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Schematic of photosynthesis in plants. The carbohydrates produced are stored in or used by the plant.
Overall equation for the type of photosynthesis that occurs in plants
Composite image showing the global distribution of photosynthesis, including both oceanic
and terrestrial . Dark red and blue-green indicate regions of high photosynthetic activity in ocean and land respectively.
Photosynthesis is a process used by plants and other organisms to convert
energy, normally from the , into
that can be later
to fuel the organisms' activities (). This chemical energy is stored in , such as , which are synthesized from
– hence the name photosynthesis, from the
, phōs, "light", and , synthesis, "putting together". In most cases, oxygen is also released as a waste product. Most , most , and
pe such organisms are called . Photosynthesis maintains
levels and supplies all of the organic compounds and most of the energy necessary for .
Although photosynthesis is performed differently by different species, the process always begins when energy from light is absorbed by
that contain green
pigments. In plants, these proteins are held inside
called , which are most abundant in leaf cells, while in bacteria they are embedded in the . In these light-dependent reactions, some energy is used to strip
from suitable substances, such as water, producing oxygen gas. The hydrogen freed by water splitting is used in the creation of two further compounds: reduced
(NADPH) and
(ATP), the "energy currency" of cells.
In plants, algae and cyanobacteria, sugars are produced by a subsequent sequence of light-independent reactions called the , but some bacteria use different mechanisms, such as the . In the Calvin cycle, atmospheric carbon dioxide is
into already existing organic carbon compounds, such as
(RuBP). Using the ATP and NADPH produced by the light-dependent reactions, the resulting compounds are then
and removed to form further carbohydrates, such as .
The first photosynthetic organisms probably
early in the
and most likely used
or , rather than water, as sources of electrons. Cyanobac the excess oxygen they produced contributed to the , which rendered the
possible. Today, the average rate of energy capture by photosynthesis globally is approximately 130 , which is about three times the current . Photosynthetic organisms also convert around 100–115 thousand million metric tonnes of carbon into
Photosynthesis changes sunlight into chemical energy, splits water to liberate O2, and fixes CO2 into sugar.
Photosynthetic organisms are , which means that they are able to
food directly from carbon dioxide and water using energy from light. However, not all organisms that use light as a source of energy carry out photosynthesis, since
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 , , and , the overall process is quite similar in these organisms. However, there are some types of bacteria that carry out . These consume carbon dioxide but do not release oxygen.
Carbon dioxide is converted into sugars in a process called . Carbon fixation is an
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
via a reduction reaction. The addition of electrons to a chemical species is called a . In general outline and in effect, photosynthesis is the opposite of , in which glucose and other compounds are oxidized to produce carbon dioxide and water, and to release chemical energy (an
reaction) to drive the organism's . The two processes, of reduction of carbon dioxide to carbohydrate and then the later oxidation of the carbohydrate, take place through a different sequence of chemical reactions and in different cellular compartments.
The general
for photosynthesis as first proposed by
is therefore:
CO2 + 2H2A +
→ [] + 2A + H2O
carbon dioxide + electron donor + light energy → carbohydrate + oxidized electron donor + water
Since water is used as the electron donor in oxygenic photosynthesis, the equation for this process is:
CO2 + 2H2O + photons → [CH2O] + O2 + H2O
carbon dioxide + water + light energy → carbohydrate + oxygen + water
This equation emphasizes that water is both a reactant in the
and a product of the , but canceling n water molecules from each side gives the net equation:
CO2 + H2O + photons → [CH2O] + O2
carbon dioxide + water + light energy → carbohydrate + oxygen
Other processes substitute other compounds (such as ) for water in the electron- for example some microbes use sunlight to oxidize arsenite to : 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
and . 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
to do so, although at least three use shortwave
or, more specifically, far-red radiation.
use a simpler method using a pigment similar to the pigments used for vision. The
changes its configuration in response to sunlight, acting as a proton pump. This produces a proton gradient more directly which is then converted to chemical energy. The process does not involve carbon dioxide fixation and does not release oxygen. It seems to have evolved separately.
Chloroplast ultrastructure:
1. outer membrane
2. intermembrane space
3. inner membrane (1+2+3: envelope)
4. stroma (aqueous fluid)
5. thylakoid lumen (inside of thylakoid)
6. thylakoid membrane
7. granum (stack of thylakoids)
8. thylakoid (lamella)
10. ribosome
11. plastidial DNA
12. plastoglobule (drop of lipids)
Main articles:
In photosynthetic bacteria, the proteins that gather light for photosynthesis are embedded in . In its simplest form, this involves the membrane surrounding the cell itself. However, the membrane may be tightly folded into cylindrical sheets called , or bunched up into round
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
called . A typical
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. Enclosed by the membrane is an aqueous fluid called the stroma. Embedded within the stroma are stacks of thylakoids (grana), which are the site of photosynthesis. The thylakoids appear as flattened disks. The thylakoid itself is enclosed by the thylakoid membrane, and within the enclosed volume is a lumen or thylakoid space. Embedded in the thylakoid membrane are integral and
complexes of the photosynthetic system, including the pigments that absorb light energy.
Plants absorb light primarily using the
. The green part of the light spectrum is not absorbed but is reflected which is the reason that most plants have a green color. Besides chlorophyll, plants also use pigments such as
and . Algae also use chlorophyll, but various other pigments are present, such as , , and
(rhodophytes) and
resulting in a wide variety of colors.
These pigments are embedded in plants and algae in complexes called antenna proteins. In such proteins, the pigments are arranged to work together. Such a combination of proteins is also called a .
Although all cells in the green parts of a plant have chloroplasts, the majority of those are found in specially adapted structures called . Certain species adapted to conditions of strong sunlight and , such as many
species, have their main photosynthetic organs in their stems. The cells in the interior tissues of a leaf, called the , can contain between 450,000 and 800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is coated with a water-resistant
that protects the leaf from excessive
of water and decreases the absorption of
to reduce . The transparent
layer allows light to pass through to the
mesophyll cells where most of the photosynthesis takes place.
Light-dependent reactions of photosynthesis at the thylakoid membrane
Main article:
In the , one molecule of the
absorbs one
and loses one . This electron is passed to a modified form of chlorophyll called , which passes the electron to a
molecule, starting the flow of electrons down an
that leads to the ultimate reduction of
to . In addition, this creates a
(energy gradient) across the , which is used by
in the synthesis of . The chlorophyll molecule ultimately regains the electron it lost when a water molecule is split in a process called , which releases a
(O2) molecule as a waste product.
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
of light can support photosynthesis. The photosynthetic action spectrum depends on the type of
present. For example, in green plants, the
resembles the
with peaks for violet-blue and red light. In red algae, the action spectrum is blue-green light, which allows these algae to use the blue end of the spectrum to grow in the deeper waters that filter out the longer wavelengths (red light) used by above ground green plants. The non-absorbed part of the
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.
The "Z scheme"
In plants,
occur in the
where they drive the synthesis of ATP and NADPH. The light-dependent reactions are of two forms: cyclic and non-cyclic.
In the non-cyclic reaction, the
are captured in the light-harvesting
(see diagram at right). The absorption of a photon by the antenna complex frees an electron by a process called . The antenna system is at the core of the chlorophyll molecule of the photosystem II reaction center. That freed electron is transferred to the primary electron-acceptor molecule, pheophytin. As the electrons are shuttled through an
(the so-called Z-scheme shown in the diagram), it initially functions to generate a
by pumping proton cations (H+) across the membrane and into the thylakoid space. An
enzyme uses that chemiosmotic potential to make ATP during photophosphorylation, whereas
is a product of the terminal
reaction in the Z-scheme. The electron enters a chlorophyll molecule in . There it is further excited by the light absorbed by that . The electron is then passed along a chain of
to which it transfers some of its energy. The energy delivered to the electron acceptors is used to move hydrogen ions across the thylakoid membrane into the lumen. The electron is eventually used to reduce the co-enzyme NADP with a H+ to NADPH (which has functions in the light-independent reaction); at that point, the path of that electron ends.
The cyclic reaction is similar to that of the non-cyclic, but differs in that it generates only ATP, and no reduced NADP (NADPH) is created. The cyclic reaction takes place only at photosystem I. Once the electron is displaced from the photosystem, the electron is passed down the electron acceptor molecules and returns to photosystem I, from where it was emitted, hence the name cyclic reaction.
Main articles:
The NADPH is the main
produced by chloroplasts, which then goes on to provide a source of energetic electrons in other cellular reactions. Its production leaves chlorophyll in photosystem I with a deficit of electrons (chlorophyll has been oxidized), which must be balanced by some other reducing agent that will supply the missing electron. The excited electrons lost from chlorophyll from
are supplied from the electron transport chain by . However, since
is the first step 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 diat the electrons yielded are transferred to a redox-active
residue that then reduces the oxidized chlorophyll a (called P680) that serves as the primary light-driven electron donor in the photosystem II reaction center. That photo receptor is in effect reset and is then able to repeat the absorption of another photon and the release of another photo-dissociated electron. The oxidation of water is
in photosystem II by a redox-active structure that contains four
binds two water molecules and contains the four oxidizing equivalents that are used to drive the water-oxidizing reaction. Photosystem II is the only known biological
that carries out this oxidation of water. The hydrogen ions released 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 , including photosynthetic organisms.
Main articles: , , and
(or "dark") reactions, the
and, in a process called the , it uses the newly formed NADPH and releases three-carbon sugars, which are later combined to form sucrose and starch. The overall equation for the light-independent reactions in green plants is:128
3 CO2 + 9 ATP + 6 NADPH + 6 H+ → C3H6O3-phosphate + 9 ADP + 8 Pi + 6 NADP+ + 3 H2O
Overview of the Calvin cycle and carbon fixation
Carbon fixation produces the intermediate three-carbon sugar product, which is then converted to the final carbohydrate products. The simple carbon sugars produced by photosynthesis are then used in the forming of other organic compounds, such as the building material , the precursors for
biosynthesis, or as a fuel in . The latter occurs not only in plants but also in
when the energy from plants is passed through a .
The fixation or reduction of carbon dioxide is a process in which
combines with a five-carbon sugar, , to yield two molecules of a three-carbon compound, , also known as 3-phosphoglycerate. Glycerate 3-phosphate, in the presence of
produced during the light-dependent stages, is reduced to . This product is also referred to as 3-phosphoglyceraldehyde () or, more generically, as
phosphate. Most (5 out of 6 molecules) of the glyceraldehyde 3-phosphate produced is used to regenerate ribulose 1,5-bisphosphate so the process can continue. The triose phosphates not thus "recycled" often condense to form
phosphates, which ultimately yield ,
and . The sugars produced during carbon
yield carbon skeletons that can be used for other metabolic reactions like the production of
Overview of
In hot and dry conditions, plants close their
to prevent water loss. Under these conditions, CO2 will decrease and oxygen gas, produced by the light reactions of photosynthesis, will increase, causing an increase of
activity of
and decrease in carbon fixation. Some plants have
mechanisms to increase the CO2 concentration in the leaves under these conditions.
Main article:
Plants that use the
carbon fixation process chemically fix carbon dioxide in the cells of the mesophyll by adding it to the three-carbon molecule , a reaction catalyzed by an enzyme called , creating the four-carbon organic acid . Oxaloacetic acid or
synthesized by this process is then translocated to specialized
cells where the enzyme
and other Calvin cycle enzymes are located, and where CO2 released by
of the four-carbon acids is then fixed by RuBisCO activity to the three-carbon . The physical separation of RuBisCO from the oxygen-generating light reactions reduces photorespiration and increases CO2 fixation and, thus, the
of the leaf. C4 plants can produce more sugar than C3 plants in conditions of high light and temperature. Many important crop plants are C4 plants, including maize, sorghum, sugarcane, and millet. Plants that do not use PEP-carboxylase in carbon fixation are called
because the primary carboxylation reaction, catalyzed by RuBisCO, produces the three-carbon 3-phosphoglyceric acids directly in the Calvin-Benson cycle. Over 90% of plants use C3 carbon fixation, compared to 3% that use C4 however, the evolution of C4 in over 60 plant lineages makes it a striking example of .
Main article:
and most , also use PEP carboxylase to capture carbon dioxide in a process called
(CAM). In contrast to C4 metabolism, which physically separates the CO2 fixation to PEP from the Calvin cycle, CAM temporally separates these two processes. CAM plants have a different leaf anatomy from C3 plants, and fix the CO2 at night, when their stomata are open. CAM plants store the CO2 mostly in the form of
via carboxylation of
to oxaloacetate, which is then reduced to malate. Decarboxylation of malate during the day releases CO2 inside the leaves, thus allowing carbon fixation to 3-phosphoglycerate by RuBisCO. Sixteen thousand species of plants use CAM.
possess , which increase the concentration of CO2 around RuBisCO to increase the rate of photosynthesis. An enzyme, , located within the carboxysome releases CO2 from the dissolved hydrocarbonate ions (HCO3-). Before the CO2 diffuses out it is quickly sponged up by RuBisCO, which is concentrated within the carboxysomes. HCO3- ions are made from CO2 outside the cell by another carbonic anhydrase and are actively pumped into the cell by a membrane protein. They cannot cross the membrane as they are charged, and within the cytosol they turn back into CO2 very slowly without the help of carbonic anhydrase. This causes the HCO3- ions to accumulate within the cell from where they diffuse into the carboxysomes.
also act to concentrate CO2 around rubisco.
The overall process of photosynthesis takes place in four stages:
Description
Time scale
Energy transfer in antenna chlorophyll (thylakoid membranes)
Transfer of electrons in photochemical reactions (thylakoid membranes)
Electron transport chain and ATP synthesis (thylakoid membranes)
Carbon fixation and export of stable products
Probability distribution resulting from one-dimensional discrete time random walks. The quantum walk created using the Hadamard coin is plotted (blue) vs a classical walk (red) after 50 time steps.
Main article:
usually convert light into
of 3–6%. Absorbed light that is unconverted is dissipated primarily as heat, with a small fraction (1–2%) re-emitted as
at longer (redder) wavelengths.
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,
convert light into
at an efficiency of approximately 6–20% for mass-produced panels, and above 40% in laboratory devices.
Photosynthesis measurement systems are not designed to directly measure the amount of light absorbed by the leaf. But analysis of chlorophyll-fluorescence, P700- and P515-absorbance and gas exchange measurements reveal detailed information about e.g. the photosystems, quantum efficiency and the CO2 assimilation rates. With some instruments even wavelength-dependency of the photosynthetic efficiency can be analyzed.
A phenomenon known as
increases the efficiency of the energy transport of light significantly. In the photosynthetic cell of an algae, bacterium, or plant, there are light-sensitive molecules called
arranged in an antenna-shaped structure named a photocomplex. When a photon is absorbed by a chromophore, it is converted into a
referred to as an , which jumps from chromophore to chromophore towards the reaction center of the photocomplex, a collection of molecules that traps its energy in a chemical form that makes it accessible for the cell's metabolism. The exciton's wave properties enable it to cover a wider area and try out several possible paths simultaneously, allowing it to instantaneously "choose" the most efficient route, where it will have the highest probability of arriving at its destination in the minimum possible time. Because that quantum walking takes place at temperatures far higher than quantum phenomena usually occur, it is only possible over very short distances, due to obstacles in the form of destructive interference that come into play. These obstacles cause the particle to lose its wave properties for an instant before it regains them once again after it is freed from its locked position through a classic "hop". The movement of the electron towards the photo center is therefore covered in a series of conventional hops and quantum walks.
-500 —
also see {} and {}
Main article:
Early photosynthetic systems, such as those in
and , are thought to have been anoxygenic, and used various other molecules as
rather than water. Green and purple sulfur bacteria are thought to have used
as electron donors. Green nonsulfur bacteria used various
as an electron donor. Purple nonsulfur bacteria used a variety of nonspecific organic molecules. The use of these molecules is consistent with the geological evidence that Earth's early atmosphere was highly
Fossils of what are thought to be
photosynthetic organisms have been dated at 3.4 billion years old.
The main source of
derives from , and its first appearance is sometimes referred to as the . Geological evidence suggests that oxygenic photosynthesis, such as that in , became important during the
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
to molecular oxygen (O
2) in the .
Plant cells with visible chloroplasts (from a moss, )
Several groups of animals have formed
relationships with photosynthetic algae. These are most common in ,
and . It is presumed that this is due to the particularly simple
and large surface areas of these animals compared to their volumes. In addition, a few marine
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
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 , including a circular , prokaryotic-type , and similar proteins in the photosynthetic reaction center. The
suggests that photosynthetic bacteria were acquired (by ) by early
cells to form the first
cells. Therefore, chloroplasts may be photosynthetic bacteria that adapted to life inside plant cells. Like , chloroplasts possess their own DNA, separate from the
of their plant host cells and the genes in this chloroplast DNA resemble those found in . DNA in chloroplasts codes for
proteins such as those found in the photosynthetic reaction centers. The
proposes that this Co-location is required for Redox Regulation.[]
The biochemical capacity to use water as the source for electrons in photosynthesis evolved once, in a
of extant . The geological record indicates that this transforming event took place early in Earth's history, at least
million years ago (Ma), and, it is speculated, much earlier. Because the Earth's atmosphere contained almost no oxygen during the estimated development of photosynthesis, it is believed that the first photosynthetic cyanobacteria did not generate oxygen. Available evidence from geobiological studies of
(&2500 Ma)
indicates that life existed 3500 Ma, but the question of when oxygenic photosynthesis evolved is still unanswered. A clear paleontological window on cyanobacterial
opened about 2000 Ma, revealing an already-diverse biota of blue-green algae.
remained the principal
of oxygen throughout the
( Ma), in part because the redox structure of the oceans favored photoautotrophs capable of .[]
joined blue-green algae as the major primary producers of oxygen on
near the end of the , but it was only with the
(251–65 Ma) radiations of dinoflagellates, coccolithophorids, and diatoms did the
of oxygen in marine shelf waters take modern form. Cyanobacteria remain critical to
as primary producers of oxygen in oceanic gyres, as agents of biological nitrogen fixation, and, in modified form, as the
of marine algae.
(Vespa orientalis) converts sunlight into electric power using a pigment called . This is the first evidence of a member of the animal kingdom engaging in photosynthesis.
Although some of the steps in photosynthesis are still not completely understood, the overall photosynthetic equation has been known since the 19th century.
began the research of the process in the mid-17th century when he carefully measured the
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
comes from the inputs of photosynthesis, not the soil itself.
, 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, , 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, , a Swiss pastor, botanist, and naturalist, demonstrated that green plants consume carbon dioxide and release oxygen under the influence of light. Soon afterward,
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.
made key discoveries explaining the chemistry of photosynthesis. By studying
and green bacteria he was the first to demonstrate that photosynthesis is a light-dependent
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 the 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.
Melvin Calvin works in his photosynthesis laboratory.
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
give off oxygen in the presence of unnatural reducing agents like
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.
to determine that the oxygen liberated in photosynthesis came from the water.
and , along with , elucidated the path of carbon assimilation (the photosynthetic carbon reduction cycle) in plants. The carbon reduction cycle is known as the , 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.
-winning scientist
was able to discover the function and significance of the electron transport chain.
discovered the I-quantum photosynthesis reaction that splits the CO2, activated by the respiration.
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.
proposed two terms, photosyntax and photosynthesis, for the biological process of synthesis of complex carbon compounds out of carbonic acid, in the presence of chlorophyll, under the influence of light. Over time, the term photosynthesis came into common usage as the term of choice. Later discovery of anoxygenic photosynthetic bacteria and photophosphorylation necessitated redefinition of the term.
is the primary site of photosynthesis in plants.
There are three main factors affecting photosynthesis and several corollary factors. The three main are:
spectra of free chlorophyll a (green) and b (red) in a solvent. The action spectra of chlorophyll molecules are slightly modified in vivo depending on specific pigment-protein interactions.
The process of photosynthesis provides the main input of free energy into the biosphere, and is one of four main ways in which radiation is important for plant life.
The radiation climate within plant communities is extremely variable, with both time and space.
In the early 20th century,
investigated the effects of light intensity () and temperature on the rate of carbon assimilation.
At constant temperature, the rate of carbon assimilation varies with irradiance, increasing as the irradiance increases, but reaching a plateau at higher irradiance.
At low irradiance, increasing the temperature has little influence on the rate of carbon assimilation. At constant high irradiance, the rate of carbon assimilation increases as the temperature is increased.
These two experiments illustrate several important points: First, it is known that, in general,
reactions are not affected by . 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
temperature-independent stage, and the
stage. Second, Blackman's experiments illustrate the concept of . 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 .[]
Photorespiration
As carbon dioxide concentrations rise, the rate at which sugars are made by the
increases until limited by other factors. , 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 . However, if the carbon dioxide concentration is low, RuBisCO will bind oxygen instead of carbon dioxide. This process, called , uses energy, but does not produce sugars.
RuBisCO oxygenase activity is disadvantageous to plants for several reasons:
One product of oxygenase activity is phosphoglycolate (2 carbon) instead of
(3 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 .
Phosphoglycolate is quickly metabolized to glycolate that is toxic to a plant at
it inhibits photosynthesis.
Salvaging glycolate is an energetically expensive process that uses the glycolate pathway, and only 75% of the carbon is returned to the Calvin-Benson cycle as 3-phosphoglycerate. The reactions also produce
(NH3), which is able to
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 , since it is characterized by light-dependent oxygen consumption and the release of carbon dioxide.
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