photosynthesis photosynthesis photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. some of this chemical energy is stored in carbohydrate molecules, such as sugars and starches, which are synthesized from carbon dioxide and water – hence the name photosynthesis, from the greek phōs (φῶς), "light", and synthesis (σύνθεσις), "putting together". most plants, algae, and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. photosynthesis is largely responsible for producing and maintaining the oxygen content of the earth's atmosphere, and supplies most of the energy necessary for life on earth. schematic of photosynthesis in plants. the carbohydrates produced are stored in or used by the plant. composite image showing the global distribution of photosynthesis, including both oceanic phytoplankton and terrestrial vegetation. dark red and blue-green indicate regions of high photosynthetic activity in the ocean and on land, respectively. although photosynthesis is performed differently by different species, the process always begins when energy from light is absorbed by proteins called reaction centers that contain green chlorophyll (and other colored) pigments/chromophores. in plants, these proteins are held inside organelles called chloroplasts, which are most abundant in leaf cells, while in bacteria they are embedded in 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 further compounds that serve as short-term stores of energy, enabling its transfer to drive other reactions: these compounds are reduced nicotinamide adenine dinucleotide phosphate (nadph) and adenosine triphosphate (atp), the "energy currency" of cells. in plants, algae and cyanobacteria, sugars are synthesized by a subsequent sequence of light-independent reactions called the calvin cycle. in the calvin cycle, atmospheric carbon dioxide is incorporated into already existing organic carbon 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 such as the reverse krebs cycle are used to achieve the same end. the first photosynthetic organisms probably evolved early in the evolutionary history of life and most likely used 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 the evolution of complex life possible. today, the average rate of energy capture by photosynthesis globally is approximately 130 terawatts, which is about eight times the current power consumption of human civilization. photosynthetic organisms also convert around 100–115 billion tons (91–104 pg petagrams, or billion metric tons), of carbon into biomass per year. that plants receive some energy from light – in addition to air, soil, and water – was first discovered in 1779 by jan ingenhousz. photosynthesis is vital for climate processes, as it captures carbon dioxide from the air and then binds carbon in plants and further in soils and harvested products. cereals alone are estimated to bind 3,825 tg (teragrams) or 3.825 pg (petagrams) of carbon dioxide every year, i.e. 3.825 billion metric tons. overview photosynthesis changes sunlight into chemical energy, splits water to liberate o2, and fixes co2 into sugar. 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 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. 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 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 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. the general equation for photosynthesis as first proposed by cornelis van niel is: co2carbondioxide + 2h2aelectron donor + photonslight energy → carbohydrate + 2aoxidizedelectrondonor + h2owater since water is used as the electron donor in oxygenic photosynthesis, the equation for this process is: co2carbondioxide + 2h2owater + photonslight energy → carbohydrate + o2oxygen + h2owater 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: co2carbondioxide + h2o water + photonslight energy → carbohydrate + o2 oxygen 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: co2carbondioxide + (aso3−3)arsenite + photonslight energy → (aso3−4)arsenate + cocarbonmonoxide(used to build other compounds in subsequent reactions) photosynthesis occurs in two stages. in 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 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 the more common types of photosynthesis. photosynthetic membranes and organelles chloroplast ultrastructure:outer membraneintermembrane spaceinner membrane (1+2+3: envelope)stroma (aqueous fluid)thylakoid lumen (inside of thylakoid)thylakoid membranegranum (stack of thylakoids)thylakoid (lamella)starchribosomeplastidial dnaplastoglobule (drop of lipids) 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 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, 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 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 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 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 the palisade mesophyll cells where most of the photosynthesis takes place. light-dependent reactions light-dependent reactions of photosynthesis at the thylakoid membrane 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 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 the conditions of non-cyclic electron flow in green plants is: 2 h2o + 2 nadp+ + 3 adp + 3 pi + light → 2 nadph + 2 h+ + 3 atp + o2 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, 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 the respective organisms. z scheme the "z scheme" 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 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 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 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