In the process of photosynthesis, the phosphorylation of ADP to form ATP using the energy of sunlight is called photophosphorylation. Only two sources of energy are available to living organisms: sunlight and reduction-oxidation (redox) reactions. All organisms produce ATP, which is the universal energy currency of life. In photosynthesis this commonly involves photolysis, or photodissociation, of water and a continuous unidirectional flow of electrons from water to photosystem II.
In photophosphorylation, light energy is used to create a high-energy electron donor and a lower-energy electron acceptor. Electrons then move spontaneously from donor to acceptor through an electron transport chain.
ATP and reactions
ATP is made by an enzyme called ATP synthase. Both the structure of this enzyme and its underlying gene are remarkably similar in all known forms of life. The Calvin cycle is one of the most important part of photosynthesis.
ATP synthase is powered by a transmembrane electrochemical potential gradient, usually in the form of a proton gradient. The function of the electron transport chain is to produce this gradient. In all living organisms, a series of redox reactions is used to produce a transmembrane electrochemical potential gradient, or a so-called proton motive force (pmf).
Redox reactions are chemical reactions in which electrons are transferred from a donor molecule to an acceptor molecule. The underlying force driving these reactions is the Gibbs free energy of the reactants and products. The Gibbs free energy is the energy available (“free”) to do work. Any reaction that decreases the overall Gibbs free energy of a system will proceed spontaneously (given that the system is isobaric and also adiabatic), although the reaction may proceed slowly if it is kinetically inhibited.
The transfer of electrons from a high-energy molecule (the donor) to a lower-energy molecule (the acceptor) can be spatially separated into a series of intermediate redox reactions. This is an electron transport chain.
The fact that a reaction is thermodynamically possible does not mean that it will actually occur. A mixture of hydrogen gas and oxygen gas does not spontaneously ignite. It is necessary either to supply an activation energy or to lower the intrinsic activation energy of the system, in order to make most biochemical reactions proceed at a useful rate. Living systems use complex macromolecular structures to lower the activation energies of biochemical reactions.
It is possible to couple a thermodynamically favorable reaction (a transition from a high-energy state to a lower-energy state) to a thermodynamically unfavorable reaction (such as a separation of charges, or the creation of an osmotic gradient), in such a way that the overall free energy of the system decreases (making it thermodynamically possible), while useful work is done at the same time. The principle that biological macromolecules catalyze a thermodynamically unfavorable reaction if and only if a thermodynamically favorable reaction occurs simultaneously, underlies all known forms of life.
Electron transport chains (most known as ETC) produce energy in the form of a transmembrane electrochemical potential gradient. This energy is used to do useful work. The gradient can be used to transport molecules across membranes. It can be used to do mechanical work, such as rotating bacterial flagella. It can be used to produce ATP and NADPH, high-energy molecules that are necessary for growth.
This form of photophosphorylation occurs on the stroma lamella or fret channels. In cyclic photophosphorylation, the high energy electron released from P700 to ps1 flow down in a cyclic pathway. In cyclic electron flow, the electron begins in a pigment complex called photosystem I, passes from the primary acceptor to ferredoxin and then to plastoquinone, then to cytochrome b6f (a similar complex to that found in mitochondria), and then to plastocyanin before returning to Photosystem-1. This transport chain produces a proton-motive force, pumping H+ ions across the membrane; this produces a concentration gradient that can be used to power ATP synthase during chemiosmosis. This pathway is known as cyclic photophosphorylation, and it produces neither O2 nor NADPH. Unlike non-cyclic photophosphorylation, NADP+ does not accept the electrons; they are instead sent back to cytochrome b6f complex.
In bacterial photosynthesis, a single photosystem is used, and therefore is involved in cyclic photophosphorylation. It is favored in anaerobic conditions and conditions of high irradiance and CO2 compensation points. 
The other pathway, non-cyclic photophosphorylation, is a two-stage process involving two different chlorophyll photosystems. Being a light reaction, non-cyclic photophosphorylation occurs in the thylakoid membrane. First, a water molecule is broken down into 2H+ + 1/2 O2 + 2e− by a process called photolysis (or light-splitting). The two electrons from the water molecule are kept in photosystem II, while the 2H+ and 1/2O2 are left out for further use. Then a photon is absorbed by chlorophyll pigments surrounding the reaction core center of the photosystem. The light excites the electrons of each pigment, causing a chain reaction that eventually transfers energy to the core of photosystem II, exciting the two electrons that are transferred to the primary electron acceptor, pheophytin. The deficit of electrons is replenished by taking electrons from another molecule of water. The electrons transfer from pheophytin to plastoquinone, which takes the 2e− from Pheophytin, and two H+ Ions from the stroma and forms PQH2, which later is broken into PQ, the 2e− is released to Cytochrome b6f complex and the two H+ ions are released into thylakoid lumen. The electrons then pass through the Cyt b6 and Cyt f. Then they are passed to plastocyanin, providing the energy for hydrogen ions (H+) to be pumped into the thylakoid space. This creates a gradient, making H+ ions flow back into the stroma of the chloroplast, providing the energy for the regeneration of ATP.
The photosystem II complex replaced its lost electrons from an external source; however, the two other electrons are not returned to photosystem II as they would in the analogous cyclic pathway. Instead, the still-excited electrons are transferred to a photosystem I complex, which boosts their energy level to a higher level using a second solar photon. The highly excited electrons are transferred to the acceptor molecule, but this time are passed on to an enzyme called Ferredoxin-NADP+ reductase which uses them to catalyse the reaction (as shown):
- NADP+ + 2H+ + 2e− → NADPH + H+
This consumes the H+ ions produced by the splitting of water, leading to a net production of 1/2O2, ATP, and NADPH+H+ with the consumption of solar photons and water.
The concentration of NADPH in the chloroplast may help regulate which pathway electrons take through the light reactions. When the chloroplast runs low on ATP for the Calvin cycle, NADPH will accumulate and the plant may shift from noncyclic to cyclic electron flow.
Early history of research
In 1950, first experimental evidence for the existence of photophosphorylation in vivo was presented by Otto Kandler using intact Chlorella cells and interpreting his findings as light-dependent ATP formation. In 1954, Daniel I. Arnon et.al. discovered photophosphorylation in vitro in isolated chloroplasts with the help of P32. His first review on the early research of photophosphorylation was published in 1956.
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- Citation needed
- Kandler, Otto (1950). "Über die Beziehungen zwischen Phosphathaushalt und Photosynthese. I. Phosphatspiegelschwankungen bei Chlorella pyrenoidosa als Folge des Licht-Dunkel-Wechsels" [On the relationship between the phosphate metabolism and photosynthesis I. Variations in phosphate levels in Chlorella pyrenoidosa as a consequence of light-dark changes] (PDF). Zeitschrift für Naturforschung. 5b (8): 423–437. doi:10.1515/znb-1950-0806.
- Arnon, Daniel I.; Allen, M.B.; Whatley, F.R. (1954). "Photosynthesis by isolated chloroplasts. II. Photophosphorylation, the conversion of light into phosphate bond energy". J Am Chem Soc. 76 (24): 6324–6329. doi:10.1021/ja01653a025 – via https://pubs.acs.org/doi/abs/10.1021/ja01653a025?journalCode=jacsat.
- Arnon, Daniel I. (1956). "Phosphorus metabolism and photosynthesis". Review of Plant Physiology. 7: 325–354. doi:10.1146/annurev.pp.07.060156.001545.
- Professor Luis Gordillo
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