedings of the Royal Society of London. Series B, Biological Sciences. 127 (847): 192–210. doi:10.1098/rspb.1939.0017.
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