125-400: Incl. Dictyochophyceae sensu lato is a photosynthetic lineage of heterokont algae. Photosynthetic Photosynthesis ( / ˌ f oʊ t ə ˈ s ɪ n θ ə s ɪ s / FOH -tə- SINTH -ə-sis ) is a system of biological processes by which photosynthetic organisms , such as most plants, algae , and cyanobacteria , convert light energy , typically from sunlight, into
250-440: A polarimeter since pure α- d -glucose has a specific rotation angle of +112.2° mL/(dm·g), pure β- d -glucose of +17.5° mL/(dm·g). When equilibrium has been reached after a certain time due to mutarotation, the angle of rotation is +52.7° mL/(dm·g). By adding acid or base, this transformation is much accelerated. The equilibration takes place via the open-chain aldehyde form. In dilute sodium hydroxide or other dilute bases,
375-500: A biochemical pump that collects carbon from the organ interior (or from the soil ) and not from the atmosphere. Cyanobacteria possess carboxysomes , which increase the concentration of CO 2 around RuBisCO to increase the rate of photosynthesis. An enzyme, carbonic anhydrase , located within the carboxysome, releases CO 2 from dissolved hydrocarbonate ions (HCO 3 ). Before the CO 2 can diffuse out, RuBisCO concentrated within
500-471: A component of starch), cellulases (named after cellulose), chitinases (named after chitin), and more. Furthermore, for the cleavage of disaccharides, there are maltase, lactase, sucrase, trehalase , and others. In humans, about 70 genes are known that code for glycosidases. They have functions in the digestion and degradation of glycogen, sphingolipids , mucopolysaccharides , and poly( ADP-ribose ). Humans do not produce cellulases, chitinases, or trehalases, but
625-409: A different leaf anatomy from C 3 plants, and fix the CO 2 at night, when their stomata are open. CAM plants store the CO 2 mostly in the form of malic acid via carboxylation of phosphoenolpyruvate to oxaloacetate , which is then reduced to malate. Decarboxylation of malate during the day releases CO 2 inside the leaves, thus allowing carbon fixation to 3-phosphoglycerate by RuBisCO. CAM
750-468: A general advancement in organic chemistry . This understanding occurred largely as a result of the investigations of Emil Fischer , a German chemist who received the 1902 Nobel Prize in Chemistry for his findings. The synthesis of glucose established the structure of organic material and consequently formed the first definitive validation of Jacobus Henricus van 't Hoff 's theories of chemical kinetics and
875-447: A lower tendency than other aldohexoses to react nonspecifically with the amine groups of proteins . This reaction— glycation —impairs or destroys the function of many proteins, e.g. in glycated hemoglobin . Glucose's low rate of glycation can be attributed to its having a more stable cyclic form compared to other aldohexoses, which means it spends less time than they do in its reactive open-chain form . The reason for glucose having
1000-566: A net gain of two ATP molecules (four ATP molecules are produced during glycolysis through substrate-level phosphorylation, but two are required by enzymes used during the process). In aerobic respiration, a molecule of glucose is much more profitable in that a maximum net production of 30 or 32 ATP molecules (depending on the organism) is generated. Click on genes, proteins and metabolites below to link to respective articles. Tumor cells often grow comparatively quickly and consume an above-average amount of glucose by glycolysis, which leads to
1125-598: A nutrition supplement in production of foodstuffs. Dextrose monohydrate is primarily consumed in North America as a corn syrup or high-fructose corn syrup . Anhydrous dextrose , on the other hand, is glucose that does not have any water molecules attached to it. Anhydrous chemical substances are commonly produced by eliminating water from a hydrated substance through methods such as heating or drying up (desiccation). Dextrose monohydrate can be dehydrated to anhydrous dextrose in industrial setting. Dextrose monohydrate
1250-480: A phospholipid outer membrane, and an intermembrane space. Enclosed by the membrane is an aqueous fluid called the stroma. Embedded within the stroma are stacks of thylakoids (grana), which are the site of photosynthesis. The thylakoids appear as flattened disks. The thylakoid itself is enclosed by the thylakoid membrane, and within the enclosed volume is a lumen or thylakoid space. Embedded in the thylakoid membrane are integral and peripheral membrane protein complexes of
1375-479: A photocomplex. When a photon is absorbed by a chromophore, it is converted into a quasiparticle referred to as an exciton , which jumps from chromophore to chromophore towards the reaction center of the photocomplex, a collection of molecules that traps its energy in a chemical form accessible to the cell's metabolism. The exciton's wave properties enable it to cover a wider area and try out several possible paths simultaneously, allowing it to instantaneously "choose"
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#17330927737741500-405: A photon by the antenna complex loosens an electron by a process called photoinduced charge separation . The antenna system is at the core of the chlorophyll molecule of the photosystem II reaction center. That loosened electron is taken up by the primary electron-acceptor molecule, pheophytin . As the electrons are shuttled through an electron transport chain (the so-called Z-scheme shown in
1625-422: A plant's photosynthetic response. Integrated chlorophyll fluorometer – gas exchange systems allow a more precise measure of photosynthetic response and mechanisms. While standard gas exchange photosynthesis systems can measure Ci, or substomatal CO 2 levels, the addition of integrated chlorophyll fluorescence measurements allows a more precise measurement of C C, the estimation of CO 2 concentration at
1750-450: A redox-active tyrosine residue that is oxidized by the energy of P680 . This resets the ability of P680 to absorb another photon and release another photo-dissociated electron. The oxidation of water is catalyzed in photosystem II by a redox-active structure that contains four manganese ions and a calcium ion ; this oxygen-evolving complex binds two water molecules and contains the four oxidizing equivalents that are used to drive
1875-550: A residue of carbon . Glucose has a pKa value of 12.16 at 25 °C (77 °F) in water. With six carbon atoms, it is classed as a hexose , a subcategory of the monosaccharides . d -Glucose is one of the sixteen aldohexose stereoisomers . The d - isomer , d -glucose, also known as dextrose, occurs widely in nature, but the l -isomer, l -glucose , does not. Glucose can be obtained by hydrolysis of carbohydrates such as milk sugar ( lactose ), cane sugar (sucrose), maltose , cellulose , glycogen , etc. Dextrose
2000-425: A ring of carbons closed by one oxygen atom. In aqueous solution, however, more than 99% of glucose molecules exist as pyranose forms. The open-chain form is limited to about 0.25%, and furanose forms exist in negligible amounts. The terms "glucose" and " D -glucose" are generally used for these cyclic forms as well. The ring arises from the open-chain form by an intramolecular nucleophilic addition reaction between
2125-399: A simpler method that employs a pigment similar to those used for vision in animals. The bacteriorhodopsin changes its configuration in response to sunlight, acting as a proton pump. This produces a proton gradient more directly, which is then converted to chemical energy. The process does not involve carbon dioxide fixation and does not release oxygen, and seems to have evolved separately from
2250-481: A source of carbon atoms to carry out photosynthesis; photoheterotrophs use organic compounds, rather than carbon dioxide, as a source of carbon. In plants, algae, and cyanobacteria, photosynthesis releases oxygen. This oxygenic photosynthesis is by far the most common type of photosynthesis used by living organisms. Some shade-loving plants (sciophytes) produce such low levels of oxygen during photosynthesis that they use all of it themselves instead of releasing it to
2375-413: A subgroup of the glycosidases, first catalyze the hydrolysis of long-chain glucose-containing polysaccharides, removing terminal glucose. In turn, disaccharides are mostly degraded by specific glycosidases to glucose. The names of the degrading enzymes are often derived from the particular poly- and disaccharide; inter alia, for the degradation of polysaccharide chains there are amylases (named after amylose,
2500-534: A subsequent sequence of light-independent reactions called the Calvin cycle . In this process, atmospheric carbon dioxide is incorporated into already existing organic compounds, such as ribulose bisphosphate (RuBP). Using the ATP and NADPH produced by the light-dependent reactions, the resulting compounds are then reduced and removed to form further carbohydrates, such as glucose . In other bacteria, different mechanisms like
2625-405: A type of non- carbon-fixing anoxygenic photosynthesis, where the simpler photopigment retinal and its microbial rhodopsin derivatives are used to absorb green light and power proton pumps to directly synthesize adenosine triphosphate (ATP), the "energy currency" of cells. Such archaeal photosynthesis might have been the earliest form of photosynthesis that evolved on Earth, as far back as
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#17330927737742750-591: A wide variety of colors. These pigments are embedded in plants and algae in complexes called antenna proteins. In such proteins, the pigments are arranged to work together. Such a combination of proteins is also called a light-harvesting complex . Although all cells in the green parts of a plant have chloroplasts, the majority of those are found in specially adapted structures called leaves . Certain species adapted to conditions of strong sunlight and aridity , such as many Euphorbia and cactus species, have their main photosynthetic organs in their stems. The cells in
2875-441: Is dextrorotatory , meaning it will rotate the direction of polarized light clockwise as seen looking toward the light source. The effect is due to the chirality of the molecules, and indeed the mirror-image isomer, l -(−)-glucose, is levorotatory (rotates polarized light counterclockwise) by the same amount. The strength of the effect is different for each of the five tautomers . The d - prefix does not refer directly to
3000-406: Is 198.17 g/mol, that for anhydrous D-glucose is 180.16 g/mol The density of these two forms of glucose is also different. In terms of chemical structure, glucose is a monosaccharide, that is, a simple sugar. Glucose contains six carbon atoms and an aldehyde group , and is therefore an aldohexose . The glucose molecule can exist in an open-chain (acyclic) as well as ring (cyclic) form—due to
3125-473: Is 31 °C (88 °F) and the Gordon–Taylor constant (an experimentally determined constant for the prediction of the glass transition temperature for different mass fractions of a mixture of two substances) is 4.5. A open-chain form of glucose makes up less than 0.02% of the glucose molecules in an aqueous solution at equilibrium. The rest is one of two cyclic hemiacetal forms. In its open-chain form,
3250-462: Is a chemical classifier denoting a sugar. Glucose was first isolated from raisins in 1747 by the German chemist Andreas Marggraf . Glucose was discovered in grapes by another German chemist – Johann Tobias Lowitz – in 1792, and distinguished as being different from cane sugar ( sucrose ). Glucose is the term coined by Jean Baptiste Dumas in 1838, which has prevailed in
3375-408: Is a glucose molecule with an additional water molecule attached. Its chemical formula is C 6 H 12 O 6 · H 2 O . Dextrose monohydrate is also called hydrated D-glucose , and commonly manufactured from plant starches. Dextrose monohydrate is utilized as the predominant type of dextrose in food applications, such as beverage mixes—it is a common form of glucose widely used as
3500-490: Is an endothermic redox reaction. In general outline, photosynthesis is the opposite of cellular respiration : while photosynthesis is a process of reduction of carbon dioxide to carbohydrates, cellular respiration is the oxidation of carbohydrates or other nutrients to carbon dioxide. Nutrients used in cellular respiration include carbohydrates, amino acids and fatty acids. These nutrients are oxidized to produce carbon dioxide and water, and to release chemical energy to drive
3625-508: Is an open-chain to a small extent and is present predominantly as α- or β- pyranose , which interconvert. From aqueous solutions, the three known forms can be crystallized: α-glucopyranose, β-glucopyranose and α-glucopyranose monohydrate. Glucose is a building block of the disaccharides lactose and sucrose (cane or beet sugar), of oligosaccharides such as raffinose and of polysaccharides such as starch , amylopectin , glycogen , and cellulose . The glass transition temperature of glucose
3750-498: Is both an evolutionary precursor to C 4 and a useful carbon-concentrating mechanism in its own right. Xerophytes , such as cacti and most succulents , also use PEP carboxylase to capture carbon dioxide in a process called Crassulacean acid metabolism (CAM). In contrast to C 4 metabolism, which spatially separates the CO 2 fixation to PEP from the Calvin cycle, CAM temporally separates these two processes. CAM plants have
3875-507: Is commonly commercially manufactured from starches , such as corn starch in the US and Japan, from potato and wheat starch in Europe, and from tapioca starch in tropical areas. The manufacturing process uses hydrolysis via pressurized steaming at controlled pH in a jet followed by further enzymatic depolymerization. Unbonded glucose is one of the main ingredients of honey . The term dextrose
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4000-474: Is commonly measured in μmols /( m / s ), parts per million, or volume per million; and H 2 O is commonly measured in mmols /(m /s) or in mbars . By measuring CO 2 assimilation , ΔH 2 O, leaf temperature, barometric pressure , leaf area, and photosynthetically active radiation (PAR), it becomes possible to estimate, "A" or carbon assimilation, "E" or transpiration , "gs" or stomatal conductance , and "Ci" or intracellular CO 2 . However, it
4125-646: Is composed of approximately 9.5% water by mass; through the process of dehydration, this water content is eliminated to yield anhydrous (dry) dextrose. Anhydrous dextrose has the chemical formula C 6 H 12 O 6 , without any water molecule attached which is the same as glucose. Anhydrous dextrose on open air tends to absorb moisture and transform to the monohydrate, and it is more expensive to produce. Anhydrous dextrose (anhydrous D-glucose) has increased stability and increased shelf life, has medical applications, such as in oral glucose tolerance test . Whereas molecular weight (molar mass) for D-glucose monohydrate
4250-430: Is converted to CO 2 by an oxalate oxidase enzyme, and the produced CO 2 can support the Calvin cycle reactions. Reactive hydrogen peroxide (H 2 O 2 ), the byproduct of oxalate oxidase reaction, can be neutralized by catalase . Alarm photosynthesis represents a photosynthetic variant to be added to the well-known C4 and CAM pathways. However, alarm photosynthesis, in contrast to these pathways, operates as
4375-419: Is freed from its locked position through a classic "hop". The movement of the electron towards the photo center is therefore covered in a series of conventional hops and quantum walks. Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old. More recent studies also suggest that photosynthesis may have begun about 3.4 billion years ago, though
4500-412: Is further excited by the light absorbed by that photosystem . The electron is then passed along a chain of electron acceptors to which it transfers some of its energy . The energy delivered to the electron acceptors is used to move hydrogen ions across the thylakoid membrane into the lumen . The electron is eventually used to reduce the coenzyme NADP with an H to NADPH (which has functions in
4625-451: Is more common to use chlorophyll fluorescence for plant stress measurement , where appropriate, because the most commonly used parameters FV/FM and Y(II) or F/FM' can be measured in a few seconds, allowing the investigation of larger plant populations. Gas exchange systems that offer control of CO 2 levels, above and below ambient , allow the common practice of measurement of A/Ci curves, at different CO 2 levels, to characterize
4750-594: Is naturally occurring and is found in its free state in fruits and other parts of plants. In animals, it is released from the breakdown of glycogen in a process known as glycogenolysis . Glucose, as intravenous sugar solution , is on the World Health Organization's List of Essential Medicines . It is also on the list in combination with sodium chloride (table salt). The name glucose is derived from Ancient Greek γλεῦκος ( gleûkos ) 'wine, must', from γλυκύς ( glykýs ) 'sweet'. The suffix -ose
4875-423: Is often used in a clinical (related to patient's health status) or nutritional context (related to dietary intake, such as food labels or dietary guidelines), while "glucose" is used in a biological or physiological context (chemical processes and molecular interactions), but both terms refer to the same molecule, specifically D-glucose. Dextrose monohydrate is the hydrated form of D-glucose, meaning that it
5000-406: Is passed through a food chain . The fixation or reduction of carbon dioxide is a process in which carbon dioxide combines with a five-carbon sugar , ribulose 1,5-bisphosphate , to yield two molecules of a three-carbon compound, glycerate 3-phosphate , also known as 3-phosphoglycerate. Glycerate 3-phosphate, in the presence of ATP and NADPH produced during the light-dependent stages,
5125-483: Is produced by conversion of food, but it is also synthesized from other metabolites in the body's cells. In humans, the breakdown of glucose-containing polysaccharides happens in part already during chewing by means of amylase , which is contained in saliva , as well as by maltase , lactase , and sucrase on the brush border of the small intestine . Glucose is a building block of many carbohydrates and can be split off from them using certain enzymes. Glucosidases ,
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5250-440: Is produced by most cell types and is of particular importance for nerve cells and pancreatic β-cells . GLUT3 is highly expressed in nerve cells. Glucose from the bloodstream is taken up by GLUT4 from muscle cells (of the skeletal muscle and heart muscle ) and fat cells . GLUT14 is expressed exclusively in testicles . Excess glucose is broken down and converted into fatty acids, which are stored as triglycerides . In
5375-623: Is reduced to glyceraldehyde 3-phosphate . This product is also referred to as 3-phosphoglyceraldehyde (PGAL) or, more generically, as triose phosphate. Most (five out of six molecules) of the glyceraldehyde 3-phosphate produced are used to regenerate ribulose 1,5-bisphosphate so the process can continue. The triose phosphates not thus "recycled" often condense to form hexose phosphates, which ultimately yield sucrose , starch , and cellulose , as well as glucose and fructose . The sugars produced during carbon metabolism yield carbon skeletons that can be used for other metabolic reactions like
5500-415: Is termed glycogenolysis, the cleavage of starch is called starch degradation. The metabolic pathway that begins with molecules containing two to four carbon atoms (C) and ends in the glucose molecule containing six carbon atoms is called gluconeogenesis and occurs in all living organisms. The smaller starting materials are the result of other metabolic pathways. Ultimately almost all biomolecules come from
5625-485: Is the phosphorylation of glucose by a hexokinase to form glucose 6-phosphate . The main reason for the immediate phosphorylation of glucose is to prevent its diffusion out of the cell as the charged phosphate group prevents glucose 6-phosphate from easily crossing the cell membrane . Furthermore, addition of the high-energy phosphate group activates glucose for subsequent breakdown in later steps of glycolysis. In anaerobic respiration, one glucose molecule produces
5750-467: Is the human body's key source of energy, through aerobic respiration, providing about 3.75 kilocalories (16 kilojoules ) of food energy per gram. Breakdown of carbohydrates (e.g., starch) yields mono- and disaccharides , most of which is glucose. Through glycolysis and later in the reactions of the citric acid cycle and oxidative phosphorylation , glucose is oxidized to eventually form carbon dioxide and water, yielding energy mostly in
5875-429: Is then translocated to specialized bundle sheath cells where the enzyme RuBisCO and other Calvin cycle enzymes are located, and where CO 2 released by decarboxylation of the four-carbon acids is then fixed by RuBisCO activity to the three-carbon 3-phosphoglyceric acids . The physical separation of RuBisCO from the oxygen-generating light reactions reduces photorespiration and increases CO 2 fixation and, thus,
6000-404: Is then converted into the final carbohydrate products. The simple carbon sugars photosynthesis produces are then used to form other organic compounds , such as the building material cellulose , the precursors for lipid and amino acid biosynthesis, or as a fuel in cellular respiration . The latter occurs not only in plants but also in animals when the carbon and energy from plants
6125-499: Is used by 16,000 species of plants. Calcium-oxalate -accumulating plants, such as Amaranthus hybridus and Colobanthus quitensis , show a variation of photosynthesis where calcium oxalate crystals function as dynamic carbon pools , supplying carbon dioxide (CO 2 ) to photosynthetic cells when stomata are partially or totally closed. This process was named alarm photosynthesis . Under stress conditions (e.g., water deficit ), oxalate released from calcium oxalate crystals
6250-457: Is used by plants to make cellulose —the most abundant carbohydrate in the world—for use in cell walls , and by all living organisms to make adenosine triphosphate (ATP), which is used by the cell as energy. In energy metabolism , glucose is the most important source of energy in all organisms . Glucose for metabolism is stored as a polymer , in plants mainly as amylose and amylopectin , and in animals as glycogen . Glucose circulates in
6375-474: Is vital for climate processes, as it captures carbon dioxide from the air and binds it into plants, harvested produce and soil. Cereals alone are estimated to bind 3,825 Tg or 3.825 Pg of carbon dioxide every year, i.e. 3.825 billion metric tons. Most photosynthetic organisms are photoautotrophs , which means that they are able to synthesize food directly from carbon dioxide and water using energy from light. However, not all organisms use carbon dioxide as
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#17330927737746500-476: The Crabtree effect . Glucose can also degrade to form carbon dioxide through abiotic means. This has been demonstrated to occur experimentally via oxidation and hydrolysis at 22 °C and a pH of 2.5. Glucose is a ubiquitous fuel in biology . It is used as an energy source in organisms, from bacteria to humans, through either aerobic respiration , anaerobic respiration (in bacteria), or fermentation . Glucose
6625-575: The Haworth projection , the designation "α-" means that the hydroxyl group attached to C-1 and the −CH 2 OH group at C-5 lies on opposite sides of the ring's plane (a trans arrangement), while "β-" means that they are on the same side of the plane (a cis arrangement). Therefore, the open-chain isomer D -glucose gives rise to four distinct cyclic isomers: α- D -glucopyranose, β- D -glucopyranose, α- D -glucofuranose, and β- D -glucofuranose. These five structures exist in equilibrium and interconvert, and
6750-572: The Paleoarchean , preceding that of cyanobacteria (see Purple Earth hypothesis ). While the details may differ between species , the process always begins when light energy is absorbed by the reaction centers , proteins that contain photosynthetic pigments or chromophores . In plants, these proteins are chlorophylls (a porphyrin derivative that absorbs the red and blue spectrums of light, thus reflecting green) held inside chloroplasts , abundant in leaf cells. In bacteria, they are embedded in
6875-439: The basolateral side of the intestinal epithelial cells via the glucose transporter GLUT2 , as well uptake into liver cells , kidney cells, cells of the islets of Langerhans , neurons , astrocytes , and tanycytes . Glucose enters the liver via the portal vein and is stored there as a cellular glycogen. In the liver cell, it is phosphorylated by glucokinase at position 6 to form glucose 6-phosphate , which cannot leave
7000-408: 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 (compounds containing carbon) like sugars, glycogen , cellulose and starches . To use this stored chemical energy, an organism's cells metabolize
7125-586: The equatorial position in the form of β- d -glucose, is more readily accessible to chemical reactions, for example, for esterification or acetal formation. For this reason, d -glucose is also a highly preferred building block in natural polysaccharides (glycans). Polysaccharides that are composed solely of glucose are termed glucans . Glucose is produced by plants through photosynthesis using sunlight, water and carbon dioxide and can be used by all living organisms as an energy and carbon source. However, most glucose does not occur in its free form, but in
7250-481: The kidneys , glucose in the urine is absorbed via SGLT1 and SGLT2 in the apical cell membranes and transmitted via GLUT2 in the basolateral cell membranes. About 90% of kidney glucose reabsorption is via SGLT2 and about 3% via SGLT1. In plants and some prokaryotes , glucose is a product of photosynthesis . Glucose is also formed by the breakdown of polymeric forms of glucose like glycogen (in animals and mushrooms ) or starch (in plants). The cleavage of glycogen
7375-637: The light reaction of photosynthesis by using chlorophyll fluorometers . Actual plants' photosynthetic efficiency varies with the frequency of the light being converted, light intensity , temperature , and proportion of carbon dioxide in the atmosphere , and can vary from 0.1% to 8%. By comparison, solar panels convert light into electric energy at an efficiency of approximately 6–20% for mass-produced panels, and above 40% in laboratory devices. Scientists are studying photosynthesis in hopes of developing plants with increased yield . The efficiency of both light and dark reactions can be measured, but
7500-434: The light-independent (or "dark") reactions, the enzyme RuBisCO captures CO 2 from the atmosphere and, in a process called the Calvin cycle , uses the newly formed NADPH and releases three-carbon sugars , which are later combined to form sucrose and starch . The overall equation for the light-independent reactions in green plants is Carbon fixation produces the three-carbon sugar intermediate , which
7625-417: The palisade mesophyll cells where most of the photosynthesis takes place. In the light-dependent reactions , one molecule of the pigment chlorophyll absorbs one photon and loses one electron . This electron is taken up by a modified form of chlorophyll called pheophytin , which passes the electron to a quinone molecule, starting the flow of electrons down an electron transport chain that leads to
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#17330927737747750-417: The photosynthetic capacity of the leaf . C 4 plants can produce more sugar than C 3 plants in conditions of high light and temperature . Many important crop plants are C 4 plants, including maize , sorghum , sugarcane , and millet . Plants that do not use PEP-carboxylase in carbon fixation are called C 3 plants because the primary carboxylation reaction , catalyzed by RuBisCO, produces
7875-462: The photosystems , quantum efficiency and the CO 2 assimilation rates. With some instruments, even wavelength dependency of the photosynthetic efficiency can be analyzed . A phenomenon known as quantum walk increases the efficiency of the energy transport of light significantly. In the photosynthetic cell of an alga , bacterium , or plant, there are light-sensitive molecules called chromophores arranged in an antenna-shaped structure called
8000-437: The plasma membrane . In these light-dependent reactions, some energy is used to strip electrons from suitable substances, such as water, producing oxygen gas. The hydrogen freed by the splitting of water is used in the creation of two important molecules that participate in energetic processes: reduced nicotinamide adenine dinucleotide phosphate (NADPH) and ATP. In plants, algae, and cyanobacteria, sugars are synthesized by
8125-450: The reverse Krebs cycle are used to achieve the same end. The first photosynthetic organisms probably evolved early in the evolutionary history of life using reducing agents such as hydrogen or hydrogen sulfide, rather than water, as sources of electrons. Cyanobacteria appeared later; the excess oxygen they produced contributed directly to the oxygenation of the Earth , which rendered
8250-411: The "envelope" conformations of cyclopentane . In the solid state, only the glucopyranose forms are observed. Some derivatives of glucofuranose, such as 1,2- O -isopropylidene- D -glucofuranose are stable and can be obtained pure as crystalline solids. For example, reaction of α-D-glucose with para -tolylboronic acid H 3 C−(C 6 H 4 )−B(OH) 2 reforms the normal pyranose ring to yield
8375-421: The 4-fold ester α-D-glucofuranose-1,2:3,5-bis( p -tolylboronate). Mutarotation consists of a temporary reversal of the ring-forming reaction, resulting in the open-chain form, followed by a reforming of the ring. The ring closure step may use a different −OH group than the one recreated by the opening step (thus switching between pyranose and furanose forms), or the new hemiacetal group created on C-1 may have
8500-433: The CO 2 concentration in the leaves under these conditions. Plants that use the C 4 carbon fixation process chemically fix carbon dioxide in the cells of the mesophyll by adding it to the three-carbon molecule phosphoenolpyruvate (PEP), a reaction catalyzed by an enzyme called PEP carboxylase , creating the four-carbon organic acid oxaloacetic acid . Oxaloacetic acid or malate synthesized by this process
8625-419: The action spectrum is blue-green light, which allows these algae to use the blue end of the spectrum to grow in the deeper waters that filter out the longer wavelengths (red light) used by above-ground green plants. The non-absorbed part of the light spectrum is what gives photosynthetic organisms their color (e.g., green plants, red algae, purple bacteria ) and is the least effective for photosynthesis in
8750-447: The aldehyde group (at C-1) and either the C-4 or C-5 hydroxyl group, forming a hemiacetal linkage, −C(OH)H−O− . The reaction between C-1 and C-5 yields a six-membered heterocyclic system called a pyranose, which is a monosaccharide sugar (hence "-ose") containing a derivatised pyran skeleton. The (much rarer) reaction between C-1 and C-4 yields a five-membered furanose ring, named after
8875-402: The ambient environment. Glucose concentrations in the atmosphere are detected via collection of samples by aircraft and are known to vary from location to location. For example, glucose concentrations in atmospheric air from inland China range from 0.8 to 20.1 pg/L, whereas east coastal China glucose concentrations range from 10.3 to 142 pg/L. In humans, glucose is metabolized by glycolysis and
9000-536: The arrangements of chemical bonds in carbon-bearing molecules. Between 1891 and 1894, Fischer established the stereochemical configuration of all the known sugars and correctly predicted the possible isomers , applying Van 't Hoff equation of asymmetrical carbon atoms. The names initially referred to the natural substances. Their enantiomers were given the same name with the introduction of systematic nomenclatures, taking into account absolute stereochemistry (e.g. Fischer nomenclature , d / l nomenclature). For
9125-435: The assimilation of carbon dioxide in plants and microbes during photosynthesis. The free energy of formation of α- d -glucose is 917.2 kilojoules per mole. In humans, gluconeogenesis occurs in the liver and kidney, but also in other cell types. In the liver about 150 g (5.3 oz) of glycogen are stored, in skeletal muscle about 250 g (8.8 oz). However, the glucose released in muscle cells upon cleavage of
9250-521: The atmosphere. Although there are some differences between oxygenic photosynthesis in plants , algae , and cyanobacteria , the overall process is quite similar in these organisms. There are also many varieties of anoxygenic photosynthesis , used mostly by bacteria, which consume carbon dioxide but do not release oxygen. Carbon dioxide is converted into sugars in a process called carbon fixation ; photosynthesis captures energy from sunlight to convert carbon dioxide into carbohydrates . Carbon fixation
9375-528: The bacteria in the gut microbiota do. In order to get into or out of cell membranes of cells and membranes of cell compartments, glucose requires special transport proteins from the major facilitator superfamily . In the small intestine (more precisely, in the jejunum ), glucose is taken up into the intestinal epithelium with the help of glucose transporters via a secondary active transport mechanism called sodium ion-glucose symport via sodium/glucose cotransporter 1 (SGLT1). Further transfer occurs on
9500-431: The blood of animals as blood sugar . The naturally occurring form is d -glucose, while its stereoisomer l -glucose is produced synthetically in comparatively small amounts and is less biologically active. Glucose is a monosaccharide containing six carbon atoms and an aldehyde group, and is therefore an aldohexose . The glucose molecule can exist in an open-chain (acyclic) as well as ring (cyclic) form. Glucose
9625-480: The carboxysome quickly sponges it up. HCO 3 ions are made from CO 2 outside the cell by another carbonic anhydrase and are actively pumped into the cell by a membrane protein. They cannot cross the membrane as they are charged, and within the cytosol they turn back into CO 2 very slowly without the help of carbonic anhydrase. This causes the HCO 3 ions to accumulate within the cell from where they diffuse into
9750-491: The carboxysomes. Pyrenoids in algae and hornworts also act to concentrate CO 2 around RuBisCO. The overall process of photosynthesis takes place in four stages: Plants usually convert light into chemical energy with a photosynthetic efficiency of 3–6%. Absorbed light that is unconverted is dissipated primarily as heat , with a small fraction (1–2%) reemitted as chlorophyll fluorescence at longer (redder) wavelengths . This fact allows measurement of
9875-428: The cell. Glucose 6-phosphatase can convert glucose 6-phosphate back into glucose exclusively in the liver, so the body can maintain a sufficient blood glucose concentration. In other cells, uptake happens by passive transport through one of the 14 GLUT proteins. In the other cell types, phosphorylation occurs through a hexokinase , whereupon glucose can no longer diffuse out of the cell. The glucose transporter GLUT1
10000-465: The chemical literature. Friedrich August Kekulé proposed the term dextrose (from the Latin dexter , meaning "right"), because in aqueous solution of glucose, the plane of linearly polarized light is turned to the right. In contrast, l-fructose (usually referred to as d -fructose) (a ketohexose) and l-glucose ( l -glucose) turn linearly polarized light to the left. The earlier notation according to
10125-409: The conditions of non-cyclic electron flow in green plants is: Not all wavelengths of light can support photosynthesis. The photosynthetic action spectrum depends on the type of accessory pigments present. For example, in green plants , the action spectrum resembles the absorption spectrum for chlorophylls and carotenoids with absorption peaks in violet-blue and red light. In red algae ,
10250-401: The cyclic ether furan . In either case, each carbon in the ring has one hydrogen and one hydroxyl attached, except for the last carbon (C-4 or C-5) where the hydroxyl is replaced by the remainder of the open molecule (which is −(C(CH 2 OH)HOH)−H or −(CHOH)−H respectively). The ring-closing reaction can give two products, denoted "α-" and "β-". When a glucopyranose molecule is drawn in
10375-503: The diagram), a chemiosmotic potential is generated by pumping proton cations ( H ) across the membrane and into the thylakoid space . An ATP synthase enzyme uses that chemiosmotic potential to make ATP during photophosphorylation , whereas NADPH is a product of the terminal redox reaction in the Z-scheme . The electron enters a chlorophyll molecule in Photosystem I . There it
10500-567: The discovery of the metabolism of glucose Otto Meyerhof received the Nobel Prize in Physiology or Medicine in 1922. Hans von Euler-Chelpin was awarded the Nobel Prize in Chemistry along with Arthur Harden in 1929 for their "research on the fermentation of sugar and their share of enzymes in this process". In 1947, Bernardo Houssay (for his discovery of the role of the pituitary gland in
10625-505: The equation for this process is: This equation emphasizes that water is both a reactant in the light-dependent reaction and a product of the light-independent reaction , but canceling n water molecules from each side gives the net equation: Other processes substitute other compounds (such as arsenite ) for water in the electron-supply role; for example some microbes use sunlight to oxidize arsenite to arsenate : The equation for this reaction is: Photosynthesis occurs in two stages. In
10750-518: The evolution of complex life possible. The average rate of energy captured by global photosynthesis is approximately 130 terawatts , which is about eight times the total power consumption of human civilization . Photosynthetic organisms also convert around 100–115 billion tons (91–104 Pg petagrams , or billions of metric tons), of carbon into biomass per year. Photosynthesis was discovered in 1779 by Jan Ingenhousz . He showed that plants need light, not just air, soil, and water. Photosynthesis
10875-443: The first direct evidence of photosynthesis comes from thylakoid membranes preserved in 1.75-billion-year-old cherts . Glucose Glucose is a sugar with the molecular formula C 6 H 12 O 6 . It is overall the most abundant monosaccharide , a subcategory of carbohydrates . It is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using energy from sunlight. It
11000-582: The first stage, light-dependent reactions or light reactions capture the energy of light and use it to make the hydrogen carrier NADPH and the energy-storage molecule ATP . During the second stage, the light-independent reactions use these products to capture and reduce carbon dioxide. Most organisms that use oxygenic photosynthesis use visible light for the light-dependent reactions, although at least three use shortwave infrared or, more specifically, far-red radiation. Some organisms employ even more radical variants of photosynthesis. Some archaea use
11125-441: The first step of the Z-scheme , requires an external source of electrons to reduce its oxidized chlorophyll a reaction center. The source of electrons for photosynthesis in green plants and cyanobacteria is water. Two water molecules are oxidized by the energy of four successive charge-separation reactions of photosystem II to yield a molecule of diatomic oxygen and four hydrogen ions. The electrons yielded are transferred to
11250-597: The form of adenosine triphosphate (ATP). The insulin reaction, and other mechanisms, regulate the concentration of glucose in the blood. The physiological caloric value of glucose, depending on the source, is 16.2 kilojoules per gram or 15.7 kJ/g (3.74 kcal/g). The high availability of carbohydrates from plant biomass has led to a variety of methods during evolution, especially in microorganisms, to utilize glucose for energy and carbon storage. Differences exist in which end product can no longer be used for energy production. The presence of individual genes, and their gene products,
11375-406: The form of its polymers, i.e. lactose, sucrose, starch and others which are energy reserve substances, and cellulose and chitin , which are components of the cell wall in plants or fungi and arthropods , respectively. These polymers, when consumed by animals, fungi and bacteria, are degraded to glucose using enzymes. All animals are also able to produce glucose themselves from certain precursors as
11500-455: The formation of lactate, the end product of fermentation in mammals, even in the presence of oxygen. This is called the Warburg effect . For the increased uptake of glucose in tumors various SGLT and GLUT are overly produced. In yeast , ethanol is fermented at high glucose concentrations, even in the presence of oxygen (which normally leads to respiration rather than fermentation). This is called
11625-416: The four cyclic isomers interconvert over a time scale of hours, in a process called mutarotation . Starting from any proportions, the mixture converges to a stable ratio of α:β 36:64. The ratio would be α:β 11:89 if it were not for the influence of the anomeric effect . Mutarotation is considerably slower at temperatures close to 0 °C (32 °F). Whether in water or the solid form, d -(+)-glucose
11750-722: The glucose degradation in animals occurs anaerobic to lactate via lactic acid fermentation and releases much less energy. Muscular lactate enters the liver through the bloodstream in mammals, where gluconeogenesis occurs ( Cori cycle ). With a high supply of glucose, the metabolite acetyl-CoA from the Krebs cycle can also be used for fatty acid synthesis . Glucose is also used to replenish the body's glycogen stores, which are mainly found in liver and skeletal muscle. These processes are hormonally regulated. In other living organisms, other forms of fermentation can occur. The bacterium Escherichia coli can grow on nutrient media containing glucose as
11875-521: The glucose molecule has an open (as opposed to cyclic ) unbranched backbone of six carbon atoms, where C-1 is part of an aldehyde group H(C=O)− . Therefore, glucose is also classified as an aldose , or an aldohexose . The aldehyde group makes glucose a reducing sugar giving a positive reaction with the Fehling test . In solutions, the open-chain form of glucose (either " D -" or " L -") exists in equilibrium with several cyclic isomers , each containing
12000-453: The glycation of proteins or lipids . In contrast, enzyme -regulated addition of sugars to protein is called glycosylation and is essential for the function of many proteins. Ingested glucose initially binds to the receptor for sweet taste on the tongue in humans. This complex of the proteins T1R2 and T1R3 makes it possible to identify glucose-containing food sources. Glucose mainly comes from food—about 300 g (11 oz) per day
12125-519: The glycogen can not be delivered to the circulation because glucose is phosphorylated by the hexokinase, and a glucose-6-phosphatase is not expressed to remove the phosphate group. Unlike for glucose, there is no transport protein for glucose-6-phosphate . Gluconeogenesis allows the organism to build up glucose from other metabolites, including lactate or certain amino acids , while consuming energy. The renal tubular cells can also produce glucose. Glucose also can be found outside of living organisms in
12250-440: The interconversion is much more rapid with acid catalysis . The other open-chain isomer L -glucose similarly gives rise to four distinct cyclic forms of L -glucose, each the mirror image of the corresponding D -glucose. The glucopyranose ring (α or β) can assume several non-planar shapes, analogous to the "chair" and "boat" conformations of cyclohexane . Similarly, the glucofuranose ring may assume several shapes, analogous to
12375-399: The interior of a cell, giving the membrane a very large surface area and therefore increasing the amount of light that the bacteria can absorb. In plants and algae, photosynthesis takes place in organelles called chloroplasts . A typical plant cell contains about 10 to 100 chloroplasts. The chloroplast is enclosed by a membrane. This membrane is composed of a phospholipid inner membrane,
12500-433: The interior tissues of a leaf, called the mesophyll , can contain between 450,000 and 800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is coated with a water-resistant waxy cuticle that protects the leaf from excessive evaporation of water and decreases the absorption of ultraviolet or blue light to minimize heating . The transparent epidermis layer allows light to pass through to
12625-434: The light reaction, and infrared gas analyzers can measure the dark reaction . An integrated chlorophyll fluorometer and gas exchange system can investigate both light and dark reactions when researchers use the two separate systems together. Infrared gas analyzers and some moisture sensors are sensitive enough to measure the photosynthetic assimilation of CO 2 and of Δ H 2 O using reliable methods . CO 2
12750-433: The light-independent reaction); at that point, the path of that electron ends. The cyclic reaction is similar to that of the non-cyclic but differs in that it generates only ATP, and no reduced NADP (NADPH) is created. The cyclic reaction takes place only at photosystem I. Once the electron is displaced from the photosystem, the electron is passed down the electron acceptor molecules and returns to photosystem I, from where it
12875-710: The metabolism of glucose and the derived carbohydrates) as well as Carl and Gerty Cori (for their discovery of the conversion of glycogen from glucose) received the Nobel Prize in Physiology or Medicine. In 1970, Luis Leloir was awarded the Nobel Prize in Chemistry for the discovery of glucose-derived sugar nucleotides in the biosynthesis of carbohydrates. Glucose forms white or colorless solids that are highly soluble in water and acetic acid but poorly soluble in methanol and ethanol . They melt at 146 °C (295 °F) ( α ) and 150 °C (302 °F) ( beta ), decompose starting at 188 °C (370 °F) with release of various volatile products, ultimately leaving
13000-415: The monosaccharides mannose , glucose and fructose interconvert (via a Lobry de Bruyn–Alberda–Van Ekenstein transformation ), so that a balance between these isomers is formed. This reaction proceeds via an enediol : [REDACTED] Glucose is the most abundant monosaccharide. Glucose is also the most widely used aldohexose in most living organisms. One possible explanation for this is that glucose has
13125-432: The more common types of photosynthesis. In photosynthetic bacteria, the proteins that gather light for photosynthesis are embedded in cell membranes . In its simplest form, this involves the membrane surrounding the cell itself. However, the membrane may be tightly folded into cylindrical sheets called thylakoids , or bunched up into round vesicles called intracytoplasmic membranes . These structures can fill most of
13250-439: The most efficient route, where it will have the highest probability of arriving at its destination in the minimum possible time. Because that quantum walking takes place at temperatures far higher than quantum phenomena usually occur, it is only possible over very short distances. Obstacles in the form of destructive interference cause the particle to lose its wave properties for an instant before it regains them once again after it
13375-441: The most stable cyclic form of all the aldohexoses is that its hydroxy groups (with the exception of the hydroxy group on the anomeric carbon of d -glucose) are in the equatorial position . Presumably, glucose is the most abundant natural monosaccharide because it is less glycated with proteins than other monosaccharides. Another hypothesis is that glucose, being the only d -aldohexose that has all five hydroxy substituents in
13500-506: The need arises. Neurons , cells of the renal medulla and erythrocytes depend on glucose for their energy production. In adult humans, there is about 18 g (0.63 oz) of glucose, of which about 4 g (0.14 oz) is present in the blood. Approximately 180–220 g (6.3–7.8 oz) of glucose is produced in the liver of an adult in 24 hours. Many of the long-term complications of diabetes (e.g., blindness , kidney failure , and peripheral neuropathy ) are probably due to
13625-463: The optical properties of the compound. It indicates that the C-5 chiral centre has the same handedness as that of d -glyceraldehyde (which was so labelled because it is dextrorotatory). The fact that d -glucose is dextrorotatory is a combined effect of its four chiral centres, not just of C-5; some of the other d -aldohexoses are levorotatory. The conversion between the two anomers can be observed in
13750-498: 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. Some bacteria also perform anoxygenic photosynthesis , which uses bacteriochlorophyll to split hydrogen sulfide as a reductant instead of water, producing sulfur instead of oxygen. Archaea such as Halobacterium also perform
13875-410: The organism's metabolism . Photosynthesis and cellular respiration are distinct processes, as they take place through different sequences of chemical reactions and in different cellular compartments (cellular respiration in mitochondria ). The general equation for photosynthesis as first proposed by Cornelis van Niel is: Since water is used as the electron donor in oxygenic photosynthesis,
14000-434: The pentose phosphate pathway. Glycolysis is used by all living organisms, with small variations, and all organisms generate energy from the breakdown of monosaccharides. In the further course of the metabolism, it can be completely degraded via oxidative decarboxylation , the citric acid cycle (synonym Krebs cycle ) and the respiratory chain to water and carbon dioxide. If there is not enough oxygen available for this,
14125-556: The photosynthetic system. Plants absorb light primarily using the pigment chlorophyll . The green part of the light spectrum is not absorbed but is reflected, which is the reason that most plants have a green color. Besides chlorophyll, plants also use pigments such as carotenes and xanthophylls . Algae also use chlorophyll, but various other pigments are present, such as phycocyanin , carotenes , and xanthophylls in green algae , phycoerythrin in red algae (rhodophytes) and fucoxanthin in brown algae and diatoms resulting in
14250-412: The presence of alcohol and aldehyde or ketone functional groups, the form having the straight chain can easily convert into a chair-like hemiacetal ring structure commonly found in carbohydrates. Glucose is present in solid form as a monohydrate with a closed pyran ring (α-glucopyranose monohydrate, sometimes known less precisely by dextrose hydrate). In aqueous solution, on the other hand, it
14375-476: The production of amino acids and lipids . In hot and dry conditions , plants close their stomata to prevent water loss. Under these conditions, CO 2 will decrease and oxygen gas , produced by the light reactions of photosynthesis, will increase, causing an increase of photorespiration by the oxygenase activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and decrease in carbon fixation. Some plants have evolved mechanisms to increase
14500-486: The relationship between the two can be complex. For example, the light reaction creates ATP and NADPH energy molecules , which C 3 plants can use for carbon fixation or photorespiration . Electrons may also flow to other electron sinks. For this reason, it is not uncommon for authors to differentiate between work done under non-photorespiratory conditions and under photorespiratory conditions . Chlorophyll fluorescence of photosystem II can measure
14625-462: The respective organisms . In plants , light-dependent reactions occur in the thylakoid membranes of the chloroplasts where they drive the synthesis of ATP and NADPH . The light-dependent reactions are of two forms: cyclic and non-cyclic . In the non-cyclic reaction, the photons are captured in the light-harvesting antenna complexes of photosystem II by chlorophyll and other accessory pigments (see diagram at right). The absorption of
14750-468: The rotation of the plane of linearly polarized light ( d and l -nomenclature) was later abandoned in favor of the d - and l -notation , which refers to the absolute configuration of the asymmetric center farthest from the carbonyl group, and in concordance with the configuration of d - or l -glyceraldehyde. Since glucose is a basic necessity of many organisms, a correct understanding of its chemical makeup and structure contributed greatly to
14875-510: The same or opposite handedness as the original one (thus switching between the α and β forms). Thus, though the open-chain form is barely detectable in solution, it is an essential component of the equilibrium. The open-chain form is thermodynamically unstable , and it spontaneously isomerizes to the cyclic forms. (Although the ring closure reaction could in theory create four- or three-atom rings, these would be highly strained, and are not observed in practice.) In solutions at room temperature ,
15000-479: The site of carboxylation in the chloroplast, to replace Ci. CO 2 concentration in the chloroplast becomes possible to estimate with the measurement of mesophyll conductance or g m using an integrated system. Photosynthesis measurement systems are not designed to directly measure the amount of light the leaf absorbs, but analysis of chlorophyll fluorescence , P700 - and P515-absorbance, and gas exchange measurements reveal detailed information about, e.g.,
15125-491: The sole carbon source. In some bacteria and, in modified form, also in archaea, glucose is degraded via the Entner-Doudoroff pathway . With Glucose, a mechanism for gene regulation was discovered in E. coli , the catabolite repression (formerly known as glucose effect ). Use of glucose as an energy source in cells is by either aerobic respiration, anaerobic respiration, or fermentation. The first step of glycolysis
15250-461: The three-carbon 3-phosphoglyceric acids directly in the Calvin-Benson cycle . Over 90% of plants use C 3 carbon fixation, compared to 3% that use C 4 carbon fixation; however, the evolution of C 4 in over sixty plant lineages makes it a striking example of convergent evolution . C 2 photosynthesis , which involves carbon-concentration by selective breakdown of photorespiratory glycine,
15375-424: The ultimate reduction of NADP to NADPH . In addition, this creates a proton gradient (energy gradient) across the chloroplast membrane , which is used by ATP synthase in the synthesis of ATP . The chlorophyll molecule ultimately regains the electron it lost when a water molecule is split in a process called photolysis , which releases oxygen . The overall equation for the light-dependent reactions under
15500-408: The water-oxidizing reaction (Kok's S-state diagrams). The hydrogen ions are released in the thylakoid lumen and therefore contribute to the transmembrane chemiosmotic potential that leads to ATP synthesis . Oxygen is a waste product of light-dependent reactions, but the majority of organisms on Earth use oxygen and its energy for cellular respiration , including photosynthetic organisms . In
15625-462: Was emitted, hence the name cyclic reaction . Linear electron transport through a photosystem will leave the reaction center of that photosystem oxidized . Elevating another electron will first require re-reduction of the reaction center. The excited electrons lost from the reaction center ( P700 ) of photosystem I are replaced by transfer from plastocyanin , whose electrons come from electron transport through photosystem II . Photosystem II, as
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