Mehler reaction

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The Mehler reaction is named after Alan H. Mehler, who, in 1951, presented data to the effect that isolated chloroplasts reduce oxygen to form hydrogen peroxide (H
2
O
2
). [1] Mehler observed that the H
2
O
2
formed in this way does not present an active intermediate in photosynthesis; rather, as a reactive oxygen species, it can be toxic to surrounding biological processes as an oxidizing agent. In scientific literature, the Mehler reaction often is used interchangeably with the Water-Water Cycle [2] to refer to the formation of H
2
O
2
by photosynthesis. Sensu stricto, the Water Water Cycle encompasses the Hill reaction, in which water is split to form oxygen, as well as the Mehler Reaction, in which oxygen is reduced to form H
2
O
2
and, finally, the scavenging of this H
2
O
2
by antioxidants to form water.

Beginning in the 1970s, Professor Kozi Asada elucidated that oxygen can be reduced by electrons emerging from ferredoxin of photosystem I, to form superoxide, which is then reduced by superoxide dismutase to form H
2
O
2
. This photochemical H
2
O
2
is then reduced by the action of ascorbate peroxidase to form water and oxidized ascorbate. Asada argued that oxygen presents an important sink for excess excitation energy acquired during plant exposure to bright light. He would often begin seminars by asking: 'Why aren't plants sunburnt despite being exposed to light?'. [3]

How much of a photoprotective role the Water Water Cycle plays has been occasion for some debate. In terrestrial plants, transfer of electrons to oxygen from ferredoxin at PSI accounts for easily less than 10% of total photosynthetic electron transport. [4] [5] [6] In algae and other uni-cellular photosynthetic organisms, however, this amount can account for 20 to 30% of total electron transport. It is possible that the reduction of oxygen by free electrons emerging from PSI prevents components of the electron transport chain from becoming over-reduced. [7]

The Water Water Cycle is not related to photorespiration, as it comprises different reactions and results in no net oxygen consumption.

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Thylakoid

Thylakoids are membrane-bound compartments inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thylakoids frequently form stacks of disks referred to as grana. Grana are connected by intergranal/ stroma thylakoids, which join granum stacks together as a single functional compartment.

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2
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Photorespiration

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Reactive oxygen species Class of compounds

Reactive oxygen species (ROS) are highly reactive chemical molecules formed due to the electron acceptability of O2. Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.

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The Calvin cycle,light-independent reactions, bio synthetic phase,dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis are the chemical reactions that convert carbon dioxide and other compounds into glucose. These reactions occur in the stroma, the fluid-filled area of a chloroplast outside the thylakoid membranes. These reactions take the products of light-dependent reactions and perform further chemical processes on them. The Calvin cycle uses the reducing powers ATP and NADPH from the light dependent reactions to produce sugars for the plant to use. These substrates are used in a series of reduction-oxidation reactions to produce sugars in a step-wise process. There is no direct reaction that converts CO2 to a sugar because all of the energy would be lost to heat.] There are three phases to the light-independent reactions, collectively called the Calvin cycle: carbon fixation, reduction reactions, and ribulose 1,5-bisphosphate (RuBP) regeneration.

Photosystem

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Cytochrome b<sub>6</sub>f complex

The cytochrome b6f complex is an enzyme found in the thylakoid membrane in chloroplasts of plants, cyanobacteria, and green algae, that catalyzes the transfer of electrons from plastoquinol to plastocyanin. The reaction is analogous to the reaction catalyzed by cytochrome bc1 of the mitochondrial electron transport chain. During photosynthesis, the cytochrome b6f complex is one step along the chain that transfers electrons from Photosystem II to Photosystem I, and at the same time pumps protons into the thylakoid space that contribute to create an electrochemical (energy) gradient which is later used to synthesize ATP from ADP.

Photosynthetic reaction centre

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DCMU

DCMU is an algicide and herbicide of the arylurea class that inhibits photosynthesis. It was introduced by Bayer in 1954 under the trade name of Diuron.

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Sedoheptulose-bisphosphatase

Sedoheptulose-bisphosphatase is an enzyme that catalyzes the removal of a phosphate group from sedoheptulose 1,7-bisphosphate to produce sedoheptulose 7-phosphate. SBPase is an example of a phosphatase, or, more generally, a hydrolase. This enzyme participates in the Calvin cycle.

Dioxygen plays an important role in the energy metabolism of living organisms. Free oxygen is produced in the biosphere through photolysis of water during photosynthesis in cyanobacteria, green algae, and plants. During oxidative phosphorylation in cellular respiration, the chemical energy of oxygen is released as it is reduced to water, thus closing the biological water-oxygen redox cycle.

Foyer-Halliwell-Asada pathway

Light-dependent reactions Photosynthetic reactions

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.

All living cells produce reactive oxygen species (ROS) as a byproduct of metabolism. ROS are reduced oxygen intermediates that include the superoxide radical (O2) and the hydroxyl radical (OH•), as well as the non-radical species hydrogen peroxide (H2O2). These ROS are important in the normal functioning of cells, playing a role in signal transduction and the expression of transcription factors. However, when present in excess, ROS can cause damage to proteins, lipids and DNA by reacting with these biomolecules to modify or destroy their intended function. As an example, the occurrence of ROS have been linked to the aging process in humans, as well as several other diseases including Alzheimer's, rheumatoid arthritis, Parkinson's, and some cancers. Their potential for damage also makes reactive oxygen species useful in direct protection from invading pathogens, as a defense response to physical injury, and as a mechanism for stopping the spread of bacteria and viruses by inducing programmed cell death.

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Hill reaction

The Hill reaction is the light-driven transfer of electrons from water to Hill reagents in a direction against the chemical potential gradient as part of photosynthesis. Robin Hill discovered the reaction in 1937. He demonstrated that the process by which plants produce oxygen is separate from the process that converts carbon dioxide to sugars.

References

  1. Mehler, Alan (1951). "Studies on reactions of illuminated chloroplasts: I. Mechanism of the reduction of oxygen and other hill reagents". Archives of Biochemistry and Biophysics. 33 (1): 65–77. doi:10.1016/0003-9861(51)90082-3.
  2. Asada, Kozi (June 1999). "The Water-Cycle in Chloroplasts: Scavenging of Active Oxygens and Dissipation of Excess Photons". Annual Review of Plant Physiology and Plant Molecular Biology. 50: 601-639. doi:10.1146/annurev.arplant.50.1.601.
  3. Mano, Endo and Miyake (2016). "How do photosynthetic organisms manage light stress? A tribute to the late Professor Kozi Asada". Plant and Cell Physiology. 57 (7): 1351-1353. doi: 10.1093/pcp/pcw116 .
  4. Badger, M.; von Caemmerer, S.; Ruuska, S.; Nakano, H. (2000). "Electron flow to oxygen in higher plants and algae: rates and control of direct photoreduction (Mehler reaction) and rubisco oxygenase". Philosophical Transactions of the Royal Society of London B: Biological Sciences. 355 (1402): 1433–1446. doi:10.1098/rstb.2000.0704. PMC   1692866 . PMID   11127997.
  5. Ruuska, S.A.; Badger, M.R.; von Caemmerer, S. (2000). "Photosynthetic electron sinks in transgenic tobacco with reduced amounts of Rubisco: little evidence for significant Mehler reaction". Journal of Experimental Botany. 51: 357–368. doi: 10.1093/jexbot/51.suppl_1.357 .
  6. Driever, S.M.; Baker, N. (2011). "The water–water cycle in leaves is not a major alternative electron sink for dissipation of excess excitation energy when CO
    2
    assimilation is restricted". Plant, Cell & Environment. 34 (5): 837–846. doi:10.1111/j.1365-3040.2011.02288.x.
  7. Heber, Ulrich (2002-01-01). "Irrungen, Wirrungen? The Mehler reaction in relation to cyclic electron transport in C3 plants". Photosynthesis Research. 73 (1–3): 223–231. doi:10.1023/A:1020459416987. ISSN   1573-5079. PMID   16245125.