The Hill reaction is the light-driven transfer of electrons from water to Hill reagents (non-physiological oxidants) 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.
The evolution of oxygen during the light-dependent steps in photosynthesis (Hill reaction) was proposed and proven by British biochemist Robin Hill. He demonstrated that isolated chloroplasts would make oxygen (O2) but not fix carbon dioxide (CO2). This is evidence that the light and dark reactions occur at different sites within the cell. [1] [2] [3]
Hill's finding was that the origin of oxygen in photosynthesis is water (H2O) not carbon dioxide (CO2) as previously believed. Hill's observation of chloroplasts in dark conditions and in the absence of CO2, showed that the artificial electron acceptor was oxidized but not reduced, terminating the process, but without production of oxygen and sugar. This observation allowed Hill to conclude that oxygen is released during the light-dependent steps (Hill reaction) of photosynthesis. [4]
Hill also discovered Hill reagents, artificial electron acceptors that participate in the light reaction, such as Dichlorophenolindophenol (DCPIP), a dye that changes color when reduced. These dyes permitted the finding of electron transport chains during photosynthesis.
Further studies of the Hill reaction were made in 1957 by plant physiologist Daniel I. Arnon. Arnon studied the Hill reaction using a natural electron acceptor, NADP. He demonstrated the light-independent reaction, observing the reaction under dark conditions with an abundance of carbon dioxide. He found that carbon fixation was independent of light. Arnon effectively separated the light-dependent reaction, which produces ATP, NADPH, H+ and oxygen, from the light-independent reaction that produces sugars.
Photosynthesis is the process in which light energy is absorbed and converted to chemical energy. This chemical energy is eventually used in the conversion of carbon dioxide to sugar in plants.
During photosynthesis, natural electron acceptor NADP is reduced to NADPH in chloroplasts. [5] The following equilibrium reaction takes place.
A reduction reaction that stores energy as NADPH:
An oxidation reaction as NADPH's energy is used elsewhere:
Ferredoxin, also known as an NADP+ reductase, is an enzyme that catalyzes the reduction reaction. It is easy to oxidize NADPH but difficult to reduce NADP+, hence a catalyst is beneficial. Cytochromes are conjugate proteins that contain a haem group. [5] The iron atom from this group undergoes redox reactions:
The light-dependent redox reaction takes place before the light-independent reaction in photosynthesis. [6]
Isolated chloroplasts placed under light conditions but in the absence of CO2, reduce and then oxidize artificial electron acceptors, allowing the process to proceed. Oxygen (O2) is released as a byproduct, but not sugar (CH2O).
Chloroplasts placed under dark conditions and in the absence of CO2, oxidize the artificial acceptor but do not reduce it, terminating the process, without production of oxygen or sugar. [4]
The association of phosphorylation and the reduction of an electron acceptor such as ferricyanide increase similarly with the addition of phosphate, magnesium (Mg), and ADP. The existence of these three components is important for maximal reductive and phosphorylative activity. Similar increases in the rate of ferricyanide reduction can be stimulated by a dilution technique. Dilution does not cause a further increase in the rate in which ferricyanide is reduced with the accumulation of ADP, phosphate, and Mg to a treated chloroplast suspension. ATP inhibits the rate of ferricyanide reduction. Studies of light intensities revealed that the effect was largely on the light-independent steps of the Hill reaction. These observations are explained in terms of a proposed method in which phosphate esterifies during electron transport reactions, reducing ferricyanide, while the rate of electron transport is limited by the rate of phosphorylation. An increase in the rate of phosphorylation increases the rate by which electrons are transported in the electron transport system. [7]
It is possible to introduce an artificial electron acceptor into the light reaction, such as a dye that changes color when it is reduced. These are known as Hill reagents. These dyes permitted the finding of electron transport chains during photosynthesis. Dichlorophenolindophenol (DCPIP), an example of these dyes, is widely used by experimenters. DCPIP is a dark blue solution that becomes lighter as it is reduced. It provides experimenters with a simple visual test and easily observable light reaction. [8]
In another approach to studying photosynthesis, light-absorbing pigments such as chlorophyll can be extracted from chloroplasts. Like so many important biological systems in the cell, the photosynthetic system is ordered and compartmentalized in a system of membranes. [9]
Photosynthesis is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabolism. Photosynthesis usually refers to oxygenic photosynthesis, a process that produces oxygen. Photosynthetic organisms store the chemical energy so produced within intracellular organic compounds like sugars, glycogen, cellulose and starches. To use this stored chemical energy, an organism's cells metabolize the organic compounds through cellular respiration. Photosynthesis plays a critical role in producing and maintaining the oxygen content of the Earth's atmosphere, and it supplies most of the biological energy necessary for complex life on Earth.
Redox is a type of chemical reaction in which the oxidation states of the reactants change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a decrease in the oxidation state. The oxidation and reduction processes occur simultaneously in the chemical reaction.
A dehydrogenase is an enzyme belonging to the group of oxidoreductases that oxidizes a substrate by reducing an electron acceptor, usually NAD+/NADP+ or a flavin coenzyme such as FAD or FMN. Like all catalysts, they catalyze reverse as well as forward reactions, and in some cases this has physiological significance: for example, alcohol dehydrogenase catalyzes the oxidation of ethanol to acetaldehyde in animals, but in yeast it catalyzes the production of ethanol from acetaldehyde.
Cellular respiration is the process by which biological fuels are oxidized in the presence of an inorganic electron acceptor, such as oxygen, to drive the bulk production of adenosine triphosphate (ATP), which contains energy. Cellular respiration may be described as a set of metabolic reactions and processes that take place in the cells of organisms to convert chemical energy from nutrients into ATP, and then release waste products.
In chemistry, a reducing agent is a chemical species that "donates" an electron to an electron recipient.
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 or stromal thylakoids, which join granum stacks together as a single functional compartment.
C4 carbon fixation or the Hatch–Slack pathway is one of three known photosynthetic processes of carbon fixation in plants. It owes the names to the 1960s discovery by Marshall Davidson Hatch and Charles Roger Slack.
Photorespiration (also known as the oxidative photosynthetic carbon cycle or C2 cycle) refers to a process in plant metabolism where the enzyme RuBisCO oxygenates RuBP, wasting some of the energy produced by photosynthesis. The desired reaction is the addition of carbon dioxide to RuBP (carboxylation), a key step in the Calvin–Benson cycle, but approximately 25% of reactions by RuBisCO instead add oxygen to RuBP (oxygenation), creating a product that cannot be used within the Calvin–Benson cycle. This process lowers the efficiency of photosynthesis, potentially lowering photosynthetic output by 25% in C3 plants. Photorespiration involves a complex network of enzyme reactions that exchange metabolites between chloroplasts, leaf peroxisomes and mitochondria.
The term amphibolism is used to describe a biochemical pathway that involves both catabolism and anabolism. Catabolism is a degradative phase of metabolism in which large molecules are converted into smaller and simpler molecules, which involves two types of reactions. First, hydrolysis reactions, in which catabolism is the breaking apart of molecules into smaller molecules to release energy. Examples of catabolic reactions are digestion and cellular respiration, where sugars and fats are broken down for energy. Breaking down a protein into amino acids, or a triglyceride into fatty acids, or a disaccharide into monosaccharides are all hydrolysis or catabolic reactions. Second, oxidation reactions involve the removal of hydrogens and electrons from an organic molecule. Anabolism is the biosynthesis phase of metabolism in which smaller simple precursors are converted to large and complex molecules of the cell. Anabolism has two classes of reactions. The first are dehydration synthesis reactions; these involve the joining of smaller molecules together to form larger, more complex molecules. These include the formation of carbohydrates, proteins, lipids and nucleic acids. The second are reduction reactions, in which hydrogens and electrons are added to a molecule. Whenever that is done, molecules gain energy.
Nicotinamide adenine dinucleotide phosphate, abbreviated NADP or, in older notation, TPN (triphosphopyridine nucleotide), is a cofactor used in anabolic reactions, such as the Calvin cycle and lipid and nucleic acid syntheses, which require NADPH as a reducing agent ('hydrogen source'). NADPH is the reduced form, whereas NADP+ is the oxidized form. NADP+ is used by all forms of cellular life. NADP+ is essential for life because it is needed for cellular respiration.
The Calvin cycle, light-independent reactions, bio synthetic phase, dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis is a series of chemical reactions that convert carbon dioxide and hydrogen-carrier compounds into glucose. The Calvin cycle is present in all photosynthetic eukaryotes and also many photosynthetic bacteria. In plants, these reactions occur in the stroma, the fluid-filled region 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 chemical energy of ATP and reducing power of NADPH from the light dependent reactions to produce sugars for the plant to use. These substrates are used in a series of reduction-oxidation (redox) reactions to produce sugars in a step-wise process; there is no direct reaction that converts several molecules of CO2 to a sugar. There are three phases to the light-independent reactions, collectively called the Calvin cycle: carboxylation, reduction reactions, and ribulose 1,5-bisphosphate (RuBP) regeneration.
Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.
In the process of photosynthesis, the phosphorylation of ADP to form ATP using the energy of sunlight is called photophosphorylation. Cyclic photophosphorylation occurs in both aerobic and anaerobic conditions, driven by the main primary source of energy available to living organisms, which is sunlight. All organisms produce a phosphate compound, ATP, which is the universal energy currency of life. In photophosphorylation, light energy is used to pump protons across a biological membrane, mediated by flow of electrons through an electron transport chain. This stores energy in a proton gradient. As the protons flow back through an enzyme called ATP synthase, ATP is generated from ADP and inorganic phosphate. ATP is essential in the Calvin cycle to assist in the synthesis of carbohydrates from carbon dioxide and NADPH.
Artificial photosynthesis is a chemical process that biomimics the natural process of photosynthesis. The term artificial photosynthesis is used loosely, referring to any scheme for capturing and then storing energy from sunlight by producing a fuel, specifically a solar fuel. An advantage of artificial photosynthesis would be that the solar energy could converted and stored. By contrast, using photovoltaic cells, sunlight is converted into electricity and then converted again into chemical energy for storage, with some necessary losses of energy associated with the second conversion. The byproducts of these reactions are environmentally friendly. Artificially photosynthesized fuel would be a carbon-neutral source of energy, but it has never been demonstrated in any practical sense. The economics of artificial photosynthesis are noncompetitive.
A photosynthetic reaction center is a complex of several proteins, biological pigments, and other co-factors that together execute the primary energy conversion reactions of photosynthesis. Molecular excitations, either originating directly from sunlight or transferred as excitation energy via light-harvesting antenna systems, give rise to electron transfer reactions along the path of a series of protein-bound co-factors. These co-factors are light-absorbing molecules (also named chromophores or pigments) such as chlorophyll and pheophytin, as well as quinones. The energy of the photon is used to excite an electron of a pigment. The free energy created is then used, via a chain of nearby electron acceptors, for a transfer of hydrogen atoms (as protons and electrons) from H2O or hydrogen sulfide towards carbon dioxide, eventually producing glucose. These electron transfer steps ultimately result in the conversion of the energy of photons to chemical energy.
Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.
2,6-Dichlorophenolindophenol is a chemical compound used as a redox dye. When oxidized, DCPIP is blue with a maximal absorption at 600 nm; when reduced, DCPIP is colorless.
Malate dehydrogenase (oxaloacetate-decarboxylating) (NADP+) (EC 1.1.1.40) or NADP-malic enzyme (NADP-ME) is an enzyme that catalyzes the chemical reaction in the presence of a bivalent metal ion:
In enzymology, a ferredoxin-NADP+ reductase (EC 1.18.1.2) abbreviated FNR, is an enzyme that catalyzes the chemical reaction
Light-dependent reactions are certain photochemical reactions involved in photosynthesis, the main process by which plants acquire energy. There are two light dependent reactions: the first occurs at photosystem II (PSII) and the second occurs at photosystem I (PSI).
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