Photosynthesis

Photosynthesis

Overview

Photosynthesis is a chemical process that converts carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 into organic compound
Organic compound
An organic compound is any member of a large class of gaseous, liquid, or solid chemical compounds whose molecules contain carbon. For historical reasons discussed below, a few types of carbon-containing compounds such as carbides, carbonates, simple oxides of carbon, and cyanides, as well as the...

s, especially sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

s, using the energy from sunlight. Photosynthesis occurs in plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s, algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

, and many species of bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

, but not in archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

. Photosynthetic organisms are called photoautotrophs, since they can create their own food. In plants, algae, and cyanobacteria, photosynthesis uses carbon dioxide and water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

, releasing oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

 as a waste product.
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Encyclopedia

Photosynthesis is a chemical process that converts carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 into organic compound
Organic compound
An organic compound is any member of a large class of gaseous, liquid, or solid chemical compounds whose molecules contain carbon. For historical reasons discussed below, a few types of carbon-containing compounds such as carbides, carbonates, simple oxides of carbon, and cyanides, as well as the...

s, especially sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

s, using the energy from sunlight. Photosynthesis occurs in plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s, algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

, and many species of bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

, but not in archaea
Archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

. Photosynthetic organisms are called photoautotrophs, since they can create their own food. In plants, algae, and cyanobacteria, photosynthesis uses carbon dioxide and water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

, releasing oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

 as a waste product. Photosynthesis is vital for all aerobic life on Earth
Life
Life is a characteristic that distinguishes objects that have signaling and self-sustaining processes from those that do not, either because such functions have ceased , or else because they lack such functions and are classified as inanimate...

. In addition to maintaining normal levels of oxygen in the atmosphere
Atmosphere
An atmosphere is a layer of gases that may surround a material body of sufficient mass, and that is held in place by the gravity of the body. An atmosphere may be retained for a longer duration, if the gravity is high and the atmosphere's temperature is low...

, photosynthesis is the source of energy for nearly all life on earth, either directly, through primary production
Primary production
400px|thumb|Global oceanic and terrestrial photoautotroph abundance, from September [[1997]] to August 2000. As an estimate of autotroph biomass, it is only a rough indicator of primary production potential, and not an actual estimate of it...

, or indirectly, as the ultimate source of the energy in their food, the exceptions being chemoautotrophs that live in rocks or around deep sea hydrothermal vents. The rate of energy capture by photosynthesis is immense, approximately 100 terawatts, which is about six times larger than the power consumption of human civilization
World energy resources and consumption
]World energy consumption in 2010: over 5% growthEnergy markets have combined crisis recovery and strong industry dynamism. Energy consumption in the G20 soared by more than 5% in 2010, after the slight decrease of 2009. This strong increase is the result of two converging trends...

. As well as energy, photosynthesis is also the source of the carbon in all the organic compounds within organisms' bodies. In all, photosynthetic organisms convert around 100–115  petagrams of carbon into biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....

 per year.

Although photosynthesis can happen in different ways in different species, some features are always the same. For example, the process always begins when energy from light is absorbed by protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

s called photosynthetic reaction centers that contain chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

s. In plants, these proteins are held inside organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....

s called chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

s, while in bacteria they are embedded in the plasma membrane. Some of the light energy gathered by chlorophylls is stored in the form of adenosine triphosphate
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

 (ATP). The rest of the energy is used to remove electron
Electron
The electron is a subatomic particle with a negative elementary electric charge. It has no known components or substructure; in other words, it is generally thought to be an elementary particle. An electron has a mass that is approximately 1/1836 that of the proton...

s from a substance such as water. These electrons are then used in the reactions that turn carbon dioxide into organic compounds. In plants, algae and cyanobacteria, this is done by a sequence of reactions called the Calvin cycle
Calvin cycle
The Calvin cycle or Calvin–Benson-Bassham cycle or reductive pentose phosphate cycle or C3 cycle or CBB cycle is a series of biochemical redox reactions that take place in the stroma of chloroplasts in photosynthetic organisms...

, but different sets of reactions are found in some bacteria, such as the reverse Krebs cycle
Reverse Krebs cycle
The reverse Krebs cycle...

 in Chlorobium
Chlorobium
Chlorobium is a genus of green sulfur bacteria. They are photolithotrophic oxidizers of sulfur and most notably utilise a noncyclic electron transport chain to reduce NAD+...

. Many photosynthetic organisms have adaptation
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....

s that concentrate or store carbon dioxide. This helps reduce a wasteful process called photorespiration
Photorespiration
Photorespiration, or "'photo-respiration'", is a process in plant metabolism by which RuBP has oxygen added to it by the enzyme , instead of carbon dioxide during normal photosynthesis. This is the beginning step of the Calvin-Benson cycle...

 that can consume part of the sugar produced during photosynthesis.

The first photosynthetic organisms probably evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 about , early in the evolutionary history of life
Evolutionary history of life
The evolutionary history of life on Earth traces the processes by which living and fossil organisms have evolved since life on Earth first originated until the present day. Earth formed about 4.5 Ga and life appeared on its surface within one billion years...

, when all forms of life on Earth were microorganism
Microorganism
A microorganism or microbe is a microscopic organism that comprises either a single cell , cell clusters, or no cell at all...

s and the atmosphere had much more carbon dioxide. They most likely used hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 or hydrogen sulfide
Hydrogen sulfide
Hydrogen sulfide is the chemical compound with the formula . It is a colorless, very poisonous, flammable gas with the characteristic foul odor of expired eggs perceptible at concentrations as low as 0.00047 parts per million...

 as sources of electrons, rather than water. Cyanobacteria appeared later, around , and drastically changed the Earth when they began to oxygenate the atmosphere
Oxygen Catastrophe
The Great Oxygenation Event , also called the Oxygen Catastrophe or Oxygen Crisis or Great Oxidation, was the biologically induced appearance of free oxygen in Earth's atmosphere. This major environmental change happened around 2.4 billion years ago.Photosynthesis was producing oxygen both before...

, beginning about . This new atmosphere allowed the evolution of complex life such as protist
Protist
Protists are a diverse group of eukaryotic microorganisms. Historically, protists were treated as the kingdom Protista, which includes mostly unicellular organisms that do not fit into the other kingdoms, but this group is contested in modern taxonomy...

s. Eventually, no later than a billion years ago, one of these protists formed a symbiotic relationship
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 with a cyanobacterium, producing the ancestor of many plants and algae. The chloroplasts in modern plants are the descendants of these ancient symbiotic cyanobacteria.

Overview



Photosynthetic organisms are photoautotrophs, which means that they are repositories of energy, they are able to synthesize
Chemical synthesis
In chemistry, chemical synthesis is purposeful execution of chemical reactions to get a product, or several products. This happens by physical and chemical manipulations usually involving one or more reactions...

 food directly from carbon dioxide, water, and using energy from light. They accrue it as part of their potential energy
Potential energy
In physics, potential energy is the energy stored in a body or in a system due to its position in a force field or due to its configuration. The SI unit of measure for energy and work is the Joule...

. However, not all organisms that use light as a source of energy carry out photosynthesis, since photoheterotroph
Photoheterotroph
Photoheterotrophs are heterotrophic organisms that use light for energy, but cannot use carbon dioxide as their sole carbon source. Consequently, they use organic compounds from the environment to satisfy their carbon requirements. They use compounds such as carbohydrates, fatty acids and...

s use organic compounds, rather than carbon dioxide, as a source of carbon. In plants, algae and cyanobacteria, photosynthesis releases oxygen. This is called oxygenic photosynthesis. Although there are some differences between oxygenic photosynthesis in plants, algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

 and cyanobacteria, the overall process is quite similar in these organisms. However, there are some types of bacteria that carry out anoxygenic photosynthesis
Anoxygenic photosynthesis
Phototrophy is the process by which organisms trap light energy and store it as chemical energy in the form of ATP. There are three major types of phototrophy: Oxygenic photosynthesis, Anoxygenic photosynthesis, and Rhodopsin-based phototrophy...

, which consumes carbon dioxide but does not release oxygen.

Carbon dioxide is converted into sugars in a process called carbon fixation
Carbon fixation
In biology, carbon fixation is the reduction of carbon dioxide to organic compounds by living organisms. The obvious example is photosynthesis. Carbon fixation requires both a source of energy such as sunlight, and an electron donor such as water. All life depends on fixed carbon. Organisms that...

. Carbon fixation is a redox
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

 reaction, so photosynthesis needs to supply both a source of energy to drive this process, and the electrons needed to convert carbon dioxide into a carbohydrate
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

, which is a reduction reaction. In general outline, photosynthesis is the opposite of cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

, where glucose and other compounds are oxidized to produce carbon dioxide, water, and release chemical energy. However, the two processes take place through a different sequence of chemical reactions and in different cellular compartments.

The general equation
Chemical equation
A chemical equation is the symbolic representation of a chemical reaction where the reactant entities are given on the left hand side and the product entities on the right hand side. The coefficients next to the symbols and formulae of entities are the absolute values of the stoichiometric numbers...

 for photosynthesis is therefore:
2n CO2 + 2n DH2 + photons → 2(CH2O)n
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

 + 2n DO


Carbon dioxide + electron donor + light energy → carbohydrate + oxidized electron donor

In oxygenic photosynthesis water is the electron donor and, since its hydrolysis
Hydrolysis
Hydrolysis is a chemical reaction during which molecules of water are split into hydrogen cations and hydroxide anions in the process of a chemical mechanism. It is the type of reaction that is used to break down certain polymers, especially those made by condensation polymerization...

 releases oxygen, the equation for this process is:
2n CO2 + 4n H2O + photons → 2(CH2O)n
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

 + 2n O2 + 2n H2O
carbon dioxide + water + light energy → carbohydrate + oxygen + water

Often 2n water molecules are cancelled on both sides, yielding:
2n CO2 + 2n H2O + photons → 2(CH2O)n
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

 + 2n O2
carbon dioxide + water + light energy → carbohydrate + oxygen


Other processes substitute other compounds (such as arsenite
Arsenite
In chemistry an arsenite is a chemical compound containing an arsenic oxoanion where arsenic has oxidation state +3.The different forms of the anion are the next ones:* ortho-arsenite: AsO33-* meta-arsenite: AsO2-...

) for water in the electron-supply role; the microbes use sunlight to oxidize arsenite to arsenate
Arsenate
The arsenate ion is AsO43−.An arsenate is any compound that contains this ion. Arsenates are salts or esters of arsenic acid.The arsenic atom in arsenate has a valency of 5 and is also known as pentavalent arsenic or As[V]....

: The equation for this reaction is:
CO2 + (AsO33–) + photons → (AsO43–) + CO
carbon dioxide + arsenite + light energy → arsenate + carbon monoxide (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 energy-storage molecules ATP
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

 and NADPH. During the second stage, the light-independent reactions use these products to capture and reduce carbon dioxide.

Most organisms that utilize photosynthesis to produce oxygen use visible light
Visible spectrum
The visible spectrum is the portion of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called visible light or simply light. A typical human eye will respond to wavelengths from about 390 to 750 nm. In terms of...

 to do so, although at least three use infrared radiation
Infrared
Infrared light is electromagnetic radiation with a wavelength longer than that of visible light, measured from the nominal edge of visible red light at 0.74 micrometres , and extending conventionally to 300 µm...

.

Photosynthetic membranes and organelles




The proteins that gather light for photosynthesis are embedded within cell membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

s. The simplest way these are arranged is in photosynthetic bacteria, where these proteins are held within the plasma membrane. However, this membrane may be tightly folded into cylindrical sheets called thylakoid
Thylakoid
A thylakoid is a membrane-bound compartment inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thylakoids frequently form stacks of disks referred to as...

s, or bunched up into round vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

 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 organelle
Organelle
In cell biology, an organelle is a specialized subunit within a cell that has a specific function, and is usually separately enclosed within its own lipid bilayer....

s called chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

s. A typical plant cell
Plant cell
Plant cells are eukaryotic cells that differ in several key respects from the cells of other eukaryotic organisms. Their distinctive features include:...

 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 between them. Within the membrane is an aqueous fluid called the stroma. The stroma contains stacks (grana) of thylakoids, which are the site of photosynthesis. The thylakoids are flattened disks, bounded by a membrane with a lumen or thylakoid space within it. The site of photosynthesis is the thylakoid membrane, which contains integral and peripheral membrane protein
Peripheral membrane protein
Peripheral membrane proteins are proteins that adhere only temporarily to the biological membrane with which they are associated. These molecules attach to integral membrane proteins, or penetrate the peripheral regions of the lipid bilayer. The regulatory protein subunits of many ion channels and...

 complexes, including the pigments that absorb light energy, which form the photosystems.

Plants absorb light primarily using the pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

 chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

, which is the reason that most plants have a green color. Besides chlorophyll, plants also use pigments such as carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

s and xanthophyll
Xanthophyll
Xanthophylls are yellow pigments that form one of two major divisions of the carotenoid group. The name is from Greek xanthos + phyllon , due to their formation of the yellow band seen in early chromatography of leaf pigments...

s.
Algae also use chlorophyll, but various other pigments are present as phycocyanin
Phycocyanin
Phycocyanin is a pigment from the light-harvesting phycobiliprotein family, along with allophycocyanin and phycoerythrin. It is an accessory pigment to chlorophyll. All phycobiliproteins are water-soluble and therefore cannot exist within the membrane as do carotenoids, but aggregate forming...

, carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

s, and xanthophyll
Xanthophyll
Xanthophylls are yellow pigments that form one of two major divisions of the carotenoid group. The name is from Greek xanthos + phyllon , due to their formation of the yellow band seen in early chromatography of leaf pigments...

s in green algae
Green algae
The green algae are the large group of algae from which the embryophytes emerged. As such, they form a paraphyletic group, although the group including both green algae and embryophytes is monophyletic...

, phycoerythrin
Phycoerythrin
Phycoerythrin is a red protein from the light-harvesting phycobiliprotein family, present in cyanobacteria, red algae and cryptomonads.Like all phycobiliproteins, phycoerythrin is composed of a protein part, organised in a hexameric structure of alpha and beta chains, covalently binding...

 in red algae
Red algae
The red algae are one of the oldest groups of eukaryotic algae, and also one of the largest, with about 5,000–6,000 species  of mostly multicellular, marine algae, including many notable seaweeds...

 (rhodophytes) and fucoxanthin
Fucoxanthin
Fucoxanthin is a xanthophyll, with formula C42H58O6. It is found as an accessory pigment in the chloroplasts of brown algae and most other heterokonts, giving them a brown or olive-green color...

 in brown algae
Brown algae
The Phaeophyceae or brown algae , is a large group of mostly marine multicellular algae, including many seaweeds of colder Northern Hemisphere waters. They play an important role in marine environments, both as food and for the habitats they form...

 and diatoms resulting in a wide variety of colors.

These pigments are embedded in plants and algae in special antenna-proteins. In such proteins all the pigments are ordered to work well together. Such a protein is also called a light-harvesting complex
Light-harvesting complex
A light-harvesting complex is a complex of subunit proteins that may be part of a larger supercomplex of a photosystem, the functional unit in photosynthesis. It is used by plants and photosynthetic bacteria to collect more of the incoming light than would be captured by the photosynthetic reaction...

.

Although all cells in the green parts of a plant have chloroplasts, most of the energy is captured in the leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

. The cells in the interior tissues of a leaf, called the mesophyll
Mesophyll
Mesophyll can refer to:* Mesophyll tissue, in plant anatomy, photosynthetic parenchyma cells that lie between the upper and lower epidermis layers of a leaf...

, can contain between 450,000 and 800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is uniformly coated with a water-resistant wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...

y cuticle
Plant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...

 that protects the leaf from excessive evaporation
Evaporation
Evaporation is a type of vaporization of a liquid that occurs only on the surface of a liquid. The other type of vaporization is boiling, which, instead, occurs on the entire mass of the liquid....

 of water and decreases the absorption of ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...

 or blue
Blue
Blue is a colour, the perception of which is evoked by light having a spectrum dominated by energy with a wavelength of roughly 440–490 nm. It is considered one of the additive primary colours. On the HSV Colour Wheel, the complement of blue is yellow; that is, a colour corresponding to an equal...

 light
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

 to reduce heat
Heat
In physics and thermodynamics, heat is energy transferred from one body, region, or thermodynamic system to another due to thermal contact or thermal radiation when the systems are at different temperatures. It is often described as one of the fundamental processes of energy transfer between...

ing. The transparent epidermis layer allows light to pass through to the palisade mesophyll cells where most of the photosynthesis takes place.

Light reactions




In the light reaction
Light-dependent reactions
The 'light-dependent reactions', or light reactions, are the first stage of photosynthesis, the process by which plants capture and store energy from sunlight. In this process, light energy is converted into chemical energy, in the form of the energy-carrying molecules ATP and NADPH...

s, one molecule of the pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

 chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

 absorbs one photon
Photon
In physics, a photon is an elementary particle, the quantum of the electromagnetic interaction and the basic unit of light and all other forms of electromagnetic radiation. It is also the force carrier for the electromagnetic force...

 and loses one electron
Electron
The electron is a subatomic particle with a negative elementary electric charge. It has no known components or substructure; in other words, it is generally thought to be an elementary particle. An electron has a mass that is approximately 1/1836 that of the proton...

. This electron is passed to a modified form of chlorophyll called pheophytin
Pheophytin
Pheophytin or phaeophytin is a chemical compound that serves as the first electron carrier intermediate in the electron transfer pathway of photosystem II in plants, and the photosynthetic reaction center found in purple bacteria...

, which passes the electron to a quinone
Quinone
A quinone is a class of organic compounds that are formally "derived from aromatic compounds [such as benzene or naphthalene] by conversion of an even number of –CH= groups into –C– groups with any necessary rearrangement of double bonds," resulting in "a fully conjugated cyclic dione structure."...

 molecule, allowing the start of a flow of electrons down an electron transport chain
Electron transport chain
An electron transport chain couples electron transfer between an electron donor and an electron acceptor with the transfer of H+ ions across a membrane. The resulting electrochemical proton gradient is used to generate chemical energy in the form of adenosine triphosphate...

 that leads to the ultimate reduction of NADP
Nicotinamide adenine dinucleotide phosphate
Nicotinamide adenine dinucleotide phosphate, abbreviated NADP or TPN in older notation , is a coenzyme used in anabolic reactions, such as lipid and nucleic acid synthesis, which require NADPH as a reducing agent....

 to NADPH
Nicotinamide adenine dinucleotide phosphate
Nicotinamide adenine dinucleotide phosphate, abbreviated NADP or TPN in older notation , is a coenzyme used in anabolic reactions, such as lipid and nucleic acid synthesis, which require NADPH as a reducing agent....

. In addition, this creates a proton gradient
Electrochemical gradient
An electrochemical gradient is a spatial variation of both electrical potential and chemical concentration across a membrane; that is, a combination of the membrane potential and the pH gradient...

 across the chloroplast membrane
Chloroplast membrane
Chloroplasts contain several important membranes, vital for their function. Like mitochondria, chloroplasts have a double-membrane envelope, called the chloroplast envelope. Each membrane is a phospholipid bilayer, between 6 and 8 nm thick, and the two are separated by a gap of 10-20 nm,...

; its dissipation is used by ATP synthase
ATP synthase
right|thumb|300px|Molecular model of ATP synthase by X-ray diffraction methodATP synthase is an important enzyme that provides energy for the cell to use through the synthesis of adenosine triphosphate . ATP is the most commonly used "energy currency" of cells from most organisms...

 for the concomitant synthesis of ATP
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

. The chlorophyll molecule regains the lost electron from a water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

 molecule through a process called photolysis
Photodissociation
Photodissociation, photolysis, or photodecomposition is a chemical reaction in which a chemical compound is broken down by photons. It is defined as the interaction of one or more photons with one target molecule....

, which releases a dioxygen (O2) molecule.
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 wavelength
Wavelength
In physics, the wavelength of a sinusoidal wave is the spatial period of the wave—the distance over which the wave's shape repeats.It is usually determined by considering the distance between consecutive corresponding points of the same phase, such as crests, troughs, or zero crossings, and is a...

s of light can support photosynthesis. The photosynthetic action spectrum depends on the type of accessory pigment
Accessory pigment
Accessory pigments are light-absorbing compounds, found in photosynthetic organisms, that work in conjunction with chlorophyll a. They include other forms of this pigment, such as chlorophyll b in green algal and higher plant antennae, while other algae may contain chlorophyll c or d...

s present. For example, in green plants, the action spectrum
Action spectrum
An action spectrum is the rate of a physiological activity plotted against wavelength of light. It shows which wavelength of light is most effectively used in a specific chemical reaction. Some reactants are able to use specific wavelengths of light more effectively to complete their reactions...

 resembles the absorption spectrum for chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

s and carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

s with peaks for violet-blue and red light. In red algae, the action spectrum overlaps with the absorption spectrum of phycobilin
Phycobilin
Phycobilins are chromophores found in cyanobacteria and in the chloroplasts of red algae, glaucophytes and some cryptomonads...

s for red blue-green light, which allows these algae to grow in deeper waters that filter out the longer wavelengths used by green plants. The non-absorbed part of the light spectrum
Electromagnetic spectrum
The electromagnetic spectrum is the range of all possible frequencies of electromagnetic radiation. The "electromagnetic spectrum" of an object is the characteristic distribution of electromagnetic radiation emitted or absorbed by that particular object....

 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



In plants, light-dependent reactions occur in the thylakoid membranes of the chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

s and use light energy to synthesize ATP and NADPH. The light-dependent reaction has two forms: cyclic and non-cyclic. In the non-cyclic reaction, the photon
Photon
In physics, a photon is an elementary particle, the quantum of the electromagnetic interaction and the basic unit of light and all other forms of electromagnetic radiation. It is also the force carrier for the electromagnetic force...

s are captured in the light-harvesting antenna complex
Antenna complex
The light-harvesting complex of plants is an array of protein and chlorophyll molecules embedded in the thylakoid membrane which transfer light energy to one chlorophyll a molecule at the reaction center of a photosystem....

es of photosystem II
Photosystem
Photosystems are functional and structural units of protein complexes involved in photosynthesis that together carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons...

 by chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

 and other accessory pigments (see diagram at right). When a chlorophyll molecule at the core of the photosystem II reaction center obtains sufficient excitation energy from the adjacent antenna pigments, an electron is transferred to the primary electron-acceptor molecule, pheophytin, through a process called photoinduced charge separation
Photoinduced charge separation
Photoinduced charge separation is the process of an electron in an atom being excited to a higher energy level by the absorption of a photon and then leaving the atom to a nearby electron acceptor.-Rutherford model:...

. These electrons are shuttled through an electron transport chain, the so-called Z-scheme shown in the diagram, that initially functions to generate a chemiosmotic potential across the membrane. An ATP synthase
ATP synthase
right|thumb|300px|Molecular model of ATP synthase by X-ray diffraction methodATP synthase is an important enzyme that provides energy for the cell to use through the synthesis of adenosine triphosphate . ATP is the most commonly used "energy currency" of cells from most organisms...

 enzyme uses the chemiosmotic potential to make ATP during photophosphorylation, whereas NADPH is a product of the terminal redox
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

 reaction in the Z-scheme. The electron enters a chlorophyll molecule in Photosystem I
Photosystem I
Photosystem I is the second photosystem in the photosynthetic light reactions of algae, plants, and some bacteria. Photosystem I is so named because it was discovered before photosystem II. Aspects of PS I were discovered in the 1950s, but the significances of these discoveries was not yet known...

. The electron is excited due to the light absorbed by the photosystem
Photosystem
Photosystems are functional and structural units of protein complexes involved in photosynthesis that together carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons...

. A second electron carrier accepts the electron, which again is passed down lowering energies of electron acceptor
Electron acceptor
An electron acceptor is a chemical entity that accepts electrons transferred to it from another compound. It is an oxidizing agent that, by virtue of its accepting electrons, is itself reduced in the process....

s. The energy created by the electron acceptors is used to move hydrogen ions across the thylakoid membrane into the lumen. The electron is used to reduce the co-enzyme NADP, which has functions in the light-independent reaction. The cyclic reaction is similar to that of the non-cyclic, but differs in the form 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 was emitted, hence the name cyclic reaction.

Water photolysis



The NADPH is the main reducing agent
Reducing agent
A reducing agent is the element or compound in a reduction-oxidation reaction that donates an electron to another species; however, since the reducer loses an electron we say it is "oxidized"...

 in chloroplasts, providing a source of energetic electrons to other reactions. Its production leaves chlorophyll with a deficit of electrons (oxidized), which must be obtained from some other reducing agent. The excited electrons lost from chlorophyll in photosystem I
Photosystem I
Photosystem I is the second photosystem in the photosynthetic light reactions of algae, plants, and some bacteria. Photosystem I is so named because it was discovered before photosystem II. Aspects of PS I were discovered in the 1950s, but the significances of these discoveries was not yet known...

 are replaced from the electron transport chain by plastocyanin
Plastocyanin
Plastocyanin is an important copper-containing protein involved in electron-transfer. The protein is monomeric, with a molecular weight around 10,500 Daltons, and 99 amino acids in most vascular plants...

. However, since photosystem II
Photosystem II
Photosystem II is the first protein complex in the Light-dependent reactions. It is located in the thylakoid membrane of plants, algae, and cyanobacteria. The enzyme uses photons of light to energize electrons that are then transferred through a variety of coenzymes and cofactors to reduce...

 includes the first steps of the Z-scheme, an external source of electrons is required to reduce its oxidized chlorophyll a molecules. The source of electrons in green-plant and cyanobacterial photosynthesis is water. Two water molecules are oxidized by four successive charge-separation reactions by photosystem II to yield a molecule of diatomic oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

 and four hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 ions; the electron yielded in each step is transferred to a redox-active tyrosine
Tyrosine
Tyrosine or 4-hydroxyphenylalanine, is one of the 22 amino acids that are used by cells to synthesize proteins. Its codons are UAC and UAU. It is a non-essential amino acid with a polar side group...

 residue that then reduces the photoxidized paired-chlorophyll a species called P680 that serves as the primary (light-driven) electron donor in the photosystem II reaction center. The oxidation of water is catalyzed
Catalysis
Catalysis is the change in rate of a chemical reaction due to the participation of a substance called a catalyst. Unlike other reagents that participate in the chemical reaction, a catalyst is not consumed by the reaction itself. A catalyst may participate in multiple chemical transformations....

 in photosystem II by a redox-active structure that contains four manganese
Manganese
Manganese is a chemical element, designated by the symbol Mn. It has the atomic number 25. It is found as a free element in nature , and in many minerals...

 ions and a calcium ion; this oxygen-evolving complex binds two water molecules and stores the four oxidizing equivalents that are required to drive the water-oxidizing reaction. Photosystem II is the only known biological enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

 that carries out this oxidation of water. The hydrogen ions 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 for cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

, including photosynthetic organisms.

The Calvin Cycle



In the light-independent or dark reactions the enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

 RuBisCO
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...

 captures CO2
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 from the atmosphere
Earth's atmosphere
The atmosphere of Earth is a layer of gases surrounding the planet Earth that is retained by Earth's gravity. The atmosphere protects life on Earth by absorbing ultraviolet solar radiation, warming the surface through heat retention , and reducing temperature extremes between day and night...

 and in a process that requires the newly formed NADPH, called the Calvin-Benson Cycle, 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:
3 CO2 + 9 ATP + 6 NADPH + 6 H+ → C3H6O3-phosphate + 9 ADP + 8 Pi + 6 NADP+ + 3 H2O

To be more specific, carbon fixation produces an intermediate product, which is then converted to the final carbohydrate products. The carbon skeletons produced by photosynthesis are then variously used to form other organic compounds, such as the building material cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

, as precursors for lipid
Lipid
Lipids constitute a broad group of naturally occurring molecules that include fats, waxes, sterols, fat-soluble vitamins , monoglycerides, diglycerides, triglycerides, phospholipids, and others...

 and amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

 biosynthesis, or as a fuel in cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

. The latter occurs not only in plants but also in animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s when the energy from plants gets passed through a food chain
Food chain
A food web depicts feeding connections in an ecological community. Ecologists can broadly lump all life forms into one of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs...

.

The fixation or reduction of carbon dioxide is a process in which carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 combines with a five-carbon sugar, ribulose 1,5-bisphosphate (RuBP), to yield two molecules of a three-carbon compound, glycerate 3-phosphate
Glycerate 3-phosphate
3-Phosphoglyceric acid , or glycerate 3-phosphate , is a biochemically significant 3-carbon molecule that is a metabolic intermediate in both glycolysis and the Calvin cycle. This chemical is often termed PGA when referring to the Calvin cycle...

 (GP), also known as 3-phosphoglycerate (PGA). GP, in the presence of ATP
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

 and NADPH from the light-dependent stages, is reduced to glyceraldehyde 3-phosphate
Glyceraldehyde 3-phosphate
Glyceraldehyde 3-phosphate, also known as triose phosphate or 3-phosphoglyceraldehyde and abbreviated as G3P, GADP, GAP, TP, GALP or PGAL, is a chemical compound that occurs as an intermediate in several central metabolic pathways of all organisms...

 (G3P). This product is also referred to as 3-phosphoglyceraldehyde (PGAL) or even as triose phosphate. Triose
Triose
A triose is a monosaccharide, or simple sugar, containing three carbon atoms. There are only three possible trioses: L-Glyceraldehyde and D-Glyceraldehyde, both aldotrioses because the carbonyl group is at the end of the chain, and dihydroxyacetone, a ketotriose because the carbonyl group is in...

 is a 3-carbon sugar (see carbohydrate
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

s). Most (5 out of 6 molecules) of the G3P produced is used to regenerate RuBP so the process can continue (see Calvin-Benson cycle). The 1 out of 6 molecules of the triose phosphates not "recycled" often condense to form hexose
Hexose
In organic chemistry, a hexose is a monosaccharide with six carbon atoms, having the chemical formula C6H12O6. Hexoses are classified by functional group, with aldohexoses having an aldehyde at position 1, and ketohexoses having a ketone at position 2....

 phosphates, which ultimately yield sucrose
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...

, starch
Starch
Starch or amylum is a carbohydrate consisting of a large number of glucose units joined together by glycosidic bonds. This polysaccharide is produced by all green plants as an energy store...

 and cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

. The sugars produced during carbon metabolism
Metabolism
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...

 yield carbon skeletons that can be used for other metabolic reactions like the production of amino acids and lipids.

On land



In hot and dry conditions, plants close their stomata to prevent the loss of water. Under these conditions, will decrease, and oxygen gas, produced by the light reactions of photosynthesis, will decrease in the stem, not leaves, causing an increase of photorespiration
Photorespiration
Photorespiration, or "'photo-respiration'", is a process in plant metabolism by which RuBP has oxygen added to it by the enzyme , instead of carbon dioxide during normal photosynthesis. This is the beginning step of the Calvin-Benson cycle...

 by the oxygenase
Oxygenase
An oxygenase is any enzyme that oxidizes a substrate by transferring the oxygen from molecular oxygen O2 to it. The oxygenases form a class of oxidoreductases; their EC number is EC 1.13 or EC 1.14....

 activity of ribulose-1,5-bisphosphate carboxylase/oxygenase
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...

 and decrease in carbon fixation. Some plants have evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 mechanisms to increase the concentration in the leaves under these conditions.
C4 plants
C4 carbon fixation
C4 carbon fixation is one of three biochemical mechanisms, along with and CAM photosynthesis, used in carbon fixation. It is named for the 4-carbon molecule present in the first product of carbon fixation in these plants, in contrast to the 3-carbon molecule products in plants. fixation is an...

 chemically fix carbon dioxide in the cells of the mesophyll
Mesophyll
Mesophyll can refer to:* Mesophyll tissue, in plant anatomy, photosynthetic parenchyma cells that lie between the upper and lower epidermis layers of a leaf...

 by adding it to the three-carbon molecule phosphoenolpyruvate (PEP)
Phosphoenolpyruvate
Phosphoenolpyruvic acid , or phosphoenolpyruvate as the anion, is an important chemical compound in biochemistry. It has the high-energy phosphate bond found in living organisms, and is involved in glycolysis and gluconeogenesis...

, a reaction catalyzed by an enzyme called PEP carboxylase, creating the four-carbon organic acid oxaloacetic acid
Oxaloacetic acid
Oxaloacetic acid is an organic compound with the chemical formula C4H4O5 or HOOC---COOH. It also has other names ....

. Oxaloacetic acid or malate
Malate
Malate is the ionized form of malic acid. It is an important chemical compound in biochemistry. In the C4 carbon fixation process, malate is a source of CO2 in the Calvin cycle....

 synthesized by this process is then translocated to specialized bundle sheath cells where the enzyme RuBisCO
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...

 and other Calvin cycle enzymes are located, and where released by decarboxylation
Decarboxylation
Decarboxylation is a chemical reaction that releases carbon dioxide . Usually, decarboxylation refers to a reaction of carboxylic acids, removing a carbon atom from a carbon chain. The reverse process, which is the first chemical step in photosynthesis, is called carbonation, the addition of CO2 to...

 of the four-carbon acids is then fixed by RuBisCO activity to the three-carbon sugar 3-phosphoglyceric acids. The physical separation of RuBisCO from the oxygen-generating light reactions reduces photorespiration and increases fixation and, thus, photosynthetic capacity
Photosynthetic capacity
Photosynthetic capacity is a measure of the maximum rate at which leaves are able to fix carbon during photosynthesis. It is typically measured as the amount of carbon dioxide that is fixed per metre squared per second, for example as μmol m-2 sec-1....

 of the leaf. plants can produce more sugar than plants in conditions of high light and temperature. Many important crop plants are plants, including maize, sorghum, sugarcane, and millet. Plants that do not use PEP-carboxylase in carbon fixation are called C3 plants
C3 carbon fixation
carbon fixation is a metabolic pathway for carbon fixation in photosynthesis. This process converts carbon dioxide and ribulose bisphosphate into 3-phosphoglycerate through the following reaction:...

 because the primary carboxylation reaction, catalyzed by RuBisCO, produces the three-carbon sugar 3-phosphoglyceric acids directly in the Calvin-Benson cycle. Over 90% of plants use carbon fixation, compared to 3% that use carbon fixation.
Xerophytes, such as cacti
Cacti
-See also:* RRDtool The underlying software upon which Cacti is built* MRTG The original Multi Router Traffic Grapher from which RRDtool was "extracted".* Munin -External links:******...

 and most succulents, also use PEP carboxylase to capture carbon dioxide in a process called Crassulacean acid metabolism (CAM). In contrast to metabolism, which physically separates the fixation to PEP from the Calvin cycle, CAM temporally separates these two processes. CAM plants have a different leaf anatomy from plants, and fix the at night, when their stomata are open. CAM plants store the mostly in the form of malic acid
Malic acid
Malic acid is an organic compound with the formula HO2CCH2CHOHCO2H. It is a dicarboxylic acid which is made by all living organisms, contributes to the pleasantly sour taste of fruits, and is used as a food additive. Malic acid has two stereoisomeric forms , though only the L-isomer exists...

 via carboxylation of phosphoenolpyruvate
Phosphoenolpyruvate
Phosphoenolpyruvic acid , or phosphoenolpyruvate as the anion, is an important chemical compound in biochemistry. It has the high-energy phosphate bond found in living organisms, and is involved in glycolysis and gluconeogenesis...

 to oxaloacetate, which is then reduced to malate. Decarboxylation of malate during the day releases inside the leaves, thus allowing carbon fixation to 3-phosphoglycerate by RuBisCO. Sixteen thousand species of plants use CAM.

In water


Cyanobacteria possess carboxysome
Carboxysome
Carboxysomes are bacterial microcompartments that contain enzymes involved in carbon fixation. Carboxysomes are made of polyhedral protein shells about 80 to 140 nanometres in diameter. These compartments are thought to concentrate carbon dioxide to overcome the inefficiency of RuBisCo - the...

s, which increase the concentration of around RuBisCO to increase the rate of photosynthesis. This operates by carbonic anhydrase
Carbonic anhydrase
The carbonic anhydrases form a family of enzymes that catalyze the rapid interconversion of carbon dioxide and water to bicarbonate and protons , a reversible reaction that occurs rather slowly in the absence of a catalyst...

, producing hydrocarbonate ions (HCO3), which are then pumped into the carboxysome, before being processed by a different carbonic anhydrase to produce . Pyrenoid
Pyrenoid
In cell biology, pyrenoids are organelles, centers of carbon dioxide fixation within the chloroplasts of algae and hornworts. Pyrenoids are not membrane-bound, but specialized areas of the plastid that contain high levels of ribulose-1,5-bisphosphate carboxylase/oxygenase .RubisCO fixes carbon...

s in algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

 and hornwort
Hornwort
Hornworts are a group of bryophytes, or non-vascular plants, comprising the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. The flattened, green plant body of a hornwort is the gametophyte plant.Hornworts may be found worldwide,...

s also act to concentrate around rubisco.

Order and kinetics


The overall process of photosynthesis takes place in four stages:
Stage Description Time scale
1 Energy transfer in antenna chlorophyll (thylakoid membranes) femtosecond to picosecond
Picosecond
A picosecond is 10−12 of a second. That is one trillionth, or one millionth of one millionth of a second, or 0.000 000 000 001 seconds. A picosecond is to one second as one second is to 31,700 years....

2 Transfer of electrons in photochemical reactions (thylakoid membranes) picosecond
Picosecond
A picosecond is 10−12 of a second. That is one trillionth, or one millionth of one millionth of a second, or 0.000 000 000 001 seconds. A picosecond is to one second as one second is to 31,700 years....

 to nanosecond
Nanosecond
A nanosecond is one billionth of a second . One nanosecond is to one second as one second is to 31.7 years.The word nanosecond is formed by the prefix nano and the unit second. Its symbol is ns....

3 Electron transport chain and ATP synthesis (thylakoid membranes) microsecond
Microsecond
A microsecond is an SI unit of time equal to one millionth of a second. Its symbol is µs.A microsecond is equal to 1000 nanoseconds or 1/1000 millisecond...

 to millisecond
Millisecond
A millisecond is a thousandth of a second.10 milliseconds are called a centisecond....

4 Carbon fixation and export of stable products millisecond
Millisecond
A millisecond is a thousandth of a second.10 milliseconds are called a centisecond....

 to second
Second
The second is a unit of measurement of time, and is the International System of Units base unit of time. It may be measured using a clock....


Efficiency



Plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s usually convert light into chemical energy
Chemical energy
Chemical energy is the potential of a chemical substance to undergo a transformation through a chemical reaction or, to transform other chemical substances...

 with a photosynthetic efficiency
Photosynthetic efficiency
The photosynthetic efficiency is the fraction of light energy converted into chemical energy during photosynthesis in plants and algae. Photosynthesis can be described by the simplified chemical reactionwhere CH2O represents carbohydrates such as sugars, cellulose, and lignin.The value of the...

 of 3–6%. 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 panel
Photovoltaic module
A solar panel is a packaged, connected assembly of solar cells, also known as photovoltaic cells...

s convert light into electric energy at an efficiency of approximately 6–20% for mass-produced panels, and above 40% in laboratory devices.

Evolution



Early photosynthetic systems, such as those from green
Green sulfur bacteria
The green sulfur bacteria are a family of obligately anaerobic photoautotrophic bacteria. Most closely related to the distant Bacteroidetes, they are accordingly assigned their own phylum....

 and purple sulfur
Purple sulfur bacteria
The purple sulfur bacteria are a group of Proteobacteria capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in hot springs or stagnant water. Unlike plants, algae, and cyanobacteria, they do not use water as their...

 and green
Chloroflexi
Chloroflexi is one of four classes of bacteria in the phylum Chloroflexi, known as filamentous green non-sulfur bacteria. They produce energy from light and are named for their green pigment, usually found in photosynthetic bodies called chlorosomes....

 and purple nonsulfur bacteria
Purple bacteria
Purple bacteria or purple photosynthetic bacteria are proteobacteria that are phototrophic, that is capable of producing energy through photosynthesis...

, are thought to have been anoxygenic, using various molecules as electron donor
Electron donor
An electron donor is a chemical entity that donates electrons to another compound. It is a reducing agent that, by virtue of its donating electrons, is itself oxidized in the process....

s. Green and purple sulfur bacteria are thought to have used hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 and sulfur
Sulfur
Sulfur or sulphur is the chemical element with atomic number 16. In the periodic table it is represented by the symbol S. It is an abundant, multivalent non-metal. Under normal conditions, sulfur atoms form cyclic octatomic molecules with chemical formula S8. Elemental sulfur is a bright yellow...

 as an electron donor. Green nonsulfur bacteria used various amino
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

 and other organic acid
Organic acid
An organic acid is an organic compound with acidic properties. The most common organic acids are the carboxylic acids, whose acidity is associated with their carboxyl group –COOH. Sulfonic acids, containing the group –SO2OH, are relatively stronger acids. The relative stability of the conjugate...

s. Purple nonsulfur bacteria used a variety of nonspecific organic molecules. The use of these molecules is consistent with the geological evidence that the atmosphere was highly reduced at that time.

Fossils of what are thought to be filamentous
Protein filament
In biology, a filament is a "long chain of proteins, such as those found in hair, muscle, or in flagella". They are often bundled together for strength and rigidity. Some cellular examples include:*Actin filaments*Microtubules*Intermediate filaments...

 photosynthetic organisms have been dated at 3.4 billion years old.

The main source of oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

 in the atmosphere
Earth's atmosphere
The atmosphere of Earth is a layer of gases surrounding the planet Earth that is retained by Earth's gravity. The atmosphere protects life on Earth by absorbing ultraviolet solar radiation, warming the surface through heat retention , and reducing temperature extremes between day and night...

 is oxygenic photosynthesis
Oxygen evolution
Oxygen evolution is the process of generating molecular oxygen through chemical reaction. Mechanisms of oxygen evolution include the oxidation of water during oxygenic photosynthesis, electrolysis of water into oxygen and hydrogen, and electrocatalytic oxygen evolution from oxides and...

, and its first appearance is sometimes referred to as the oxygen catastrophe
Oxygen Catastrophe
The Great Oxygenation Event , also called the Oxygen Catastrophe or Oxygen Crisis or Great Oxidation, was the biologically induced appearance of free oxygen in Earth's atmosphere. This major environmental change happened around 2.4 billion years ago.Photosynthesis was producing oxygen both before...

. Geological evidence suggests that oxygenic photosynthesis, such as that in cyanobacteria, became important during the Paleoproterozoic
Paleoproterozoic
The Paleoproterozoic is the first of the three sub-divisions of the Proterozoic occurring between . This is when the continents first stabilized...

 era around 2 billion years ago. Modern photosynthesis in plants and most photosynthetic prokaryotes is oxygenic. Oxygenic photosynthesis uses water as an electron donor, which is oxidized
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

 to molecular oxygen in the photosynthetic reaction center.

Symbiosis and the origin of chloroplasts


Several groups of animals have formed symbiotic
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 relationships with photosynthetic algae. These are most common in coral
Coral
Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...

s, sponges and sea anemone
Sea anemone
Sea anemones are a group of water-dwelling, predatory animals of the order Actiniaria; they are named after the anemone, a terrestrial flower. Sea anemones are classified in the phylum Cnidaria, class Anthozoa, subclass Zoantharia. Anthozoa often have large polyps that allow for digestion of larger...

s. It is presumed that this is due to the particularly simple body plan
Body plan
A body plan is the blueprint for the way the body of an organism is laid out. An organism's symmetry, its number of body segments and number of limbs are all aspects of its body plan...

s and large surface areas of these animals compared to their volumes. In addition, a few marine mollusk
Mollusca
The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest...

s Elysia viridis
Elysia viridis
Elysia viridis or the sap-sucking slug is a small-to-medium-sized species of green sea slug, a marine opisthobranch gastropod mollusc in the family Placobranchidae....

 and Elysia chlorotica
Elysia chlorotica
Elysia chlorotica, common name the eastern emerald elysia, is a small-to-medium-sized species of green sea slug, a marine opisthobranch gastropod mollusc. This sea slug superficially resembles a nudibranch, yet it does not belong to that suborder of gastropods. Instead it is a member of the...

 also maintain a symbiotic relationship with chloroplasts they capture from the algae in their diet and then store in their bodies. This allows the mollusks to survive solely by photosynthesis for several months at a time. Some of the genes from the plant cell nucleus
Cell nucleus
In cell biology, the nucleus is a membrane-enclosed organelle found in eukaryotic cells. It contains most of the cell's genetic material, organized as multiple long linear DNA molecules in complex with a large variety of proteins, such as histones, to form chromosomes. The genes within these...

 have even been transferred to the slugs, so that the chloroplasts can be supplied with proteins that they need to survive.

An even closer form of symbiosis may explain the origin of chloroplasts. Chloroplasts have many similarities with photosynthetic bacteria, including a circular chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

, prokaryotic-type ribosome
Ribosome
A ribosome is a component of cells that assembles the twenty specific amino acid molecules to form the particular protein molecule determined by the nucleotide sequence of an RNA molecule....

s, and similar proteins in the photosynthetic reaction center. The endosymbiotic theory
Endosymbiotic theory
The endosymbiotic theory concerns the mitochondria, plastids , and possibly other organelles of eukaryotic cells. According to this theory, certain organelles originated as free-living bacteria that were taken inside another cell as endosymbionts...

 suggests that photosynthetic bacteria were acquired (by endocytosis
Endocytosis
Endocytosis is a process by which cells absorb molecules by engulfing them. It is used by all cells of the body because most substances important to them are large polar molecules that cannot pass through the hydrophobic plasma or cell membrane...

) by early eukaryotic
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

 cells to form the first plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

 cells. Therefore, chloroplasts may be photosynthetic bacteria that adapted to life inside plant cells. Like mitochondria
Mitochondrion
In cell biology, a mitochondrion is a membrane-enclosed organelle found in most eukaryotic cells. These organelles range from 0.5 to 1.0 micrometers in diameter...

, chloroplasts still possess their own DNA, separate from the nuclear DNA
Nuclear DNA
Nuclear DNA, nuclear deoxyribonucleic acid , is DNA contained within a nucleus of eukaryotic organisms. In mammals and vertebrates, nuclear DNA encodes more of the genome than the mitochondrial DNA and is composed of information inherited from two parents, one male, and one female, rather than...

 of their plant host cells and the genes in this chloroplast DNA resemble those in cyanobacteria. DNA in chloroplasts codes for redox
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

 proteins such as photosynthetic reaction centers. The CoRR Hypothesis
CoRR Hypothesis
The CoRR hypothesis states that the location of genetic information in cytoplasmic organelles permits regulation of its expression by the reduction-oxidation state of its gene products....

 proposes that this Co-location is required for Redox Regulation.

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
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....

 of extant cyanobacteria. 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. Available evidence from geobiological studies of Archean
Archean
The Archean , also spelled Archeozoic or Archæozoic) is a geologic eon before the Paleoproterozoic Era of the Proterozoic Eon, before 2.5 Ga ago. Instead of being based on stratigraphy, this date is defined chronometrically...

 (>2500 Ma) sedimentary rock
Sedimentary rock
Sedimentary rock are types of rock that are formed by the deposition of material at the Earth's surface and within bodies of water. Sedimentation is the collective name for processes that cause mineral and/or organic particles to settle and accumulate or minerals to precipitate from a solution....

s indicates that life existed 3500 Ma, but the question of when oxygenic photosynthesis evolved is still unanswered. A clear paleontological window on cyanobacterial evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 opened about 2000 Ma, revealing an already-diverse biota of blue-greens. Cyanobacteria remained principal primary producers
Primary producers
Primary producers are those organisms in an ecosystem that produce biomass from inorganic compounds . In almost all cases these are photosynthetically active organisms...

 throughout the Proterozoic Eon (2500–543 Ma), in part because the redox structure of the oceans favored photoautotrophs capable of nitrogen fixation
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

. Green algae
Green algae
The green algae are the large group of algae from which the embryophytes emerged. As such, they form a paraphyletic group, although the group including both green algae and embryophytes is monophyletic...

 joined blue-greens as major primary producers on continental shelves
Continental shelf
The continental shelf is the extended perimeter of each continent and associated coastal plain. Much of the shelf was exposed during glacial periods, but is now submerged under relatively shallow seas and gulfs, and was similarly submerged during other interglacial periods. The continental margin,...

 near the end of the Proterozoic
Proterozoic
The Proterozoic is a geological eon representing a period before the first abundant complex life on Earth. The name Proterozoic comes from the Greek "earlier life"...

, but only with the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...

 (251–65 Ma) radiations of dinoflagellates, coccolithophorids, and diatoms did primary production
Primary production
400px|thumb|Global oceanic and terrestrial photoautotroph abundance, from September [[1997]] to August 2000. As an estimate of autotroph biomass, it is only a rough indicator of primary production potential, and not an actual estimate of it...

 in marine shelf waters take modern form. Cyanobacteria remain critical to marine ecosystem
Marine ecosystem
Marine ecosystems are among the largest of Earth's aquatic ecosystems. They include oceans, salt marsh and intertidal ecology, estuaries and lagoons, mangroves and coral reefs, the deep sea and the sea floor. They can be contrasted with freshwater ecosystems, which have a lower salt content. Marine...

s as primary producers in oceanic gyres, as agents of biological nitrogen fixation, and, in modified form, as the plastid
Plastid
Plastids are major organelles found in the cells of plants and algae. Plastids are the site of manufacture and storage of important chemical compounds used by the cell...

s of marine algae.

A 2010 study by researchers at Tel Aviv University
Tel Aviv University
Tel Aviv University is a public university located in Ramat Aviv, Tel Aviv, Israel. With nearly 30,000 students, TAU is Israel's largest university.-History:...

 discovered that the Oriental hornet
Oriental hornet
The Oriental hornet Vespa orientalis, is a hornet which looks very similar to the European hornet. It should not be confused with the Asian giant hornet ....

 (Vespa orientalis) converts sunlight into electric power using a pigment called xanthopterin
Xanthopterin
Xanthopterin is a yellow, crystalline solid, that occurs mainly in the wings of butterflies and in the urine of mammals. Small microorganisms convert it into folic acid. It is the end product of a non-conjugated pteridine compound, and inhibits the growth of lymphocytes produced by concanovalin...

. This is the first scientific evidence of a member of the animal kingdom engaging in photosynthesis.

Discovery


Although some of the steps in photosynthesis are still not completely understood, the overall photosynthetic equation has been known since the 19th century.

Jan van Helmont began the research of the process in the mid-17th century when he carefully measured the mass
Mass
Mass can be defined as a quantitive measure of the resistance an object has to change in its velocity.In physics, mass commonly refers to any of the following three properties of matter, which have been shown experimentally to be equivalent:...

 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 also 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
Biomass (ecology)
Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms,...

 comes from the inputs of photosynthesis, not the soil itself.

Joseph Priestley
Joseph Priestley
Joseph Priestley, FRS was an 18th-century English theologian, Dissenting clergyman, natural philosopher, chemist, educator, and political theorist who published over 150 works...

, a chemist and minister, discovered that, when he isolated a volume of air under an inverted jar, and burned a candle in it, 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.

In 1778, Jan Ingenhousz
Jan Ingenhousz
Jan Ingenhousz or Ingen-Housz FRS was a Dutch physiologist, biologist and chemist. He is best known for showing that light is essential to photosynthesis and thus having discovered photosynthesis. He also discovered that plants, like animals, have cellular respiration...

, court physician to the Austria
Austria
Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...

n Empress, 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.

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
Nicolas-Théodore de Saussure
Nicolas-Théodore de Saussure was a Swiss chemist and student of plant physiology who made seminal advances in phytochemistry....

 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.

Cornelis Van Niel made key discoveries explaining the chemistry of photosynthesis. By studying purple sulfur bacteria
Purple sulfur bacteria
The purple sulfur bacteria are a group of Proteobacteria capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in hot springs or stagnant water. Unlike plants, algae, and cyanobacteria, they do not use water as their...

 and green bacteria he was the first scientist to demonstrate that photosynthesis is a light-dependent redox
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

 reaction, in which hydrogen reduces 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 pigment. 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 absorption of quanta are equal in both the PSII and PSI, assuring that input energy from the antenna complex is divided between the PSI and PSII system, which in turn powers the photochemistry.

Robert Hill thought that a complex of reactions consisting of an intermediate to cytochrome b6 (now a plastoquinone), another is from cytochrome f to a step in the carbohydrate-generating mechanisms. These are linked by plastoquinone, which does require energy to reduce cytochrome f for it is a sufficient reductant. 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 chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...

s give off oxygen in the presence of unnatural reducing agents like iron
Iron
Iron is a chemical element with the symbol Fe and atomic number 26. It is a metal in the first transition series. It is the most common element forming the planet Earth as a whole, forming much of Earth's outer and inner core. It is the fourth most common element in the Earth's crust...

 oxalate
Oxalate
Oxalate , is the dianion with formula C2O42− also written 22−. Either name is often used for derivatives, such as disodium oxalate, 2C2O42−, or an ester of oxalic acid Oxalate (IUPAC: ethanedioate), is the dianion with formula C2O42− also written (COO)22−. Either...

, ferricyanide
Ferricyanide
Ferricyanide is the anion [Fe6]3−.  It is also called hexacyanoferrate and in rare, but systematic nomenclature, hexacyanidoferrate...

 or benzoquinone after exposure to light. The Hill reaction is as follows:
2 H2O + 2 A + (light, chloroplasts) → 2 AH2 + O2


where A is the electron acceptor. Therefore, in light, the electron acceptor is reduced and oxygen is evolved.

Samuel Ruben
Sam Ruben
Samuel Ruben , the son of Herschel and Frieda Penn Rubenstein – the name was officially shortened to Ruben in 1930...

 and Martin Kamen
Martin Kamen
Martin David Kamen , a physicist inside the Manhattan project. Together with Sam Ruben, he co-discovered the isotope carbon-14 on February 27, 1940, at the University of California Radiation Laboratory, Berkeley....

 used radioactive isotopes
Radionuclide
A radionuclide is an atom with an unstable nucleus, which is a nucleus characterized by excess energy available to be imparted either to a newly created radiation particle within the nucleus or to an atomic electron. The radionuclide, in this process, undergoes radioactive decay, and emits gamma...

 to determine that the oxygen liberated in photosynthesis came from the water.

Melvin Calvin
Melvin Calvin
Melvin Ellis Calvin was an American chemist most famed for discovering the Calvin cycle along with Andrew Benson and James Bassham, for which he was awarded the 1961 Nobel Prize in Chemistry. He spent most of his five-decade career at the University of California, Berkeley.- Life :Calvin was born...

 and Andrew Benson
Andrew Benson
Andrew Alm Benson is an American biologist and a professor of biology at the University of California, San Diego until his retirement in 1989...

, along with James Bassham
James Bassham
James Alan Bassham James Alan Bassham James Alan Bassham (born November 26, 1922 in Sacramento, California is an American scientist known for his work on photosynthesis.He received a B.S. degree in chemistry in 1945 from the University of California and his Ph.D. degree in 1949...

, elucidated the path of carbon assimilation (the photosynthetic carbon reduction cycle) in plants. The carbon reduction cycle is known as the Calvin cycle
Calvin cycle
The Calvin cycle or Calvin–Benson-Bassham cycle or reductive pentose phosphate cycle or C3 cycle or CBB cycle is a series of biochemical redox reactions that take place in the stroma of chloroplasts in photosynthetic organisms...

, which ignores the contribution of Bassham and Benson. Many scientists refer to the cycle as the Calvin-Benson Cycle, Benson-Calvin, and some even call it the Calvin-Benson-Bassham (or CBB) Cycle.

Nobel Prize
Nobel Prize
The Nobel Prizes are annual international awards bestowed by Scandinavian committees in recognition of cultural and scientific advances. The will of the Swedish chemist Alfred Nobel, the inventor of dynamite, established the prizes in 1895...

-winning scientist Rudolph A. Marcus
Rudolph A. Marcus
Rudolph "Rudy" Arthur Marcus is a Canadian-born chemist who received the 1992 Nobel Prize in Chemistry for his theory of electron transfer. Marcus theory, named after him, provides a thermodynamic and kinetic framework for describing one electron outer-sphere electron transfer.He was born in...

 was able to discover the function and significance of the electron transport chain.

Otto Heinrich Warburg
Otto Heinrich Warburg
Otto Heinrich Warburg , son of physicist Emil Warburg, was a German physiologist, medical doctor and Nobel laureate. He served as an officer in the elite Uhlan during the First World War and won the Iron Cross for bravery. Warburg was one of the twentieth century's leading biochemists...

 and Dean Burk
Dean Burk
Dean Burk was an American biochemist: a co-discoverer of biotin, medical researcher, and a cancer researcher at the Kaiser Wilhelm Institute and the National Cancer Institute...

 discovered the I-quantum photosynthesis reaction that splits the CO2, activated by the respiration.

Louis N.M. Duysens and Jan Amesz discovered that chlorophyll a will absorb one light, oxidize cytochrome f, chlorophyll a (and other pigments) will absorb another light, but will reduce this same oxidized cytochrome, stating the two light reactions are in series.

Factors



There are three main factors affecting photosynthesis and several corollary factors. The three main are:
  • Light irradiance
    Irradiance
    Irradiance is the power of electromagnetic radiation per unit area incident on a surface. Radiant emittance or radiant exitance is the power per unit area radiated by a surface. The SI units for all of these quantities are watts per square meter , while the cgs units are ergs per square centimeter...

     and wavelength
    Wavelength
    In physics, the wavelength of a sinusoidal wave is the spatial period of the wave—the distance over which the wave's shape repeats.It is usually determined by considering the distance between consecutive corresponding points of the same phase, such as crests, troughs, or zero crossings, and is a...

  • Carbon dioxide
    Carbon dioxide
    Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

     concentration
    Concentration
    In chemistry, concentration is defined as the abundance of a constituent divided by the total volume of a mixture. Four types can be distinguished: mass concentration, molar concentration, number concentration, and volume concentration...

  • Temperature
    Temperature
    Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...

    .

Light intensity (irradiance), wavelength and temperature


In the early 20th century, Frederick Frost Blackman
Frederick Blackman
Frederick Frost Blackman was a British plant physiologist.Frederick Blackman was born in Lambeth, London to a doctor. He studied medicine at St. Bartholomew's Hospital, graduating MA...

 along with Albert Einstein
Albert Einstein
Albert Einstein was a German-born theoretical physicist who developed the theory of general relativity, effecting a revolution in physics. For this achievement, Einstein is often regarded as the father of modern physics and one of the most prolific intellects in human history...

 investigated the effects of light intensity (irradiance
Irradiance
Irradiance is the power of electromagnetic radiation per unit area incident on a surface. Radiant emittance or radiant exitance is the power per unit area radiated by a surface. The SI units for all of these quantities are watts per square meter , while the cgs units are ergs per square centimeter...

) and temperature on the rate of carbon assimilation.
  • At constant temperature, the rate of carbon assimilation varies with irradiance, initially increasing as the irradiance increases. However, at higher irradiance, this relationship no longer holds and the rate of carbon assimilation reaches a plateau.
  • At constant irradiance, the rate of carbon assimilation increases as the temperature is increased over a limited range. This effect is seen only at high irradiance levels. At low irradiance, increasing the temperature has little influence on the rate of carbon assimilation.

These two experiments illustrate vital points: First, from research
Research
Research can be defined as the scientific search for knowledge, or as any systematic investigation, to establish novel facts, solve new or existing problems, prove new ideas, or develop new theories, usually using a scientific method...

 it is known that, in general, photochemical reactions are not affected by temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...

. 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, of course, the light-dependent 'photochemical' stage and the light-independent, temperature-dependent
Light-independent reaction
The light-independent reactions of photosynthesis are chemical reactions that convert carbon dioxide and other compounds into glucose. These reactions occur in the stroma, the fluid-filled area of a chloroplast outside of the thylakoid membranes. These reactions take the light-dependent reactions...

 stage. Second, Blackman's experiments illustrate the concept of limiting factor
Limiting factor
A limiting factor or limiting resource is a factor that controls a process, such as organism growth or species population, size, or distribution. The availability of food, predation pressure, or availability of shelter are examples of factors that could be limiting for an organism...

s. 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
Phycobilisome
Phycobilisomes are light harvesting antennae of photosystem II in cyanobacteria, red algae and glaucophytes.- General structure :Phycobilisomes are protein complexes anchored to thylakoid membranes. They are made of stacks of chromophorylated proteins, the phycobiliproteins, and their associated...

.

Carbon dioxide levels and photorespiration


As carbon dioxide concentrations rise, the rate at which sugars are made by the light-independent reaction
Light-independent reaction
The light-independent reactions of photosynthesis are chemical reactions that convert carbon dioxide and other compounds into glucose. These reactions occur in the stroma, the fluid-filled area of a chloroplast outside of the thylakoid membranes. These reactions take the light-dependent reactions...

s increases until limited by other factors. RuBisCO
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...

, 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
Carbon fixation
In biology, carbon fixation is the reduction of carbon dioxide to organic compounds by living organisms. The obvious example is photosynthesis. Carbon fixation requires both a source of energy such as sunlight, and an electron donor such as water. All life depends on fixed carbon. Organisms that...

. However, if the carbon dioxide concentration is low, RuBisCO will bind oxygen instead of carbon dioxide. This process, called photorespiration
Photorespiration
Photorespiration, or "'photo-respiration'", is a process in plant metabolism by which RuBP has oxygen added to it by the enzyme , instead of carbon dioxide during normal photosynthesis. This is the beginning step of the Calvin-Benson cycle...

, uses energy, but does not produce sugars.

RuBisCO oxygenase activity is disadvantageous to plants for several reasons:
  1. 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.
  2. Phosphoglycolate is quickly metabolized to glycolate that is toxic to a plant at a high concentration; it inhibits photosynthesis.
  3. 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
    Ammonia
    Ammonia is a compound of nitrogen and hydrogen with the formula . It is a colourless gas with a characteristic pungent odour. Ammonia contributes significantly to the nutritional needs of terrestrial organisms by serving as a precursor to food and fertilizers. Ammonia, either directly or...

     (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
Photorespiration
Photorespiration, or "'photo-respiration'", is a process in plant metabolism by which RuBP has oxygen added to it by the enzyme , instead of carbon dioxide during normal photosynthesis. This is the beginning step of the Calvin-Benson cycle...

, since it is characterized by light-dependent oxygen consumption and the release of carbon dioxide.

See also



  • Jan Anderson (scientist)
    Jan Anderson (scientist)
    Jan Mary Anderson FRS is a scientist from New Zealand, distinguished by her investigation of photosynthesis. She was the first to show that the photosynthetic mechanism comprises two fundamental components: photosystem I and photosytem II....

  • Artificial photosynthesis
    Artificial photosynthesis
    Artificial photosynthesis is a chemical process that replicates the natural process of photosynthesis, a process that converts sunlight, water, and carbon dioxide into carbohydrates and oxygen. The term is commonly used to refer to any scheme for capturing and storing the energy from sunlight in...

  • Calvin-Benson cycle
  • Carbon fixation
    Carbon fixation
    In biology, carbon fixation is the reduction of carbon dioxide to organic compounds by living organisms. The obvious example is photosynthesis. Carbon fixation requires both a source of energy such as sunlight, and an electron donor such as water. All life depends on fixed carbon. Organisms that...

  • Cellular respiration
    Cellular respiration
    Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

  • Chemosynthesis
    Chemosynthesis
    In biochemistry, chemosynthesis is the biological conversion of one or more carbon molecules and nutrients into organic matter using the oxidation of inorganic molecules or methane as a source of energy, rather than sunlight, as in photosynthesis...

  • Light-dependent reaction
  • Photobiology
    Photobiology
    Photobiology is the scientific study of the interactions of light and living organisms. The field includes the study of photosynthesis, photomorphogenesis, visual processing, circadian rhythms, bioluminescence, and ultraviolet radiation effects...

  • Photoinhibition
    Photoinhibition
    Photoinhibition is light-induced reduction in the photosynthetic capacity of a plant, alga, or cyanobacterium. Photosystem II is more sensitive to light than the rest of the photosynthetic machinery, and most researchers define the term as light-induced damage to PSII...

  • Photosystem
    Photosystem
    Photosystems are functional and structural units of protein complexes involved in photosynthesis that together carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons...

  • Photosystem I
    Photosystem I
    Photosystem I is the second photosystem in the photosynthetic light reactions of algae, plants, and some bacteria. Photosystem I is so named because it was discovered before photosystem II. Aspects of PS I were discovered in the 1950s, but the significances of these discoveries was not yet known...

  • Photosystem II
    Photosystem II
    Photosystem II is the first protein complex in the Light-dependent reactions. It is located in the thylakoid membrane of plants, algae, and cyanobacteria. The enzyme uses photons of light to energize electrons that are then transferred through a variety of coenzymes and cofactors to reduce...

  • Photosynthetic reaction center
  • Photosynthetically active radiation
    Photosynthetically active radiation
    Photosynthetically active radiation, often abbreviated PAR, designates the spectral range of solar radiation from 400 to 700 nanometers that photosynthetic organisms are able to use in the process of photosynthesis. This spectral region corresponds more or less with the range of light visible to...

  • Quantum biology
    Quantum biology
    Quantum biology refers to applications of quantum mechanics to biological objects and problems. Usually, it is taken to refer to applications of the "non-trivial" quantum features such as superposition, nonlocality, entanglement and tunneling, as opposed to the "trivial" applications such as...

  • Red edge
    Red edge
    Red edge refers to the region of rapid change in reflectance of chlorophyll in the near infrared range. Vegetation absorbs most of the light in the visible part of the spectrum but is strongly reflective at wavelengths greater than 700 nm. The change can be from 5% to 50% reflectance between 680 nm...

  • Vitamin D
    Vitamin D
    Vitamin D is a group of fat-soluble secosteroids. In humans, vitamin D is unique both because it functions as a prohormone and because the body can synthesize it when sun exposure is adequate ....



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