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[Unsubscribe]( Efficiency Main article: Photosynthetic efficiency Plants usually convert light into chemical energy with a photosynthetic efficiency of 3–6%.[38][39] Absorbed light that is unconverted is dissipated primarily as heat, with a small fraction (1–2%)[40] re-emitted as chlorophyll fluorescence at longer (redder) wavelengths. This fact allows measurement of the light reaction of photosynthesis by using chlorophyll fluorometers.[40] 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%.[41] 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.[39] The efficiency of both light and dark reactions can be measured but the relationship between the two can be complex.[42] For example, the ATP and NADPH energy molecules, created by the light reaction, can be used for carbon fixation or for photorespiration in C3 plants.[42] Electrons may also flow to other electron sinks.[43][44][45] For this reason, it is not uncommon for authors to differentiate between work done under non-photorespiratory conditions and under photorespiratory conditions.[46][47][48] Chlorophyll fluorescence of photosystem II can measure the light reaction, and infrared gas analyzers can measure the dark reaction.[49] It is also possible to investigate both at the same time using an integrated chlorophyll fluorometer and gas exchange system, or by using two separate systems together.[50] Infrared gas analyzers and some moisture sensors are sensitive enough to measure the photosynthetic assimilation of CO2, and of ΔH2O using reliable methods[51] CO2 is commonly measured in μmols/(m2/s), parts per million or volume per million and H2O is commonly measured in mmol/(m2/s) or in mbars.[51] By measuring CO2 assimilation, ΔH2O, leaf temperature, barometric pressure, leaf area, and photosynthetically active radiation or PAR, it becomes possible to estimate, "A" or carbon assimilation, "E" or transpiration, "gs" or stomatal conductance, and Ci or intracellular CO2.[51] However, it is more common to used 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.[48] Gas exchange systems that offer control of CO2 levels, above and below ambient, allow the common practice of measurement of A/Ci curves, at different CO2 levels, to characterize a plant's photosynthetic response.[51] Integrated chlorophyll fluorometer – gas exchange systems allow a more precise measure of photosynthetic response and mechanisms.[49][50] While standard gas exchange photosynthesis systems can measure Ci, or substomatal CO2 levels, the addition of integrated chlorophyll fluorescence measurements allows a more precise measurement of CC to replace Ci.[50][52] The estimation of CO2 at the site of carboxylation in the chloroplast, or CC, becomes possible with the measurement of mesophyll conductance or gm using an integrated system.[49][50][53] Photosynthesis measurement systems are not designed to directly measure the amount of light absorbed by the leaf. But analysis of chlorophyll-fluorescence, P700- and P515-absorbance and gas exchange measurements reveal detailed information about e.g. the photosystems, quantum efficiency and the CO2 assimilation rates. With some instruments, even wavelength-dependency of the photosynthetic efficiency can be analyzed.[54] 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 named 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" 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 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.[55][56][57] Evolution Main article: Evolution of photosynthesis Life timeline This box: viewtalkedit −4500 —–—–−4000 —–—–−3500 —–—–−3000 —–—–−2500 —–—–−2000 —–—–−1500 —–—–−1000 —–—–−500 —–—–0 — Water Single-celled life Photosynthesis Eukaryotes Multicellular life P l a n t s Arthropods Molluscs Flowers Dinosaurs Mammals Birds Primates H a d e a n A r c h e a n P r o t e r o z o i c P h a n e r o z o i c ← Earth formed ← Earliest water ← LUCA ← Earliest fossils ← LHB meteorites ← Earliest oxygen ← Pongola glaciation* ← Atmospheric oxygen ← Huronian glaciation* ← Sexual reproduction ← Earliest multicellular life ← Earliest fungi ← Earliest plants ← Earliest animals ← Cryogenian ice age* ← Ediacaran biota ← Cambrian explosion ← Andean glaciation* ← Earliest tetrapods ← Karoo ice age* ← Earliest apes / humans ← Quaternary ice age* (million years ago)*Ice Ages Early photosynthetic systems, such as those in green and purple sulfur and green and purple nonsulfur bacteria, are thought to have been anoxygenic, and used various other molecules than water as electron donors. Green and purple sulfur bacteria are thought to have used hydrogen and sulfur as electron donors. Green nonsulfur bacteria used various amino and other organic acids as electron donors. Purple nonsulfur bacteria used a variety of nonspecific organic molecules. The use of these molecules is consistent with the geological evidence that Earth's early atmosphere was highly reducing at that time.[58] Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old.[59][60] More recent studies also suggest that photosynthesis may have begun about 3.4 billion years ago.[61][62] The main source of oxygen in the Earth's atmosphere derives from oxygenic photosynthesis, and its appearance is sometimes referred to as the oxygen catastrophe. Geological evidence suggests that oxygenic photosynthesis, such as that in cyanobacteria, became important during the Paleoproterozoic era around 2 billion years ago. Modern photosynthesis in plants and most photosynthetic prokaryotes is oxygenic, using water as an electron donor, which is oxidized to molecular oxygen in the photosynthetic reaction center. Symbiosis and the origin of chloroplasts Plant cells with visible chloroplasts (from a moss, Plagiomnium affine) Several groups of animals have formed symbiotic relationships with photosynthetic algae. These are most common in corals, sponges and sea anemones. It is presumed that this is due to the particularly simple body plans and large surface areas of these animals compared to their volumes.[63] In addition, a few marine mollusks Elysia viridis and Elysia chlorotica also maintain a symbiotic relationship with chloroplasts they capture from the algae in their diet and then store in their bodies (see Kleptoplasty). This allows the mollusks to survive solely by photosynthesis for several months at a time.[64][65] Some of the genes from the plant cell nucleus have even been transferred to the slugs, so that the chloroplasts can be supplied with proteins that they need to survive.[66] An even closer form of symbiosis may explain the origin of chloroplasts. Chloroplasts have many similarities with photosynthetic bacteria, including a circular chromosome, prokaryotic-type ribosome, and similar proteins in the photosynthetic reaction center.[67][68] The endosymbiotic theory suggests that photosynthetic bacteria were acquired (by endocytosis) by early eukaryotic cells to form the first plant cells. Therefore, chloroplasts may be photosynthetic bacteria that adapted to life inside plant cells. Like mitochondria, chloroplasts possess their own DNA, separate from the nuclear DNA of their plant host cells and the genes in this chloroplast DNA resemble those found in cyanobacteria.[69] DNA in chloroplasts codes for redox proteins such as those found in the photosynthetic reaction centers. The CoRR Hypothesis proposes that this co-location of genes with their gene products is required for redox regulation of gene expression, and accounts for the persistence of DNA in bioenergetic organelles.[70] Photosynthetic eukaryotic lineages Symbiotic and kleptoplastic organisms excluded: The glaucophytes and the red and green algae—clade Archaeplastida (uni- and multicellular) The cryptophytes—clade Cryptista (unicellular) The haptophytes—clade Haptista (unicellular) The dinoflagellates and chromerids in the superphylum Myzozoa, and Pseudoblepharisma in the phylum Ciliophora—clade Alveolata (unicellular) The ochrophytes—clade Heterokonta (uni- and multicellular) The chlorarachniophytes and 3 species of Paulinella in the phylum Cercozoa—clade Rhizaria (unicellular) The euglenids—clade Excavata (unicellular) Except for the euglenids, which are found within the Excavata, all of these belong to the Diaphoretickes. Archaeplastida and the photosynthetic Paulinella got their plastids, which are surrounded by two membranes, through primary endosymbiosis in two separate events, by engulfing a cyanobacterium. The plastids in all the other groups have either a red or green algal origin, and are referred to as the "red lineages" and the "green lineages". The only known exception is the ciliate Pseudoblepharisma tenue, which in addition to its plastids that originated from green algae also has a purple sulfur bacterium as symbiont. In dinoflagellates and euglenids the plastids are surrounded by three membranes, and in the remaining lines by four. A nucleomorph, remnants of the original algal nucleus located between the inner and outer membranes of the plastid, is present in the cryptophytes (from a red alga) and chlorarachniophytes (from a green alga).[71] Some dinoflagellates that lost their photosynthetic ability later regained it again through new endosymbiotic events with different algae. While able to perform photosynthesis, many of these eukaryotic groups are mixotrophs and practice heterotrophy to various degrees. Cyanobacteria and the evolution of photosynthesis The biochemical capacity to use water as the source for electrons in photosynthesis evolved once, in a common ancestor of extant cyanobacteria (formerly called blue-green algae), which are the only prokaryotes performing oxygenic photosynthesis. The geological record indicates that this transforming event took place early in Earth's history, at least 2450–2320 million years ago (Ma), and, it is speculated, much earlier.[72][73] Because the Earth's atmosphere contained almost no oxygen during the estimated development of photosynthesis, it is believed that the first photosynthetic cyanobacteria did not generate oxygen.[74] Available evidence from geobiological studies of Archean (>2500 Ma) sedimentary rocks indicates that life existed 3500 Ma, but the question of when oxygenic photosynthesis evolved is still unanswered. A clear paleontological window on cyanobacterial evolution opened about 2000 Ma, revealing an already-diverse biota of cyanobacteria. Cyanobacteria remained the principal primary producers of oxygen throughout the Proterozoic Eon (2500–543 Ma), in part because the redox structure of the oceans favored photoautotrophs capable of nitrogen fixation.[citation needed] Green algae joined cyanobacteria as the major primary producers of oxygen on continental shelves near the end of the Proterozoic, but only with the Mesozoic (251–66 Ma) radiations of dinoflagellates, coccolithophorids, and diatoms did the primary production of oxygen in marine shelf waters take modern form. Cyanobacteria remain critical to marine ecosystems as primary producers of oxygen in oceanic gyres, as agents of biological nitrogen fixation, and, in modified form, as the plastids of marine algae.[75] Experimental history Discovery Although some of the steps in photosynthesis are still not completely understood, the overall photosynthetic equation has been known since the 19th century. Portrait of Jan Baptist van Helmont by Mary Beale, c.1674 Jan van Helmont began the research of the process in the mid-17th century when he carefully measured the mass of the soil used by a plant and the mass of the plant as it grew. After noticing that the soil mass changed very little, he hypothesized that the mass of the growing plant must come from the water, the only substance he added to the potted plant. His hypothesis was partially accurate – much of the gained mass comes from carbon dioxide as well as water. However, this was a signaling point to the idea that the bulk of a plant's biomass comes from the inputs of photosynthesis, not the soil itself. Joseph Priestley, a chemist and minister, discovered that when he isolated a volume of air under an inverted jar and burned a candle in it (which gave off CO2), the candle would burn out very quickly, much before it ran out of wax. He further discovered that a mouse could similarly "injure" air. He then showed that the air that had been "injured" by the candle and the mouse could be restored by a plant.[76] In 1779, Jan Ingenhousz repeated Priestley's experiments. He discovered that it was the influence of sunlight on the plant that could cause it to revive a mouse in a matter of hours.[76][77] In 1796, Jean Senebier, a Swiss pastor, botanist, and naturalist, demonstrated that green plants consume carbon dioxide and release oxygen under the influence of light. Soon afterward, Nicolas-Théodore de Saussure showed that the increase in mass of the plant as it grows could not be due only to uptake of CO2 but also to the incorporation of water. Thus, the basic reaction by which photosynthesis is used to produce food (such as glucose) was outlined.[78] Refinements Cornelis Van Niel made key discoveries explaining the chemistry of photosynthesis. By studying purple sulfur bacteria and green bacteria he was the first to demonstrate that photosynthesis is a light-dependent redox reaction, in which hydrogen reduces (donates its atoms as electrons and protons to) carbon dioxide. Robert Emerson discovered two light reactions by testing plant productivity using different wavelengths of light. With the red alone, the light reactions were suppressed. When blue and red were combined, the output was much more substantial. Thus, there were two photosystems, one absorbing up to 600 nm wavelengths, the other up to 700 nm. The former is known as PSII, the latter is PSI. PSI contains only chlorophyll "a", PSII contains primarily chlorophyll "a" with most of the available chlorophyll "b", among other pigments. These include phycobilins, which are the red and blue pigments of red and blue algae, respectively, and fucoxanthol for brown algae and diatoms. The process is most productive when the absorption of quanta is equal in both PSII and PSI, assuring that input energy from the antenna complex is divided between the PSI and PSII systems, which in turn powers the photochemistry.[13] Robert Hill thought that a complex of reactions consisted of an intermediate to cytochrome b6 (now a plastoquinone), and that another was from cytochrome f to a step in the carbohydrate-generating mechanisms. These are linked by plastoquinone, which does require energy to reduce cytochrome f. Further experiments to prove that the oxygen developed during the photosynthesis of green plants came from water were performed by Hill in 1937 and 1939. He showed that isolated chloroplasts give off oxygen in the presence of unnatural reducing agents like iron oxalate, ferricyanide or benzoquinone after exposure to light. In the Hill reaction:[79] 2 H2O + 2 A + (light, chloroplasts) → 2 AH2 + O2 A is the electron acceptor. Therefore, in light, the electron acceptor is reduced and oxygen is evolved. Samuel Ruben and Martin Kamen used radioactive isotopes to determine that the oxygen liberated in photosynthesis came from the water. Melvin Calvin works in his photosynthesis laboratory. Melvin Calvin and Andrew Benson, along with James Bassham, elucidated the path of carbon assimilation (the photosynthetic carbon reduction cycle) in plants. The carbon reduction cycle is known as the Calvin cycle, but many scientists refer to it as the Calvin-Benson, Benson-Calvin, or even Calvin-Benson-Bassham (or CBB) Cycle. Nobel Prize-winning scientist Rudolph A. Marcus was later able to discover the function and significance of the electron transport chain. Otto Heinrich Warburg and Dean Burk discovered the I-quantum photosynthesis reaction that splits CO2, activated by the respiration.[80] In 1950, first experimental evidence for the existence of photophosphorylation in vivo was presented by Otto Kandler using intact Chlorella cells and interpreting his findings as light-dependent ATP formation.[81] In 1954, Daniel I. Arnon et al. discovered photophosphorylation in vitro in isolated chloroplasts with the help of P32.[82][83] Louis N.M. Duysens and Jan Amesz discovered that chlorophyll "a" will absorb one light, oxidize cytochrome f, while chlorophyll "a" (and other pigments) will absorb another light but will reduce this same oxidized cytochrome, stating the two light reactions are in series. Development of the concept In 1893, Charles Reid Barnes proposed two terms, photosyntax and photosynthesis, for the biological process of synthesis of complex carbon compounds out of carbonic acid, in the presence of chlorophyll, under the influence of light. Over time, the term photosynthesis came into common usage. Later discovery of anoxygenic photosynthetic bacteria and photophosphorylation necessitated redefinition of the term.[84] C3 : C4 photosynthesis research In the late 1940s at the University of California, Berkeley, the details of photosynthetic carbon metabolism were sorted out by the chemists Melvin Calvin, Andrew Benson, James Bassham and a score of students and researchers utilizing the carbon-14 isotope and paper chromatography techniques.[85] The pathway of CO2 fixation by the algae Chlorella in a fraction of a second in light resulted in a 3 carbon molecule called phosphoglyceric acid (PGA). For that original and ground-breaking work, a Nobel Prize in Chemistry was awarded to Melvin Calvin in 1961. In parallel, plant physiologists studied leaf gas exchanges using the new method of infrared gas analysis and a leaf chamber where the net photosynthetic rates ranged from 10 to 13 μmol CO2·m−2·s−1, with the conclusion that all terrestrial plants have the same photosynthetic capacities, that are light saturated at less than 50% of sunlight.[86][87] Later in 1958–1963 at Cornell University, field grown maize was reported to have much greater leaf photosynthetic rates of 40 μmol CO2·m−2·s−1 and not be saturated at near full sunlight.[88][89] This higher rate in maize was almost double of those observed in other species such as wheat and soybean, indicating that large differences in photosynthesis exist among higher plants. At the University of Arizona, detailed gas exchange research on more than 15 species of monocot and dicot uncovered for the first time that differences in leaf anatomy are crucial factors in differentiating photosynthetic capacities among species.[90][91] In tropical grasses, including maize, sorghum, sugarcane, Bermuda grass and in the dicot amaranthus, leaf photosynthetic rates were around 38−40 μmol CO2·m−2·s−1, and the leaves have two types of green cells, i. e. outer layer of mesophyll cells surrounding a tightly packed cholorophyllous vascular bundle sheath cells. This type of anatomy was termed Kranz anatomy in the 19th century by the botanist Gottlieb Haberlandt while studying leaf anatomy of sugarcane.[92] Plant species with the greatest photosynthetic rates and Kranz anatomy showed no apparent photorespiration, very low CO2 compensation point, high optimum temperature, high stomatal resistances and lower mesophyll resistances for gas diffusion and rates never saturated at full sun light.[93] The research at Arizona was designated a Citation Classic in 1986.[91] These species were later termed C4 plants as the first stable compound of CO2 fixation in light has 4 carbons as malate and aspartate.[94][95][96] Other species that lack Kranz anatomy were termed C3 type such as cotton and sunflower, as the first stable carbon compound is the 3-carbon PGA. At 1000 ppm CO2 in measuring air, both the C3 and C4 plants had similar leaf photosynthetic rates around 60 μmol CO2·m−2·s−1 indicating the suppression of photorespiration in C3 plants.[90][91] Factors The leaf is the primary site of photosynthesis in plants. There are three main factors affecting photosynthesis[clarification needed] and several corollary factors. The three main are:[citation needed] Light irradiance and wavelength Carbon dioxide concentration Temperature. Total photosynthesis is limited by a range of environmental factors. These include the amount of light available, the amount of leaf area a plant has to capture light (shading by other plants is a major limitation of photosynthesis), the rate at which carbon dioxide can be supplied to the chloroplasts to support photosynthesis, the availability of water, and the availability of suitable temperatures for carrying out photosynthesis.[97] Light intensity (irradiance), wavelength and temperature See also: PI (photosynthesis-irradiance) curve Absorbance spectra of free chlorophyll a (blue) and b (red) in a solvent. The action spectra of chlorophyll molecules are slightly modified in vivo depending on specific pigment–protein interactions. The process of photosynthesis provides the main input of free energy into the biosphere, and is one of four main ways in which radiation is important for plant life.[98] The radiation climate within plant communities is extremely variable, in both time and space. In the early 20th century, Frederick Blackman and Gabrielle Matthaei investigated the effects of light intensity (irradiance) and temperature on the rate of carbon assimilation. At constant temperature, the rate of carbon assimilation varies with irradiance, increasing as the irradiance increases, but reaching a plateau at higher irradiance. At low irradiance, increasing the temperature has little influence on the rate of carbon assimilation. At constant high irradiance, the rate of carbon assimilation increases as the temperature is increased. These two experiments illustrate several important points: First, it is known that, in general, photochemical reactions are not affected by temperature. However, these experiments clearly show that temperature affects the rate of carbon assimilation, so there must be two sets of reactions in the full process of carbon assimilation. These are the light-dependent 'photochemical' temperature-independent stage, and the light-independent, temperature-dependent stage. Second, Blackman's experiments illustrate the concept of limiting factors. Another limiting factor is the wavelength of light. Cyanobacteria, which reside several meters underwater, cannot receive the correct wavelengths required to cause photoinduced charge separation in conventional photosynthetic pigments. To combat this problem, a series of proteins with different pigments surround the reaction center. This unit is called a phycobilisome.[clarification needed] Carbon dioxide levels and photorespiration Photorespiration As carbon dioxide concentrations rise, the rate at which sugars are made by the light-independent reactions increases until limited by other factors. RuBisCO, the enzyme that captures carbon dioxide in the light-independent reactions, has a binding affinity for both carbon dioxide and oxygen. When the concentration of carbon dioxide is high, RuBisCO will fix carbon dioxide. However, if the carbon dioxide concentration is low, RuBisCO will bind oxygen instead of carbon dioxide. This process, called photorespiration, uses energy, but does not produce sugars. RuBisCO oxygenase activity is disadvantageous to plants for several reasons: One product of oxygenase activity is phosphoglycolate (2 carbon) instead of 3-phosphoglycerate (3 carbon). Phosphoglycolate cannot be metabolized by the Calvin-Benson cycle and represents carbon lost from the cycle. A high oxygenase activity, therefore, drains the sugars that are required to recycle ribulose 5-bisphosphate and for the continuation of the Calvin-Benson cycle. Phosphoglycolate is quickly metabolized to glycolate that is toxic to a plant at a high concentration; it inhibits photosynthesis. Salvaging glycolate is an energetically expensive process that uses the glycolate pathway, and only 75% of the carbon is returned to the Calvin-Benson cycle as 3-phosphoglycerate. The reactions also produce ammonia (NH3), which is able to diffuse out of the plant, leading to a loss of nitrogen. A highly simplified summary is: 2 glycolate + ATP → 3-phosphoglycerate + carbon dioxide + ADP + NH3 The salvaging pathway for the products of RuBisCO oxygenase activity is more commonly known as photorespiration, since it is characterized by light-dependent oxygen consumption and the release of carbon dioxide. [New Trading View Logo](