text:
Lucky Charms
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For other uses, see Lucky Charms (disambiguation).
Lucky Charms
The cereal's logo and leprechaun mascot, Lucky.
Product type
Cereal with marshmallows
Owner
General Mills
Country
United States
Introduced
March 20, 1964; 54 years ago (1964-03-20)
Markets
United States, United Kingdom, Canada
Tagline
"They're magically delicious."
Website
luckycharms.com
Lucky Charms is a brand of cereal produced by the General Mills food company since 1964.[1] The cereal consists of toasted oat pieces and multi-colored marshmallow shapes ("marbits" or marshmallow bits). The label features a leprechaun mascot, Lucky, animated in commercials.
Contents
[hide]
1 History
2 Marshmallows
2.1 Limited Edition Marshmallows
2.2 Marshmallow-only promotion
3 Theme song
4 Taglines
5 See also
6 References
7 Further reading
8 External links
History[edit]
Lucky Charms was created in 1964 by product developer John Holahan. General Mills management challenged a team of product developers to use the available manufacturing capacity from either of General Mills' two principal cereal products—Wheaties or Cheerios—and do something unique. Holahan came up with the idea after a visit to the grocery store in which he decided to mix Cheerios with bits of Brach's Circus Peanuts.[2]
An advertising company employed by General Mills and Company suggested marketing the new cereal around the idea of charm bracelets.[3] Thus, the charms of Lucky Charms were born. Lucky Charms is the first cereal to include marshmallows in the recipe. The mascot of Lucky Charms, created in 1963, is Lucky the Leprechaun, also known as Sir Charms, and originally called L.C. Leprechaun.[4] The cartoon character's voice was supplied by the late voice actor Arthur Anderson until 1992. In 1975, Lucky the Leprechaun was briefly replaced by Waldo the Wizard in New England, while Lucky remained the mascot in the rest of the United States.[5] The oat cereal was not originally sugar coated. After initial sales failed to meet expectations, the oats were sugar coated, and the cereal's success grew. Piggy banks and plastic watches were introduced as cereal box send-away prizes as a marketing tactic to increase sales. The recipe for the cereal remained unchanged until the introduction of a new flavor: Chocolate Lucky Charms, in 2005. Later in 2012, General Mills introduced "Lucky Charms Marshmallow Treats."
Following the product launch, the General Mills marketing department found that sales performed dramatically better if the composition of the marbits changed periodically.[3] Various features of the marbits were modified to maximize their appeal to young consumers. Over the years, over 40 limited edition features such as Winter Lucky Charms, Olympic-themed Lucky Charms, and Lucky Charms featuring marshmallow landmarks from around the world, were created to drive consumer demands. In focus groups and market research, more brightly colored charms resulted in better sales than did dull or pastel colors.[3] Currently, General Mills conducts "concept-ideation" studies on Lucky Charms.[3]
Marshmallows[edit]
Lucky Charms
The first boxes of Lucky Charms cereal contained marshmallows in the shapes of pink hearts, yellow moons, orange stars, and green clovers. The lineup has changed occasionally, beginning with the introduction of blue diamonds in 1975, followed by purple horseshoes in 1983,[6] red balloons in 1989, green trees 1991, rainbows in 1992, blue moons 1995, leprechaun hats in 1997 (temporarily replaced the green clovers), orange shooting stars and around the world charms in 1998 (added blue, green, yellow, purple, and red in 2011), a crystal ball in 2001, and an hourglass in 2008.[3] In 2013, 6 new rainbow swirl moons and 2 new rainbow charms were introduced. From the original four marshmallows, the permanent roster as of 2013 includes eight marshmallows.
Older marshmallows were phased out periodically. The first shapes to disappear were the yellow moons and blue diamonds, replaced by yellow/orange pots of gold and blue moons respectively in 1994. In 2006, the assortment included purple horseshoes; red balloons; blue crescent-moons; orange and white shooting stars; yellow and orange pots of gold; pink, yellow, and blue rainbows; two-tone green leprechaun hats; pink hearts (the one shape to survive since the beginning); with the most recent addition being the return of the clovers in 2004. The hourglass shape was retired in spring 2018 and was replaced by a unicorn, which was chosen on social media by way of emojis[7][8]. The size and brightness of the marshmallows changed in 2004.[9]
Recent changes to the marshmallows include the star shape taking on a "shooting star" design, the orange five-pointed star being complemented by a white "trail." In late 2005, another marshmallow shape was added, the "Hidden Key". It is a solid yellow marshmallow that resembles an arched door (similar to the shape of a tombstone; flat at the bottom, flat sides with a round top). When liquid is added to the cereal, the sugar in the marshmallow dissolves and the shape of a skeleton key appears "as if by magic." The tagline was, "Unlock the door with milk!" This "new" marshmallow type has been used in other hot and cold cereals, but with mixed success (from characters "hidden" inside a bigger marshmallow to letters appearing). In early June 2006, General Mills introduced Magic Mirror marshmallows. In 2008, yellow and orange hourglass marshmallows were introduced with the marketing tagline of, "The Hourglass Charm has the power to Stop Time * Speed Up Time * Reverse Time". As of 2011, swirled marshmallows and rainbow-colored stars have been introduced.[citation needed] In 2018, for the first time in ten years Lucky Charms retired a marshmallow, which was the hourglass, and added a new permanent marshmallow, the Magical Unicorn. It's powers are unknown.
The marshmallows are meant to represent Lucky's magical charms, each with their own special meaning or "power." The following are explanations of the permanent marshmallows:[10]
Hearts - power to bring things to life
Shooting Stars - power to fly
Horseshoes - power to speed things up
Green Toasters - luck, but you will never know what kind of luck you will get
Blue Moons - power of invisibility
Rainbows - instantaneous travel from place to place
Balloons - power to make things float
Hourglass - power to control time
Limited Edition Marshmallows[edit]
There have been more than 30 featured limited edition marshmallow shapes over the years, with the introduction of themed Lucky Charms, such as Winter Lucky Charms. Some of these include:
In 1986, a whale-shaped marshmallow was temporarily added to the lineup.[11]
In 1990, a green pine tree-shaped marshmallow was temporarily added to the lineup.[11] During that time, the cereal promoted Earth Day with a free Colorado Blue Spruce seedling with proofs-of-purchase.[12]
In 1991, the star and balloon shape marshmallows were combined for a short time. The red balloon featured a gold six-pointed star. The star was removed at a later date to make the Red Balloon and Star marshmallows separate.[11]
In 1994, sprinkles were temporarily added to the marshmallows.
In 1999, the moon shape marshmallows were modified with the addition of the yellow curve line for a limited time.
In 2000, a "New Sparkling Rainbow" was added to the mix for a limited time. It was described by General Mills as "a sprinkling of multicolored sugar on a white rainbow marbit." This marshmallow replaced the original rainbow at this time.[13]
In 2010, the swirled marshmallows were in Lucky Charms for a limited time.
In June 2013, two new rainbow marshmallows were added for LGBT Pride Month.[14]
In 2015, new diamond shaped marshmallows were added in.[citation needed]
Introduced in 2017, limited edition cinnamon vanilla Lucky Charms include only snowman, snowball, and snowflake shaped marshmallows.
In 2018 a unicorn shaped marshmallow was added and is here to stay unofficially.
Marshmallow-only promotion[edit]
In May 2017, General Mills announced they would be promoting 10,000 boxes of cereal that contain only marshmallow pieces.[15] In order to win one of the coveted boxes, consumers would need to purchase a specially-marked box of regular Lucky Charms with a code on the inside panel. The code would be entered into an official website to see if the consumer is the winner of one of the 10,000 novelty boxes produced. The sweepstakes ran through December 2017.[16][17]
Theme song[edit]
In the earliest commercials, Lucky Charms cereal had no theme jingle; action was accompanied by a light instrumental "Irish" tune. Soon, however, a simple two-line tag was added:
Frosted Lucky Charms,
They're magically delicious!
This simple closer, with the kids usually singing the first line and Lucky singing the second, survived into the 1980s.
Then, with the addition of the purple horseshoe marbit, it was extended into a jingle describing the contents of the box.[18] This was later revised with the addition of red balloons to the now-familiar "Hearts, stars and horse-shoes, clovers and blue-moons. Pots of gold and rainbows, and tasty red balloons!" In 2008, the pot of gold was replaced with the hourglass in the theme song.
The jingle is usually accompanied by mentioning that Lucky Charms contains whole grain ingredients, and is part of a balanced meal. General Mills' market position is centered on cereals that contain "more whole grain than any other single ingredient, which is significant, because 95 percent of Americans aren't eating minimally 48 grams of whole grain per day as recommended by the U.S. Dietary Guidelines."[19]
Taglines[edit]
This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (October 2013) (Learn how and when to remove this template message)
They're Magically Delicious![20]
Frosted Lucky Charms, They're Magically Delicious!
They're Always After Me Lucky Charms!
You'll Never Get Me Lucky Charms!
Pink Hearts, Yellow Moons, Orange Stars, and Green Clovers!
Pink Hearts, Yellow Moons, Orange Stars, Green Clovers, and Blue Diamonds!
Pink Hearts, Yellow Moons, Orange Stars, Green Clovers, Blue Diamonds, and Purple Horseshoes!
Pink Hearts, Yellow Moons, Orange Stars, Green Clovers, Blue Diamonds, Purple Horseshoes, and Red Balloons!
Hearts, Stars, and Horseshoes, Clovers and Blue Moons, Pots of Gold and Rainbows, and Red Balloons!
Hearts, Stars, and Horseshoes, Clovers and Blue Moons, Hourglasses, Rainbows, and Tasty Red Balloons!
Hearts, Stars, and Horseshoes, Clovers, Red Balloons, Hourglasses, Rainbows, and Six New Swirled Moons!
Hearts, Stars and Horseshoes, Clovers, Blue Moons, Hourglasses, Rainbows, and Tasty Red Balloons in Lucky Charms!
Heart, Star, Horseshoe, Clover, Blue Moon, Hourglass, Rainbow, and also Red Balloon!
See also[edit]
Marshmallow Mateys
References[edit]
Jump up ^ "1960s" (PDF). General Mills History Timeline. General Mills. p. 3. Archived from the original (PDF) on October 23, 2007. Retrieved 2009-05-20.
Jump up ^ Hartel, Richard; Hartel, AnnaKate (October 4, 2004). "The Miracle of Orange Circus Peanuts". The Capital Times. Madison, Wisconsin. Archived from the original on September 29, 2008.
^ Jump up to: a b c d e Karen Wright (August 1999). "A Charm's Life - Lucky Charm's cereal (sic)". Discover Magazine. Archived from the original on 2008-09-15.
Jump up ^ Hoffman, David (2005). The Breakfast Cereal Gourmet. Andrew McMeel Publishing. p. 127. ISBN 9780740750298. Retrieved 2014-05-05.
Jump up ^ "Waldo the Wizard" the Forgotten Lucky Charms Mascot". http://www.neatorama.com/2015/03/18/Waldo-the-Wizard-The-Forgotten-Lucky-Charms-Mascot/ accessed September 1, 2015
Jump up ^ http://www.blog.generalmills.com/2014/03/lucky-charms/
Jump up ^ "Lucky Charms reveals new unicorn marshmallow". KABC-TV. February 19, 2018. Retrieved February 19, 2018.
Jump up ^ "Lucky Charms Unleashes The Power Of The Unicorn As The Next New Permanent Marshmallow In Its Magical Cereal Line-Up" (Press release). PR Newswire. February 19, 2018. Retrieved February 19, 2018.
Jump up ^ "General Mills Cereals". Topher's Breakfast Cereal Character Guide. Archived from the original on March 6, 2012.
Jump up ^ "Lucky Announces the Power to Control Time with New Marshmallow Charm in Lucky Charms® Cereal!" (Press release). General Mills. June 9, 2008. Archived from the original on November 23, 2010.
^ Jump up to: a b c "5 Things about Lucky Charm Cereal". Archived from the original on 2013-05-31. Retrieved 2013-02-12.
Jump up ^ "Lucky Charms and Christmas". Retrieved 2014-06-18.
Jump up ^ Elliott, Stuart (2000-06-02). "Marketers bet on the concept of good luck as a selling tool". The New York Times. Retrieved 2009-05-06.
Jump up ^ "Lucky Charms, General Mills Cereal, Celebrates LGBT Pride With #LuckyToBe Campaign". Huffingtonpost.com. 2013-06-28. Retrieved 2013-12-30.
Jump up ^ Natt Garun (26 April 2017). "Lucky Charms Gives Away 10,000 Marshmallow-Only Boxes". The VERGE. Retrieved 17 May 2017.
Jump up ^ Painter, Kristen (2017-05-15). "General Mills to make 10,000 boxes of marshmallow-only Lucky Charms". Star Tribune. Retrieved 2017-05-16.
Jump up ^ Ben Popken (16 May 2017). "Amazon's Echo Look Privacy Could Be a Big Issue Someday". NBC News. Retrieved 17 May 2017.
Jump up ^ "TV Acres ad slogans". Archived from the original on February 4, 2013. Retrieved June 21, 2007.
Jump up ^ "Lucky Charms has adult allure". Retrieved 2013-02-12.
Jump up ^ "Lucky Charms Cereal (10/02/09)". Retrieved 2013-02-12.
Further reading[edit]
An Actor's Odyssey: Orson Welles to Lucky the Leprechaun, by Arthur Anderson. Albany, 2010. BearManor Media. ISBN 1-59393-522-6
External links[edit]
General Mills corporate Lucky Charms site: includes package information and Nutrition Facts.
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Companies listed on the New York Stock Exchange (I)
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New York Stock Exchange listed stocks:
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I[edit]
This list is complete and up-to-date as of August 2, 2014.
Stock name
Symbol
Country of origin
Iamgold Corporation IAG Canada
ICICI Bank Ltd. IBN India
Idacorp Inc. IDA US
IDEX Corporation IEX US
IDT Corporation IDT US
IFM Investments Limited CTC China
IHS Inc. IHS US
Illinois Tool Works Inc. ITW US
Imation Corporation IMN US
IMAX Corporation IMAX Canada
Imperial Holdings, Inc. IFT US
Imperva, Inc. IMPV US
IMS Health Holdings, Inc. IMS US
Independence Holding Company IHC US
India Fund, Inc. IFN US
Industrias Bachoco, S.A.B. de C.V. IBA Mexico
Infoblox Inc. BLOX US
Infosys Limited INFY US
ING Groep N.V. ING Netherlands
ING Groep N.V. IDG Netherlands
ING Groep N.V. IND Netherlands
ING Groep N.V. INZ Netherlands
ING Groep N.V. ISF Netherlands
ING Groep N.V. ISG Netherlands
ING Groep N.V. ISP Netherlands
Ingersoll-Rand plc IR US
Ingram Micro Inc. IM US
Ingredion Incorporated INGR US
Inland Real Estate Corporation IRC US
Inland Real Estate Corporation IRC.PRA US
Inphi Corporation IPHI US
Insperity, Inc. NSP US
Installed Building Products, Inc. IBP US
Integrys Energy Group, Inc. IEH US
Integrys Energy Group, Inc. TEG US
Intelsat S.A. I US
Intelsat S.A. I.PRA US
Intercontinental Exchange, Inc. ICE US
InterContinental Hotels Group PLC IHG United Kingdom
International Business Machines Corporation IBM US
International Flavors & Fragrances Inc. IFF US
International Game Technology IGT US
International Paper Co. IP US
International Rectifier Corporation IRF US
International Shipholding Corporation ISH US
International Shipholding Corporation ISH.PRA US
International Shipholding Corporation ISH.PRB US
InterOil Corporation IOC Papua New Guinea
The Interpublic Group of Companies Inc. IPG US
Interstate Power and Light Company IPL.PRD US
InterXion Holding N.V. INXN US
IntraLinks Holdings, Inc. IL US
Intrawest Resorts Holdings, Inc. SNOW US
Intrepid Potash, Inc. IPI US
Intrexon Corporation XON US
Invacare Corporation IVC US
InvenSense, Inc. INVN US
Invesco Ltd. IVZ Bermuda
Invesco Bond Fund VBF US
Invesco California Value Municipal Income Trust VCV US
Invesco Credit Dynamic Opportunities Fund VTA US
Invesco High Income Trust II VLT US
Invesco Mortgage Capital Inc. IVR US
Invesco Mortgage Capital Inc. IVR.PRA US
Invesco Municipal Income Opportunities Trust OIA US
Invesco Municipal Opportunity Trust VMO US
Invesco Municipal Trust VKQ US
Invesco Pennsylvania Value Municipal Income Trust VPV US
Invesco Quality Municipal Income Trust IQI US
Invesco Senior Income Trust VVR US
Invesco Trust for Investment Grade Municipals VGM US
Invesco Trust For Investment Grade New York Municipals VTN US
Invesco Value Municipal Income Trust IIM US
Investment Technology Group Inc. ITG US
Investors Real Estate Trust IRET US
Investors Real Estate Trust IRET.PR US
Investors Real Estate Trust IRET.PRB US
Ion Geophysical Corporation IO US
Iron Mountain Inc. IRM US
IRSA Inversiones Y Representaciones S.A. IRS Argentina
iSoftStone Holdings Limited ISS US
iStar Financial Inc. STAR US
iStar Financial Inc. STAR.PRD US
iStar Financial Inc. STAR.PRE US
iStar Financial Inc. STAR.PRF US
iStar Financial Inc. STAR.PRG US
iStar Financial Inc. STAR.PRI US
Itau Unibanco Holding S.A. ITUB Brazil
ITC Holdings Corp. ITC US
ITT Corporation ITT US
ITT Educational Services Inc. ESI US
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Chemiosmosis
From Wikipedia, the free encyclopedia
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Chemiosmosis is the movement of ions across a semipermeable membrane, down their electrochemical gradient. An example of this would be the generation of adenosine triphosphate (ATP) by the movement of hydrogen ions across a membrane during cellular respiration or photosynthesis.
An ion gradient has potential energy and can be used to power chemical reactions when the ions pass through a channel (red).
Hydrogen ions, or protons, will diffuse from an area of high proton concentration to an area of lower proton concentration, and an electrochemical concentration gradient of protons across a membrane can be harnessed to make ATP. This process is related to osmosis, the diffusion of water across a membrane, which is why it is called "chemiosmosis".
ATP synthase is the enzyme that makes ATP by chemiosmosis. It allows protons to pass through the membrane and uses the free energy difference to phosphorylate adenosine diphosphate (ADP), making ATP. The generation of ATP by chemiosmosis occurs in mitochondria and chloroplasts, as well as in most bacteria and archaea, an electron transport chain pumps H+ ions in the thylakoid spaces through thylakoid membranes. The energy from the electron movement through electron transport chains cross through ATP synthase which allows the proton to pass through them and use this free energy difference to photophosphorylate ADP making ATP.
Contents
[hide]
1 The chemiosmotic theory
2 The proton-motive force
2.1 Equations
3 In mitochondria
4 In plants
5 In prokaryotes
6 See also
7 References
8 Further reading
9 External links
The chemiosmotic theory[edit]
Peter D. Mitchell proposed the chemiosmotic hypothesis in 1961.[1] The theory suggests essentially that most adenosine triphosphate (ATP) synthesis in respiring cells comes from the electrochemical gradient across the inner membranes of mitochondria by using the energy of NADH and FADH2 formed from the breaking down of energy-rich molecules such as glucose. Mitchell won the Nobel Prize in 1976 for his chemiosmotic theory. [2]
Chemiosmosis in a mitochondrion.
Molecules such as glucose are metabolized to produce acetyl CoA as an energy-rich intermediate. The oxidation of acetyl coenzyme A (acetyl-CoA) in the mitochondrial matrix is coupled to the reduction of a carrier molecule such as nicotinamide adenine dinucleotide (NAD) and flavin adenine dinucleotide (FAD).[3] The carriers pass electrons to the electron transport chain (ETC) in the inner mitochondrial membrane, which in turn pass them to other proteins in the ETC. The energy available in the electrons is used to pump protons from the matrix across the stroma, storing energy in the form of a transmembrane electrochemical gradient. The protons move back across the inner membrane through the enzyme ATP synthase. The flow of protons back into the matrix of the mitochondrion via ATP synthase provides enough energy for ADP to combine with inorganic phosphate to form ATP. The electrons and protons at the last pump in the ETC are taken up by oxygen to form water.
This was a radical proposal at the time, and was not well accepted. The prevailing view was that the energy of electron transfer was stored as a stable high potential intermediate, a chemically more conservative concept. The problem with the older paradigm is that no high energy intermediate was ever found, and the evidence for proton pumping by the complexes of the electron transfer chain grew too great to be ignored. Eventually the weight of evidence began to favor the chemiosmotic hypothesis, and in 1978, Peter Mitchell was awarded the Nobel Prize in Chemistry.[4]
Chemiosmotic coupling is important for ATP production in mitochondria, chloroplasts[5] and many bacteria and archaea.[6]
The proton-motive force[edit]
Energy conversion by the inner mitochondrial membrane and chemiosmotic coupling between the chemical energy of redox reactions in the respiratory chain and the oxidative phosphorylation catalysed by the ATP synthase [7][8] (sometimes called as "mitochondrial mushrooms").
The movement of ions across the membrane depends on a combination of two factors:
Diffusion force caused by a concentration gradient - all particles tend to diffuse from higher concentration to lower.
Electrostatic force caused by electrical potential gradient - cations like protons H+ tend to diffuse down the electrical potential, from the positive (P) side of the membrane to the negative (N) side. Anions diffuse spontaneously in the opposite direction.
These two gradients taken together can be expressed as an electrochemical gradient.
Lipid bilayers of biological membranes, however, are barriers for ions. This is why energy can be stored as a combination of these two gradients across the membrane. Only special membrane proteins like ion channels can sometimes allow ions to move across the membrane (see also: Membrane transport). In chemiosmotic theory transmembrane ATP synthases are very important. They convert energy of spontaneous flow of protons through them into chemical energy of ATP bonds.
Hence researchers created the term proton-motive force (PMF), derived from the electrochemical gradient mentioned earlier. It can be described as the measure of the potential energy stored as a combination of proton and voltage (electrical potential) gradients across a membrane. The electrical gradient is a consequence of the charge separation across the membrane (when the protons H+ move without a counterion, such as chloride Cl−).
In most cases the proton-motive force is generated by an electron transport chain which acts as a proton pump, using the Gibbs free energy of redox reactions to pump protons (hydrogen ions) out across the membrane, separating the charge across the membrane. In mitochondria, energy released by the electron transport chain is used to move protons from the mitochondrial matrix (N side) to the stroma (P side). Moving the protons out of the mitochondrion creates a lower concentration of positively charged protons inside it, resulting in excess negative charge on the inside of the membrane. The electrical potential gradient is about -170 mV [7], negative inside (N). These gradients - charge difference and the proton concentration difference both create a combined electrochemical gradient across the membrane, often expressed as the proton-motive force (PMF). In mitochondria, the PMF is almost entirely made up of the electrical component but in chloroplasts the PMF is made up mostly of the pH gradient because the charge of protons H+ is neutralized by the movement of Cl− and other anions. In either case, the PMF needs to be greater than about 460 mV (45 kJ/mol) for the ATP synthase to be able to make ATP.
Equations[edit]
The proton-motive force is derived from the Gibbs free energy. Let N denote the inside of a cell, and let P denote the outside. Then[7]
Δ G = z F Δ ψ + R T ln  [ X z + ] N [ X z + ] P {\displaystyle \Delta \!G=zF\Delta \!\psi +RT\ln {\frac {[\mathrm {X} ^{z+}]_{\text{N}}}{[\mathrm {X} ^{z+}]_{\text{P}}}}}
where
Δ G {\displaystyle \Delta \!G} is the Gibbs free energy change per unit amount of cations transferred from P to N;
z {\displaystyle z} is the charge number of the cation X z + {\displaystyle \mathrm {X} ^{z+}} ;
Δ ψ {\displaystyle \Delta \psi } is the electric potential of N relative to P;
[ X z + ] P {\displaystyle [\mathrm {X} ^{z+}]_{\text{P}}} and [ X z + ] N {\displaystyle [\mathrm {X} ^{z+}]_{\text{N}}} are the cation concentrations at P and N, respectively;
F {\displaystyle F} is the Faraday constant;
R {\displaystyle R} is the gas constant; and
T {\displaystyle T} is the temperature.
The molar Gibbs free energy change Δ G {\displaystyle \Delta \!G} is frequently interpreted as a molar electrochemical ion potential Δ μ X z + = Δ G {\displaystyle \Delta \!\mu _{\mathrm {X} ^{z+}}=\Delta \!G} .
For an electrochemical proton gradient z = 1 {\displaystyle z=1} and as a consequence:
Δ μ H + = F Δ ψ + R T ln  [ H + ] N [ H + ] P = F Δ ψ − ( ln  10 ) R T Δ p H {\displaystyle \Delta \!\mu _{\mathrm {H} ^{+}}=F\Delta \!\psi +RT\ln {\frac {[\mathrm {H} ^{+}]_{\text{N}}}{[\mathrm {H} ^{+}]_{\text{P}}}}=F\Delta \!\psi -(\ln 10)RT\Delta \mathrm {pH} }
A diagram of chemiosmotic phosphorylation
where
Δ p H = p H N − p H P {\displaystyle \Delta \!\mathrm {pH} =\mathrm {pH} _{\mathrm {N} }-\mathrm {pH} _{\mathrm {P} }} .
Mitchell defined the proton-motive force (PMF) as
Δ p = − Δ μ H + F {\displaystyle \Delta \!p=-{\frac {\Delta \!\mu _{\mathrm {H^{+}} }}{F}}} .
For example, Δ μ H + = 1 k J m o l − 1 {\displaystyle \Delta \!\mu _{\mathrm {H} ^{+}}=1\,\mathrm {kJ} \,\mathrm {mol} ^{-1}} implies Δ p = 10.4 m V {\displaystyle \Delta \!p=10.4\,\mathrm {mV} } . At 298 K {\displaystyle 298\,\mathrm {K} } this equation takes the form:
Δ p = − Δ ψ + ( 59.1 m V ) Δ p H {\displaystyle \Delta \!p=-\Delta \!\psi +\left(59.1\,\mathrm {mV} \right)\Delta \!\mathrm {pH} } .
Note that for spontaneous proton import from the P side (relatively more positive and acidic) to the N side (relatively more negative and alkaline), Δ μ H + {\displaystyle \Delta \!\mu _{\mathrm {H} ^{+}}} is negative (similar to Δ G {\displaystyle \Delta \!G} ) whereas PMF is positive (similar to redox cell potential Δ E {\displaystyle \Delta E} ).
It is worth noting that, as with any transmembrane transport process, the PMF is directional. The sign of the transmembrane electric potential difference Δ ψ {\displaystyle \Delta \!\psi } is chosen to represent the change in potential energy per unit charge flowing into the cell as above. Furthermore, due to redox-driven proton pumping by coupling sites, the proton gradient is always inside-alkaline. For both of these reasons, protons flow in spontaneously, from the P side to the N side; the available free energy is used to synthesize ATP (see below). For this reason, PMF is defined for proton import, which is spontaneous. PMF for proton export, i.e., proton pumping as catalyzed by the coupling sites, is simply the negative of PMF(import).
The spontaneity of proton import (from the P to the N side) is universal in all bioenergetic membranes.[9] This fact was not recognized before the 1990s, because the chloroplast thylakoid lumen was interpreted as an interior phase, but in fact it is topologically equivalent to the exterior of the chloroplast. Azzone et al. stressed that the inside phase (N side of the membrane) is the bacterial cytoplasm, mitochondrial matrix, or chloroplast stroma; the outside (P) side is the bacterial periplasmic space, mitochondrial intermembrane space, or chloroplast lumen. Furthermore, 3D tomography of the mitochondrial inner membrane shows its extensive invaginations to be stacked, similar to thylakoid disks; hence the mitochondrial intermembrane space is topologically quite similar to the chloroplast lumen.[10]:

The energy expressed here as Gibbs free energy, electrochemical proton gradient, or proton-motive force (PMF), is a combination of two gradients across the membrane:
the concentration gradient (via Δ p H {\displaystyle \Delta \!\mathrm {pH} } ) and
electric potential gradient Δ ψ {\displaystyle \Delta \!\psi } .
When a system reaches equilibrium, Δ p = 0 {\displaystyle \Delta \!p=0} ; nevertheless, the concentrations on either side of the membrane need not be equal. Spontaneous movement across the potential membrane is determined by both concentration and electric potential gradients.
The molar Gibbs free energy Δ G p {\displaystyle \Delta \!G_{\mathrm {p} }} of ATP synthesis
A D P 4 − + H + + H O P O 3 2 − → A T P 4 − + H 2 O {\displaystyle \mathrm {ADP} ^{4-}+\mathrm {H} ^{+}+\mathrm {HOPO} _{3}^{2-}\rightarrow \mathrm {ATP} ^{4-}+\mathrm {H_{2}O} }
is also called phosphorylation potential. The equilibrium concentration ratio [ H + ] / [ A T P ] {\displaystyle [\mathrm {H} ^{+}]/[\mathrm {ATP} ]} can be calculated by comparing Δ p {\displaystyle \Delta \!p} and Δ G p {\displaystyle \Delta \!G_{\mathrm {p} }} , for example in case of the mammalian mitochondrion[11]:
H+ / ATP = ΔGp / (Δp / 10.4 kJ·mol−1/mV) = 40.2 kJ·mol−1 / (173.5 mV / 10.4 kJ·mol−1/mV) = 40.2 / 16.7 = 2.4. The actual ratio of the proton-binding c-subunit to the ATP-synthesizing beta-subunit copy numbers is 8/3 = 2.67, showing that under these conditions, the mitochondrion functions at 90% (2.4/2.67) efficiency.[12]
In fact, the thermodynamic efficiency is lower in eukaryotic cells because ATP must be exported from the matrix to the cytoplasm, and ADP and phosphate must be imported from the cytoplasm. This "costs" one "extra" proton import per ATP,[7][8] hence the actual efficiency is only 65% (= 2.4/3.67).
In mitochondria[edit]
Directions of chemiosmotic proton transfer in the mitochondrion, chloroplast and in gram-negative bacterial cells (cellular respiration and photosynthesis). The bacterial cell wall is omitted, gram-positive bacterial cells do not have outer membrane.[7]
The complete breakdown of glucose in the presence of oxygen is called cellular respiration. The last steps of this process occur in mitochondria. The reduced molecules NADH and FADH2 are generated by the Krebs cycle, glycolysis, and pyruvate processing. These molecules pass electrons to an electron transport chain, which uses the energy released to create a proton gradient across the inner mitochondrial membrane. ATP synthase then uses the energy stored in this gradient to make ATP. This process is called oxidative phosphorylation because oxygen is the final electron acceptor and the energy released by reducing oxygen to water is used to phosphorylate ADP and generate ATP.
In plants[edit]
The light reactions of photosynthesis generate ATP by the action of chemiosmosis. The photons in sunlight are received by the antenna complex of Photosystem II, which excites electrons to a higher energy level. These electrons travel down an electron transport chain, causing protons to be actively pumped across the thylakoid membrane into the thylakoid lumen. These protons then flow down their electrochemical potential gradient through an enzyme called ATP-synthase, creating ATP by the phosphorylation of ADP to ATP. The electrons from the initial light reaction reach Photosystem I, then are raised to a higher energy level by light energy and then received by an electron acceptor and reduce NADP+ to NADPH. The electrons lost from Photosystem II get replaced by the oxidation of water, which is "split" into protons and oxygen by the oxygen-evolving complex (OEC, also known as WOC, or the water-oxidizing complex). To generate one molecule of diatomic oxygen, 10 photons must be absorbed by photosystems I and II, four electrons must move through the two photosystems, and 2 NAPDH are generated (later used for carbon dioxide fixation in the Calvin Cycle).
In prokaryotes[edit]
Chemiosmotic coupling between the energy of sunlight, bacteriorhodopsin and phosphorylation (chemical energy) during photosynthesis in halophilic bacteria Halobacterium salinarum (syn. H. halobium). The bacterial cell wall is omitted. [7][8]
Bacteria and archaea also can use chemiosmosis to generate ATP. Cyanobacteria, green sulfur bacteria, and purple bacteria synthesize ATP by a process called photophosphorylation. These bacteria use the energy of light to create a proton gradient using a photosynthetic electron transport chain. Non-photosynthetic bacteria such as E. coli also contain ATP synthase. In fact, mitochondria and chloroplasts are believed to have been formed when early eukaryotic cells ingested bacteria that could transfer energy using chemiosmosis. This is called the endosymbiotic theory.
Chemiosmotic phosphorylation is the third pathway that produces ATP from inorganic phosphate and an ADP molecule. This process is part of oxidative phosphorylation.
See also[edit]
Bacteriorhodopsin
Cellular respiration
Citric acid cycle
Electrochemical gradient
Glycolysis
Oxidative phosphorylation
References[edit]
Jump up ^ Peter Mitchell (1961). "Coupling of phosphorylation to electron and hydrogen transfer by a chemi-osmotic type of mechanism". Nature. 191 (4784): 144–148. Bibcode:1961Natur.191..144M. PMID 13771349. doi:10.1038/191144a0.
Jump up ^ "Chemiosmosis". biologydictionary.net. Retrieved 7 August 2017.
Jump up ^ Alberts, Bruce; Alexander Johnson; Julian Lewis; Martin Raff; Keith Roberts; Peter Walter (2002). "Proton Gradients Produce Most of the Cell's ATP". Molecular Biology of the Cell. Garland. ISBN 0-8153-4072-9.
Jump up ^ The Nobel Prize in Chemistry 1978.
Jump up ^ Cooper, Geoffrey M. (2000). "Figure 10.22: Electron transport and ATP synthesis during photosynthesis". The Cell: A Molecular Approach (2nd ed.). Sinauer Associates, Inc. ISBN 0-87893-119-8.
Jump up ^ Alberts, Bruce; Alexander Johnson; Julian Lewis; Martin Raff; Keith Roberts; Peter Walter (2002). "Figure 14-32: The importance of H+-driven transport in bacteria". Molecular Biology of the Cell. Garland. ISBN 0-8153-4072-9.
^ Jump up to: a b c d e f Nicholls D. G.; Ferguson S. J. (1992). Bioenergetics 2 (2nd ed.). San Diego: Academic Press. ISBN 9780125181242.
^ Jump up to: a b c Stryer, Lubert (1995). Biochemistry (fourth ed.). New York - Basingstoke: W. H. Freeman and Company. ISBN 978-0716720096.
Jump up ^ Azzone et al (1993) "Transmembrane Measurements Across Bioenergetic Membranes". Biochim. Biophys. Acta BBA - Bioenerg. 1183 (1): 1-3.
Jump up ^ Silverstein, Todd (2014). "An exploration of how the thermodynamic efficiency of bioenergetic membrane systems varies with c-subunit stoichiometry of F1F0 ATP synthases". J. Bioenerg. Biomembr. 46:229-241.
Jump up ^ Silverstein, Todd (2014). "An exploration of how the thermodynamic efficiency of bioenergetic membrane systems varies with c-subunit stoichiometry of F1F0 ATP synthases". J. Bioenerg. Biomembr. 46:229-241.
Jump up ^ Silverstein, Todd (2014). "An exploration of how the thermodynamic efficiency of bioenergetic membrane systems varies with c-subunit stoichiometry of F1F0 ATP synthases". J. Bioenerg. Biomembr.46:229-241.
Further reading[edit]
Biochemistry textbook reference, from the NCBI bookshelf – Jeremy M. Berg; John L. Tymoczko; Lubert Stryer (eds.). "18.4. A Proton Gradient Powers the Synthesis of ATP". Biochemistry (5th ed.). W. H. Freeman.
Technical reference relating one set of experiments aiming to test some tenets of the chemiosmotic theory – Seiji Ogawa & Tso Ming Lee (1984). "The Relation between the Internal Phosphorylation Potential and the Proton Motive Force in Mitochondria during ATP Synthesis and Hydrolysis". Journal of Biological Chemistry. 259 (16): 10004–10011. PMID 6469951.
External links[edit]
Chemiosmosis (University of Wisconsin)
Retrieved from "https://en.wikipedia.org/w/index.php?title=Chemiosmosis&oldid=801132350"
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/wiki/Respiratory_chain
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Respiratory chain

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/wiki/Electrostatic_force
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Electrostatic force

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/wiki/Electrical_potential
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Electrical potential

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values: {\displaystyle [\mathrm {X} ^{z+}]_{\text{P}}}

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values: {\displaystyle [\mathrm {X} ^{z+}]_{\text{N}}}

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values: {\displaystyle \Delta \!p=10.4\,\mathrm {mV} }

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values: {\displaystyle \Delta \!\mu _{\mathrm {H} ^{+}}}

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values: {\displaystyle \Delta \!G_{\mathrm {p} }}

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/wiki/Mitochondrial_membrane
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Mitochondrial membrane

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/wiki/Light-dependent_reaction
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Light-dependent reaction

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/wiki/Thylakoid_membrane
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Thylakoid membrane

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/wiki/Endosymbiotic_theory
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Endosymbiotic theory

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PubMed Identifier

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//www.ncbi.nlm.nih.gov/pubmed/13771349

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//doi.org/10.1038%2F191144a0

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NCBI bookshelf

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//www.ncbi.nlm.nih.gov/pubmed/6469951

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