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Contents

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Overview

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Photosynthetic membranes and organelles

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Light-dependent reactions
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Light-independent reactions
 Toggle Light-independent reactions subsection
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Order and kinetics

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Efficiency

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Evolution
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Experimental history
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Factors
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See also

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References

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Photosynthesis
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From Wikipedia, the free encyclopedia

Schematic of photosynthesis in
plants. The carbohydrates produced are stored in or used by the plant.

Composite image showing the global

distribution of photosynthesis, including both oceanic phytoplankton and
terrestrial vegetation. Dark red and blue-green indicate regions of high
photosynthetic activity in the ocean and on land, respectively.
Photosynthesis (/ˌfoʊtəˈsɪnθəsɪs/ FOH-tə-SINTH-ə-sis)[1] is a system of biological
processes by which photosynthetic organisms, such as most plants, algae,
and cyanobacteria, convert light energy, typically from sunlight, into the chemical
energy necessary to fuel their activities. Photosynthetic organisms use
intracellular organic compounds to store the chemical energy they produce in
photosynthesis within organic compounds
like sugars, glycogen, cellulose and starches. Photosynthesis is usually used to refer
to oxygenic photosynthesis, a process that produces oxygen. To use this stored
chemical energy, the organisms' cells metabolize the organic compounds through
another process called cellular respiration. Photosynthesis plays a critical role in
producing and maintaining the oxygen content of the Earth's atmosphere, and it
supplies most of the biological energy necessary for complex life on Earth.[2]
Some bacteria also perform anoxygenic photosynthesis, which
uses bacteriochlorophyll to split hydrogen sulfide as a reductant instead of water.
This process produces sulfur instead of oxygen. Archaea such
as Halobacterium also perform a type of non-carbon-fixing anoxygenic
photosynthesis, where the simpler photopigment retinal and its microbial
rhodopsin derivatives are used to absorb green light and power proton pumps to
directly synthesize adenosine triphosphate (ATP), the "energy currency" of cells.
Such archaeal photosynthesis might have been the earliest form of photosynthesis
that evolved on Earth, going back as far as the Paleoarchean, preceding that of
cyanobacteria (see Purple Earth hypothesis).
While the details may differ between species, the process always begins when light
energy is absorbed by the reaction centers, proteins that contain photosynthetic
pigments or chromophores. In plants, these proteins
are chlorophylls (a porphyrin derivative that absorbs the red and
blue spectrums of light, thus reflecting a green color) held inside chloroplasts,
abundant in leaf cells. In bacteria they are embedded in the plasma membrane. In
these light-dependent reactions, some energy is used to strip electrons from suitable
substances, such as water, producing oxygen gas. The hydrogen freed by the
splitting of water is used in the creation of two important molecules that participate in
energetic processes: reduced nicotinamide adenine dinucleotide
phosphate (NADPH) and ATP.
In plants, algae, and cyanobacteria, sugars are synthesized by a subsequent
sequence of light-independent reactions called the Calvin cycle. In this process,
atmospheric carbon dioxide is incorporated into already existing organic carbon
compounds, such as ribulose bisphosphate (RuBP).[3] Using the ATP and NADPH
produced by the light-dependent reactions, the resulting compounds are
then reduced and removed to form further carbohydrates, such as glucose. In other
bacteria, different mechanisms like the reverse Krebs cycle are used to achieve the
same end.
The first photosynthetic organisms probably evolved early in the evolutionary history
of life and most likely used reducing agents such as hydrogen or hydrogen sulfide,
rather than water, as sources of electrons.[4] Cyanobacteria appeared later;
the excess oxygen they produced contributed directly to the oxygenation of the
Earth,[5] which rendered the evolution of complex life possible. Today, the average
rate of energy captured by photosynthesis globally is approximately 130 terawatts,[6][7]
[8]
which is about eight times the current power consumption of human civilization.
[9]
Photosynthetic organisms also convert around 100–115 billion tons (91–104
Pg petagrams, or a billion metric tons), of carbon into biomass per year.[10]
[11]
Photosyntesis was first discovered in 1779 by Jan Ingenhousz; he showed that
plants need light, not just air, soil, and water.
Photosynthesis is vital for climate processes, as it captures carbon dioxide from the
air and then binds it in plants, harvested products and soil. Cereals alone are
estimated to bind 3,825 Tg (teragrams) or 3.825 Pg (petagrams) of carbon dioxide
every year, i.e. 3.825 billion metric tons.[12]
Overview
Main article: Biological carbon fixation

Photosynthesis changes sunlight into chemical

energy, splits water to liberate O , and fixes CO into sugar.
2 2

Most photosynthetic organisms are photoautotrophs, which means that they are able
to synthesize food directly from carbon dioxide and water using energy from light.
However, not all organisms use carbon dioxide as a source of carbon atoms to carry
out photosynthesis; photoheterotrophs use organic compounds, rather than carbon
dioxide, as a source of carbon.[2]
In plants, algae, and cyanobacteria, photosynthesis releases oxygen. This oxygenic
photosynthesis is by far the most common type of photosynthesis used by living
organisms. Some shade-loving plants (sciophytes) produce such low levels of
oxygen during photosynthesis that they use all of it themselves instead of releasing it
to the atmosphere.[13]
Although there are some differences between oxygenic photosynthesis
in plants, algae, and cyanobacteria, the overall process is quite similar in these
organisms. There are also many varieties of anoxygenic photosynthesis, used
mostly by bacteria, which consume carbon dioxide but do not release oxygen.[citation needed]
Carbon dioxide is converted into sugars in a process called carbon fixation;
photosynthesis captures energy from sunlight to convert carbon dioxide
into carbohydrates. Carbon fixation is an endothermic redox reaction. In general
outline, photosynthesis is the opposite of cellular respiration: while photosynthesis is
a process of reduction of carbon dioxide to carbohydrates, cellular respiration is the
oxidation of carbohydrates or other nutrients to carbon dioxide. Nutrients used in
cellular respiration include carbohydrates, amino acids and fatty acids. These
nutrients are oxidized to produce carbon dioxide and water, and to release chemical
energy to drive the organism's metabolism.
Photosynthesis and cellular respiration are distinct processes, as they take place
through different sequences of chemical reactions and in different cellular
compartments.[citation needed]
The general equation for photosynthesis as first proposed by Cornelis van Niel is:[14]
CO2carbon
dioxide + 2H2Aelectron donor + photonslight energy → [CH2O]carbohydrate + 2Aoxidized
 electron
donor + H2Owater
Since water is used as the electron donor in oxygenic photosynthesis, the
equation for this process is:
CO2carbon
dioxide + 2H2Owater + photonslight energy → [CH2O]carbohydrate + O2oxygen + H2Owater

This equation emphasizes that water is both a reactant in the light-dependent

reaction and a product of the light-independent reaction, but
canceling n water molecules from each side gives the net equation:
CO2carbon
dioxide + H2Owater + photonslight energy → [CH2O]carbohydrate + O2oxygen

Other processes substitute other compounds (such as arsenite) for water

in the electron-supply role; for example some microbes use sunlight to
oxidize arsenite to arsenate:[15] The equation for this reaction is:
CO2carbon
dioxide + (AsO3−
3)

arsenite + photonslight energy → (AsO3−

4)

arsenate + COcarbon
monoxide(used to build other compounds in subsequent reactions) [16]

Photosynthesis occurs in two stages. In the first stage, light-

dependent reactions or light reactions capture the energy of light and
use it to make the hydrogen carrier NADPH and the energy-storage
molecule ATP. During the second stage, the light-independent
reactions use these products to capture and reduce carbon dioxide.
Most organisms that use oxygenic photosynthesis use visible light for
the light-dependent reactions, although at least three use
shortwave infrared or, more specifically, far-red radiation.[17]
Some organisms employ even more radical variants of
photosynthesis. Some archaea use a simpler method that employs a
pigment similar to those used for vision in animals.
The bacteriorhodopsin changes its configuration in response to
sunlight, acting as a proton pump. This produces a proton gradient
more directly, which is then converted to chemical energy. The
process does not involve carbon dioxide fixation and does not release
oxygen, and seems to have evolved separately from the more
common types of photosynthesis.[18]
Photosynthetic membranes and organelles
Main articles: Chloroplast and Thylakoid
 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)
9. starch
10. ribosome
11. plastidial DNA
12. plastoglobule (drop of lipids)
In photosynthetic bacteria, the proteins that gather light for
photosynthesis are embedded in cell membranes. In its simplest form,
this involves the membrane surrounding the cell itself.[19] However, the
membrane may be tightly folded into cylindrical sheets
called thylakoids,[20] or bunched up into
round vesicles called intracytoplasmic membranes.[21] 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.[20]
In plants and algae, photosynthesis takes place
in organelles called chloroplasts. A typical plant cell contains about 10
to 100 chloroplasts. The chloroplast is enclosed by a membrane. This
membrane is composed of a phospholipid inner membrane, a
phospholipid outer membrane, and an intermembrane space.
Enclosed by the membrane is an aqueous fluid called the stroma.
Embedded within the stroma are stacks of thylakoids (grana), which
are the site of photosynthesis. The thylakoids appear as flattened
disks. The thylakoid itself is enclosed by the thylakoid membrane, and
within the enclosed volume is a lumen or thylakoid space. Embedded
in the thylakoid membrane are integral and peripheral membrane
protein complexes of the photosynthetic system.
Plants absorb light primarily using the pigment chlorophyll. The green
part of the light spectrum is not absorbed but is reflected which is the
reason that most plants have a green color. Besides chlorophyll,
plants also use pigments such as carotenes and xanthophylls.[22] Algae
also use chlorophyll, but various other pigments are present, such
as phycocyanin, carotenes, and xanthophylls in green
algae, phycoerythrin in red algae (rhodophytes)
and fucoxanthin in brown algae and diatoms resulting in a wide variety
of colors.
These pigments are embedded in plants and algae in complexes
called antenna proteins. In such proteins, the pigments are arranged
to work together. Such a combination of proteins is also called a light-
harvesting complex.[23]
Although all cells in the green parts of a plant have chloroplasts, the
majority of those are found in specially adapted structures
called leaves. Certain species adapted to conditions of strong sunlight
and aridity, such as many Euphorbia and cactus species, have their
main photosynthetic organs in their stems. The cells in the interior
tissues of a leaf, called the mesophyll, can contain between 450,000
and 800,000 chloroplasts for every square millimeter of leaf. The
surface of the leaf is coated with a water-resistant waxy cuticle that
protects the leaf from excessive evaporation of water and decreases
the absorption of ultraviolet or blue light to minimize heating. The
transparent epidermis layer allows light to pass through to
the palisade mesophyll cells where most of the photosynthesis takes
place.
Light-dependent reactions
Main article: Light-dependent reactions

Light-dependent reactions
of photosynthesis at the thylakoid membrane
In the light-dependent reactions, one molecule of the
pigment chlorophyll absorbs one photon and loses one electron. This
electron is taken up by a modified form of chlorophyll
called pheophytin, which passes the electron to a quinone molecule,
starting the flow of electrons down an electron transport chain that
leads to the ultimate reduction of NADP to NADPH. In addition, this
creates a proton gradient (energy gradient) across the chloroplast
membrane, which is used by ATP synthase in the synthesis of ATP.
The chlorophyll molecule ultimately regains the electron it lost when
a water molecule is split in a process called photolysis, which
releases oxygen.
The overall equation for the light-dependent reactions under the
conditions of non-cyclic electron flow in green plants is:[24]
2 H2O + 2 NADP+ + 3 ADP + 3 Pi + light → 2 NADPH + 2 H+ + 3 ATP +
O2
Not all wavelengths of light can support photosynthesis. The
photosynthetic action spectrum depends on the type of accessory
pigments present. For example, in green plants, the action spectrum
resembles the absorption
spectrum for chlorophylls and carotenoids with absorption peaks in
violet-blue and red light. In red algae, the action spectrum is blue-
green light, which allows these algae to use the blue end of the
spectrum to grow in the deeper waters that filter out the longer
wavelengths (red light) used by above-ground green plants. The non-
absorbed part of the light spectrum is what gives photosynthetic
organisms their color (e.g., green plants, red algae, purple bacteria)
and is the least effective for photosynthesis in the
respective organisms.
Z scheme

The "Z scheme"

In plants, light-dependent reactions occur in the thylakoid
membranes of the chloroplasts where they drive the synthesis
of ATP and NADPH. The light-dependent reactions are of two
forms: cyclic and non-cyclic.
In the non-cyclic reaction, the photons are captured in the light-
harvesting antenna complexes of photosystem II by chlorophyll and
other accessory pigments (see diagram at right). The absorption of a
photon by the antenna complex loosens an electron by a process
called photoinduced charge separation. The antenna system is at the
core of the chlorophyll molecule of the photosystem II reaction center.
That loosened electron is taken up by the primary electron-
acceptor molecule, pheophytin. As the electrons are shuttled through
an electron transport chain (the so-called Z-scheme shown in the
diagram), a chemiosmotic potential is generated by pumping proton
cations (H+) across the membrane and into the thylakoid space. An
ATP synthase enzyme uses that chemiosmotic potential to make ATP
during photophosphorylation, whereas NADPH is a product of the
terminal redox reaction in the Z-scheme. The electron enters a
chlorophyll molecule in Photosystem I. There it is further excited by
the light absorbed by that photosystem. The electron is then passed
along a chain of electron acceptors to which it transfers some of
its energy. The energy delivered to the electron acceptors is used to
move hydrogen ions across the thylakoid membrane into the lumen.
The electron is eventually used to reduce the coenzyme NADP with
an H+ to NADPH (which has functions in 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.
Water photolysis
Main articles: Photodissociation and Oxygen evolution
Linear electron transport through a photosystem will leave the reaction
center of that photosystem oxidized. Elevating another electron will
first require re-reduction of the reaction center. The excited electrons
lost from the reaction center (P700) of photosystem I are replaced by
transfer from plastocyanin, whose electrons come from electron
transport through photosystem II. Photosystem II, as the first step of
the Z-scheme, requires an external source of electrons to reduce its
oxidized chlorophyll a reaction center. The source of electrons for
photosynthesis in green plants and cyanobacteria is water. Two water
molecules are oxidized by the energy of four successive charge-
separation reactions of photosystem II to yield a molecule
of diatomic oxygen and four hydrogen ions. The electrons yielded are
transferred to a redox-active tyrosine residue that is oxidized by the
energy of P680+. This resets the ability of P680 to absorb another
photon and release another photo-dissociated electron. The oxidation
of water is catalyzed in photosystem II by a redox-active structure that
contains four manganese ions and a calcium ion; this oxygen-evolving
complex binds two water molecules and contains the four oxidizing
equivalents that are used to drive the water-oxidizing reaction (Kok's
S-state diagrams). The hydrogen ions are released in the thylakoid
lumen and therefore contribute to the transmembrane chemiosmotic
potential that leads to ATP synthesis. Oxygen is a waste product of
light-dependent reactions, but the majority of organisms on Earth use
oxygen and its energy for cellular respiration, including photosynthetic
organisms.[25][26]
Light-independent reactions
Calvin cycle
Main articles: Light-independent reactions and Carbon fixation
In the light-independent (or "dark") reactions, the
enzyme RuBisCO captures CO2 from the atmosphere and, in
a process called the Calvin cycle, uses the newly formed NADPH and
releases three-carbon sugars, which are later combined to
form sucrose and starch. The overall equation for the light-
independent reactions in green plants is[24]: 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 three-carbon sugar intermediate, which
is then converted into the final carbohydrate products. The simple
carbon sugars photosynthesis produces are then used to form
other organic compounds, such as the building material cellulose,
the precursors for lipid and amino acid biosynthesis, or as a fuel
in cellular respiration. The latter occurs not only in plants but also
in animals when the carbon and energy from plants is passed through
a food chain.
The fixation or reduction of carbon dioxide is a process in which
carbon dioxide combines with a five-carbon sugar, ribulose 1,5-
bisphosphate, to yield two molecules of a three-carbon
compound, glycerate 3-phosphate, also known as 3-
phosphoglycerate. Glycerate 3-phosphate, in the presence
of ATP and NADPH produced during the light-dependent stages, is
reduced to glyceraldehyde 3-phosphate. This product is also referred
to as 3-phosphoglyceraldehyde (PGAL) or, more generically,
as triose phosphate. Most (five out of six molecules) of the
glyceraldehyde 3-phosphate produced are used to regenerate ribulose
1,5-bisphosphate so the process can continue. The triose phosphates
not thus "recycled" often condense to form hexose phosphates, which
ultimately yield sucrose, starch, and cellulose, as well
as glucose and fructose. The sugars produced during
carbon metabolism yield carbon skeletons that can be used for
other metabolic reactions like the production of amino acids and lipids.
Carbon concentrating mechanisms
On land
Main articles: C4 carbon fixation, CAM photosynthesis, and Alarm
photosynthesis
 Overview of C4 carbon
fixation. (This image mistakenly shows lactic acid instead of pyruvate,
and all the species ending in "-ate" are shown as unionized acids,
such as malic acid and so on).
In hot and dry conditions, plants close their stomata 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 photorespiration by the oxygenase activity
of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) and
decrease in carbon fixation. Some plants have evolved mechanisms
to increase the CO2 concentration in the leaves under these
conditions.[27]
Plants that use the C4 carbon fixation process chemically fix carbon
dioxide in the cells of the mesophyll by adding it to the three-carbon
molecule phosphoenolpyruvate (PEP), a reaction catalyzed by
an enzyme called PEP carboxylase, creating the four-carbon organic
acid oxaloacetic acid. Oxaloacetic acid or malate synthesized by this
process is then translocated to specialized bundle sheath cells where
the enzyme RuBisCO and other Calvin cycle enzymes are located,
and where CO2 released by decarboxylation of the four-carbon acids is
then fixed by RuBisCO activity to the three-carbon 3-phosphoglyceric
acids. The physical separation of RuBisCO from the oxygen-
generating light reactions reduces photorespiration and increases
CO2 fixation and, thus, the photosynthetic capacity of the leaf.
[28]
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 C3 plants 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 carbon fixation;[29] however, the evolution of C4 in over
sixty plant lineages makes it a striking example of convergent
evolution.[27] C2 photosynthesis, which involves carbon-concentration by
selective breakdown of photorespiratory glycine, is both an
evolutionary precursor to C4 and a useful carbon-concentrating
mechanism in its own right.[30]
Xerophytes, such as cacti and most succulents, also use PEP
carboxylase to capture carbon dioxide in a process
called Crassulacean acid metabolism (CAM). In contrast to
C4 metabolism, which spatially 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 malic acid via carboxylation
of phosphoenolpyruvate 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. CAM is used by 16,000 species of plants.[31]
Calcium-oxalate-accumulating plants, such as Amaranthus
hybridus and Colobanthus quitensis, show a variation of
photosynthesis where calcium oxalate crystals function as
dynamic carbon pools, supplying carbon dioxide (CO2) to
photosynthetic cells when stomata are partially or totally closed. This
process was named alarm photosynthesis. Under stress conditions
(e.g., water deficit), oxalate released from calcium oxalate crystals is
converted to CO2 by an oxalate oxidase enzyme, and the produced
CO2 can support the Calvin cycle reactions. Reactive hydrogen
peroxide (H2O2), the byproduct of oxalate oxidase reaction, can
be neutralized by catalase. Alarm photosynthesis represents a
photosynthetic variant to be added to the well-known C4 and CAM
pathways. However, alarm photosynthesis, in contrast to these
pathways, operates as a biochemical pump that collects carbon from
the organ interior (or from the soil) and not from the atmosphere.[32][33]
In water
Cyanobacteria possess carboxysomes, which increase the
concentration of CO2 around RuBisCO to increase the rate of
photosynthesis. An enzyme, carbonic anhydrase, located within the
carboxysome, releases CO2 from dissolved hydrocarbonate
ions (HCO−
3). Before the CO2 can diffuse out, RuBisCO concentrated within the

carboxysome quickly sponges it up. HCO−

3 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 HCO−
3 ions to accumulate within the cell from where they diffuse into the

carboxysomes.[34] Pyrenoids in algae and hornworts also act to

concentrate CO2 around RuBisCO.[35]
Order and kinetics
The overall process of photosynthesis takes place in four stages:[11]

Stage Event Site Time scale

Energy transfer in antenna

1 Femtosecond to picosecond
chlorophyll

Transfer of Thylakoid
2 electrons in photochemical membranes in Picosecond to nanosecond
reactions the chloroplasts

Electron transport
3 Microsecond to millisecond
chain and ATP synthesis

Stroma of the
Carbon fixation and export
4 chloroplasts and Millisecond to second
of stable products
the cell cytosol

Efficiency
Main article: Photosynthetic efficiency
Plants usually convert light into chemical energy with a photosynthetic
efficiency of 3–6%.[36][37] Absorbed light that is unconverted
is dissipated primarily as heat, with a small fraction (1–2%)[38] reemitted
as chlorophyll fluorescence at longer (redder) wavelengths. This fact
allows measurement of the light reaction of photosynthesis by using
chlorophyll fluorometers.[38]
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%.
[39]
By comparison, solar panels convert light into electric energy at an
efficiency of approximately 6–20% for mass-produced panels, and
above 40% in laboratory devices. Scientists are studying
photosynthesis in hopes of developing plants with increased yield.[37]
The efficiency of both light and dark reactions can be measured, but
the relationship between the two can be complex. For example,
the light reaction creates ATP and NADPH energy molecules,
which C3 plants can use for carbon fixation or photorespiration.
[40]
Electrons may also flow to other electron sinks.[41][42][43] For this reason,
it is not uncommon for authors to differentiate between work done
under non-photorespiratory conditions and under photorespiratory
conditions.[44][45][46]
Chlorophyll fluorescence of photosystem II can measure the light
reaction, and infrared gas analyzers can measure the dark reaction.
[47]
An integrated chlorophyll fluorometer and gas exchange system can
investigate both light and dark reactions when researchers use the
two separate systems together.[48] Infrared gas analyzers and
some moisture sensors are sensitive enough to measure
the photosynthetic assimilation of CO2 and of ΔH2O using reliable
methods.[49] CO2 is commonly measured in μmols/(m2/s), parts
per million, or volume per million; and H2O is commonly measured
in mmols/(m2/s) or in mbars.[49] By measuring CO2 assimilation, ΔH2O,
leaf temperature, barometric pressure, leaf area,
and photosynthetically active radiation (PAR), it becomes possible to
estimate, "A" or carbon assimilation, "E" or transpiration, "gs"
or stomatal conductance, and "Ci" or intracellular CO2.[49] However, it is
more common to use chlorophyll fluorescence for plant stress
measurement, where appropriate, because the most commonly used
parameters FV/FM and Y(II) or F/FM' can be measured in a few
seconds, allowing the investigation of larger plant populations.[46]
Gas exchange systems that offer control of CO2 levels, above and
below ambient, allow the common practice of measurement of A/Ci
curves, at different CO2 levels, to characterize a plant's photosynthetic
response.[49]
Integrated chlorophyll fluorometer – gas exchange systems allow a
more precise measure of photosynthetic response and mechanisms.[47]
[48]
While standard gas exchange photosynthesis systems can measure
Ci, or substomatal CO2 levels, the addition of integrated chlorophyll
fluorescence measurements allows a more precise measurement of
CC, the estimation of CO2 concentration at the site of carboxylation in
the chloroplast, to replace Ci.[48][50] CO2 concentration in the chloroplast
becomes possible to estimate with the measurement of mesophyll
conductance or gm using an integrated system.[47][48][51]
Photosynthesis measurement systems are not designed to directly
measure the amount of light the leaf absorbs, but analysis
of chlorophyll fluorescence, P700- and P515-absorbance, and gas
exchange measurements reveal detailed information about, e.g.,
the photosystems, quantum efficiency and the CO2 assimilation rates.
With some instruments, even wavelength dependency of the
photosynthetic efficiency can be analyzed.[52]
A phenomenon known as quantum walk increases the efficiency of the
energy transport of light significantly. In the photosynthetic cell of
an alga, bacterium, or plant, there are light-sensitive molecules
called chromophores arranged in an antenna-shaped structure called
a photocomplex. When a photon is absorbed by a chromophore, it is
converted into a quasiparticle referred to as an exciton, which jumps
from chromophore to chromophore towards the reaction center of the
photocomplex, a collection of molecules that traps its energy in a
chemical form accessible to the cell's metabolism. The exciton's wave
properties enable it to cover a wider area and try out several possible
paths simultaneously, allowing it to instantaneously "choose" the most
efficient route, where it will have the highest probability of arriving at
its destination in the minimum possible time.
Because that quantum walking takes place at temperatures far higher
than quantum phenomena usually occur, it is only possible over very
short distances. Obstacles in the form of destructive interference
cause the particle to lose its wave properties for an instant before it
regains them once again after it is freed from its locked position
through a classic "hop". The movement of the electron towards the
photo center is therefore covered in a series of conventional hops and
quantum walks.[53][54][55]
Evolution
Main article: Evolution of photosynthesis

Life timeline
This box:
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 ← Earth formed
−4500 — ← Earliest water
– Water ← LUCA
← Earliest fossils
— ← LHB meteorites
–
← Earliest oxygen
Single-celled life ← Pongola glaciation*
−4000 — ← Atmospheric oxygen
← Huronian glaciation*
– ← Sexual reproduction
Photosynthesis
— ← Earliest multicellular life
← Earliest fungi
– ← Earliest plants
Eukaryotes
−3500 —
← Earliest animals
← Cryogenian ice age*
– ← Ediacaran biota
Multicellular life ← Cambrian explosion
— ← Andean glaciation*
– ← Earliest tetrapods
P ← Karoo ice age*
−3000 — l ← Earliest apes / humans
– a ← Quaternary ice age*
n
— t
s
–

−2500 —
Arthropods Molluscs
–
Flowers
—
Dinosaurs
–

−2000 —
Mammals
–
Birds
—
Primates
–
H
−1500 —
a
– d
e
— a
– n

−1000 —

–
A
—
r
– c
h
−500 — e
– a
n
—
 –

0—

P
r
o
t
e
r
o
z
o
i
c
P
h
a
n
e
r
o
z
o
i
c

(million years ago)

*Ice Ages

Fossils of what are thought to

be filamentous photosynthetic organisms have been dated at 3.4
billion years old.[56][57] More recent studies also suggest that
photosynthesis may have begun about 3.4 billion years ago,[58]
[59]
though the first direct evidence of photosynthesis comes
from thylakoid membranes preserved in 1.75-billion-year-old cherts.[60]
Oxygenic photosynthesis is the main source of oxygen in the Earth's
atmosphere, and its earliest appearance is sometimes referred to as
the oxygen catastrophe. Geological evidence suggests that oxygenic
photosynthesis, such as that in cyanobacteria, became important
during the Paleoproterozoic era around two billion years ago. Modern
photosynthesis in plants and most photosynthetic prokaryotes is
oxygenic, using water as an electron donor, which is oxidized to
molecular oxygen in the photosynthetic reaction center.
Symbiosis and the origin of chloroplasts
 Plant cells with visible chloroplasts
(from a moss, Plagiomnium affine)
Several groups of animals have formed symbiotic relationships with
photosynthetic algae. These are most common in corals, sponges,
and sea anemones. Scientists presume that this is due to the
particularly simple body plans and large surface areas of these
animals compared to their volumes.[61] In addition, a few
marine mollusks, such as Elysia viridis and Elysia chlorotica, also
maintain a symbiotic relationship with chloroplasts they capture from
the algae in their diet and then store in their bodies (see Kleptoplasty).
This allows the mollusks to survive solely by photosynthesis for
several months at a time.[62][63] Some of the genes from the plant cell
nucleus have even been transferred to the slugs, so that the
chloroplasts can be supplied with proteins they need to survive.[64]
An even closer form of symbiosis may explain the origin of
chloroplasts. Chloroplasts have many similarities with
photosynthetic bacteria, including a circular chromosome, prokaryotic-
type ribosome, and similar proteins in the photosynthetic reaction
center.[65][66] The endosymbiotic theory suggests that photosynthetic
bacteria were acquired (by endocytosis) by early eukaryotic cells to
form the first plant cells. Therefore, chloroplasts may be
photosynthetic bacteria that adapted to life inside plant cells.
Like mitochondria, chloroplasts possess their own DNA, separate from
the nuclear DNA of their plant host cells and the genes in this
chloroplast DNA resemble those found in cyanobacteria.[67] DNA in
chloroplasts codes for redox proteins such as those found in the
photosynthetic reaction centers. The CoRR Hypothesis proposes that
this co-location of genes with their gene products is required for redox
regulation of gene expression, and accounts for the persistence of
DNA in bioenergetic organelles.[68]
Photosynthetic eukaryotic lineages
Symbiotic and kleptoplastic organisms excluded:

 The glaucophytes and the red and green algae—

clade Archaeplastida (uni- and multicellular)
 The cryptophytes—clade Cryptista (unicellular)
 The haptophytes—clade Haptista (unicellular)
 The dinoflagellates and chromerids in the
superphylum Myzozoa, and Pseudoblepharisma in the
phylum Ciliophora—clade Alveolata (unicellular)
  The ochrophytes—clade Stramenopila (uni- and
multicellular)
 The chlorarachniophytes and three species of Paulinella in
the phylum Cercozoa—clade Rhizaria (unicellular)
 The euglenids—clade Excavata (unicellular)
Except for the euglenids, which are found within the Excavata, all of
these belong to the Diaphoretickes. Archaeplastida and the
photosynthetic Paulinella got their plastids, which are surrounded by
two membranes, through primary endosymbiosis in two separate
events, by engulfing a cyanobacterium. The plastids in all the other
groups have either a red or green algal origin, and are referred to as
the "red lineages" and the "green lineages". The only known exception
is the ciliate Pseudoblepharisma tenue, which in addition to its plastids
that originated from green algae also has a purple sulfur bacterium as
symbiont. In dinoflagellates and euglenids the plastids are surrounded
by three membranes, and in the remaining lines by four.
A nucleomorph, remnants of the original algal nucleus located
between the inner and outer membranes of the plastid, is present in
the cryptophytes (from a red alga) and chlorarachniophytes (from a
green alga).[69] Some dinoflagellates that lost their photosynthetic ability
later regained it again through new endosymbiotic events with
different algae. While able to perform photosynthesis, many of these
eukaryotic groups are mixotrophs and practice heterotrophy to various
degrees.
Photosynthetic prokaryotic lineages
Early photosynthetic systems, such as those in green and purple
sulfur and green and purple nonsulfur bacteria, are thought to have
been anoxygenic, and used various other molecules than water
as electron donors. Green and purple sulfur bacteria are thought to
have used hydrogen and sulfur as electron donors. Green nonsulfur
bacteria used various amino and other organic acids as electron
donors. Purple nonsulfur bacteria used a variety of nonspecific organic
molecules. The use of these molecules is consistent with the
geological evidence that Earth's early atmosphere was
highly reducing at that time.[70]
With a possible exception of Heimdallarchaeota, photosynthesis is not
found in archaea.[71] Haloarchaea are phototrophic and can absorb
energy from the sun, but do not harvest carbon from the atmosphere
and are therefore not photosynthetic.[72] Instead of chlorophyll they use
rhodopsins, which convert light-energy to ion gradients but cannot
mediate electron transfer reactions.[73][74]
In bacteria eight photosynthetic lineages are currently known:[75][76][77][78]

 Cyanobacteria, the only prokaryotes performing oxygenic

photosynthesis and the only prokaryotes that contain two
types of photosystems (type I (RCI), also known as Fe-S
type, and type II (RCII), also known as quinone type). The
 seven remaining prokaryotes have anoxygenic
photosynthesis and use versions of either type I or type II.
 Chlorobi (green sulfur bacteria) Type I
 Heliobacteria Type I
 Chloracidobacterium Type I
 Proteobacteria (purple sulfur bacteria and purple non-sulfur
bacteria) Type II
 Chloroflexota (green non-sulfur bacteria) Type II
 Gemmatimonadota Type II
 Eremiobacterota Type II
Cyanobacteria and the evolution of photosynthesis
The biochemical capacity to use water as the source for electrons in
photosynthesis evolved once, in a common ancestor of
extant cyanobacteria (formerly called blue-green algae). 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.[79][80] 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.[81] Available
evidence from geobiological studies of Archean (>2500
Ma) sedimentary rocks indicates that life existed 3500 Ma, but the
question of when oxygenic photosynthesis evolved is still unanswered.
A clear paleontological window on cyanobacterial evolution opened
about 2000 Ma, revealing an already-diverse biota of cyanobacteria.
Cyanobacteria remained the principal primary producers of oxygen
throughout the Proterozoic Eon (2500–543 Ma), in part because the
redox structure of the oceans favored photoautotrophs capable
of nitrogen fixation.[citation needed] Green algae joined cyanobacteria as the
major primary producers of oxygen on continental shelves near the
end of the Proterozoic, but only with the Mesozoic (251–66 Ma)
radiations of dinoflagellates, coccolithophorids, and diatoms did
the primary production of oxygen in marine shelf waters take modern
form. Cyanobacteria remain critical to marine ecosystems as primary
producers of oxygen in oceanic gyres, as agents of biological nitrogen
fixation, and, in modified form, as the plastids of marine algae.[82]
Experimental history
Discovery
Although some of the steps in photosynthesis are still not completely
understood, the overall photosynthetic equation has been known since
the 19th century.
 Portrait of Jan Baptist van
Helmont by Mary Beale, c. 1674
Jan van Helmont began the research of the process in the mid-17th
century when he carefully measured the mass of the soil a plant was
using and the mass of the plant as it grew. After noticing that the soil
mass changed very little, he hypothesized that the mass of
the growing plant must come from the water, the only substance he
added to the potted plant. His hypothesis was partially accurate –
much of the gained mass comes from carbon dioxide as well as water.
However, this was a signaling point to the idea that the bulk of a
plant's biomass comes from the inputs of photosynthesis, not the soil
itself.
Joseph Priestley, a chemist and minister, discovered that when he
isolated a volume of air under an inverted jar and burned a candle in it
(which gave off CO2), the candle would burn out very quickly, much
before it ran out of wax. He further discovered that a mouse could
similarly "injure" air. He then showed that a plant could restore the air
the candle and the mouse had "injured."[83]
In 1779, Jan Ingenhousz repeated Priestley's experiments. He
discovered that it was the influence of sunlight on the plant that could
cause it to revive a mouse in a matter of hours.[83][84]
In 1796, Jean Senebier, a Swiss pastor, botanist,
and naturalist, demonstrated that green plants consume carbon
dioxide and release oxygen under the influence of light. Soon
afterward, Nicolas-Théodore de Saussure showed that the increase in
mass of the plant as it grows could not be due only to uptake of
CO2 but also to the incorporation of water. Thus, the basic reaction by
which organisms use photosynthesis to produce food (such
as glucose) was outlined.[85]
Refinements
Cornelis Van Niel made key discoveries explaining the chemistry of
photosynthesis. By studying purple sulfur bacteria and green bacteria,
he was the first to demonstrate that photosynthesis is a light-
 dependent redox reaction in which hydrogen reduces (donates
its atoms as electrons and protons to) carbon dioxide.
Robert Emerson discovered two light reactions by testing plant
productivity using different wavelengths of light. With the red alone,
the light reactions were suppressed. When blue and red were
combined, the output was much more substantial. Thus, there were
two photosystems, one absorbing up to 600 nm wavelengths, the
other up to 700 nm. The former is known as PSII, the latter is PSI. PSI
contains only chlorophyll "a", PSII contains primarily chlorophyll "a"
with most of the available chlorophyll "b", among other pigments.
These include phycobilins, which are the red and blue pigments of red
and blue algae, respectively, and fucoxanthol for brown algae and
diatoms. The process is most productive when the absorption of
quanta is equal in both PSII and PSI, assuring that input energy from
the antenna complex is divided between the PSI and PSII systems,
which in turn powers the photochemistry.[11]
Robert Hill thought that a complex of reactions consisted of an
intermediate to cytochrome b6 (now a plastoquinone), and that another
was from cytochrome f to a step in the carbohydrate-generating
mechanisms. These are linked by plastoquinone, which does require
energy to reduce cytochrome f. Further experiments to prove that the
oxygen developed during the photosynthesis of green plants came
from water were performed by Hill in 1937 and 1939. He showed that
isolated chloroplasts give off oxygen in the presence of unnatural
reducing agents like iron oxalate, ferricyanide or benzoquinone after
exposure to light. In the Hill reaction:[86]
2 H2O + 2 A + (light, chloroplasts) → 2 AH2 + O2
A is the electron acceptor. Therefore, in light, the electron acceptor
is reduced and oxygen is evolved. Samuel Ruben and Martin
Kamen used radioactive isotopes to determine that the oxygen
liberated in photosynthesis came from the water.

Melvin Calvin works in his photosynthesis

laboratory.
Melvin Calvin and Andrew Benson, along with James Bassham,
elucidated the path of carbon assimilation (the photosynthetic
carbon reduction cycle) in plants. The carbon reduction cycle is
known as the Calvin cycle, but many scientists refer to it as the
Calvin-Benson, Benson-Calvin, or even Calvin-Benson-Bassham
(or CBB) Cycle.
Nobel Prize–winning scientist Rudolph A. Marcus was later able to
discover the function and significance of the electron transport
chain.
Otto Heinrich Warburg and Dean Burk discovered the I-quantum
photosynthesis reaction that splits CO2, activated by the
respiration.[87]
In 1950, first experimental evidence for the existence
of photophosphorylation in vivo was presented by Otto
Kandler using intact Chlorella cells and interpreting his findings as
light-dependent ATP formation.[88] In 1954, Daniel I. Arnon et al.
discovered photophosphorylation in vitro in
isolated chloroplasts with the help of P32.[89][90]
Louis N. M. Duysens and Jan Amesz discovered that chlorophyll
"a" will absorb one light, oxidize cytochrome f, while chlorophyll "a"
(and other pigments) will absorb another light but will reduce this
same oxidized cytochrome, stating the two light reactions are in
series.
Development of the concept
In 1893, the American botanist Charles Reid Barnes proposed two
terms, photosyntax and photosynthesis, for the biological process
of synthesis of complex carbon compounds out of carbonic acid, in
the presence of chlorophyll, under the influence of light. The
term photosynthesis is derived from the Greek phōs (φῶς, gleam)
and sýnthesis (σύνθεσις, arranging together),[91][92][93] while another
word that he designated was photosyntax, from sýntaxis (σύνταξις,
configuration). Over time, the term photosynthesis came into
common usage. Later discovery of anoxygenic photosynthetic
bacteria and photophosphorylation necessitated redefinition of the
term.[94]
C3 : C4 photosynthesis research
In the late 1940s at the University of California, Berkeley, the
details of photosynthetic carbon metabolism were sorted out by
the chemists Melvin Calvin, Andrew Benson, James Bassham and
a score of students and researchers utilizing the carbon-14 isotope
and paper chromatography techniques.[95] The pathway of
CO2 fixation by the algae Chlorella in a fraction of a second in light
resulted in a three carbon molecule called phosphoglyceric acid
(PGA). For that original and ground-breaking work, a Nobel Prize
in Chemistry was awarded to Melvin Calvin in 1961. In parallel,
plant physiologists studied leaf gas exchanges using the new
method of infrared gas analysis and a leaf chamber where the net
photosynthetic rates ranged from 10 to 13 μmol CO2·m−2·s−1, with
the conclusion that all terrestrial plants have the same
photosynthetic capacities, that are light saturated at less than 50%
of sunlight.[96][97]
Later in 1958–1963 at Cornell University, field grown maize was
reported to have much greater leaf photosynthetic rates of 40 μmol
CO2·m−2·s−1 and not be saturated at near full sunlight.[98][99] This
higher rate in maize was almost double of those observed in other
species such as wheat and soybean, indicating that large
differences in photosynthesis exist among higher plants. At the
University of Arizona, detailed gas exchange research on more
than 15 species of monocots and dicots uncovered for the first
time that differences in leaf anatomy are crucial factors in
differentiating photosynthetic capacities among species.[100][101] In
tropical grasses, including maize, sorghum, sugarcane, Bermuda
grass and in the dicot amaranthus, leaf photosynthetic rates were
around 38−40 μmol CO2·m−2·s−1, and the leaves have two types of
green cells, i.e. outer layer of mesophyll cells surrounding a tightly
packed cholorophyllous vascular bundle sheath cells. This type of
anatomy was termed Kranz anatomy in the 19th century by the
botanist Gottlieb Haberlandt while studying leaf anatomy of
sugarcane.[102] Plant species with the greatest photosynthetic rates
and Kranz anatomy showed no apparent photorespiration, very
low CO2 compensation point, high optimum temperature, high
stomatal resistances and lower mesophyll resistances for gas
diffusion and rates never saturated at full sun light.[103] The research
at Arizona was designated a Citation Classic in 1986.[101] These
species were later termed C4 plants as the first stable compound
of CO2 fixation in light has four carbons as malate and aspartate.[104]
[105][106]
Other species that lack Kranz anatomy were termed C3 type
such as cotton and sunflower, as the first stable carbon compound
is the three-carbon PGA. At 1000 ppm CO2 in measuring air, both
the C3 and C4 plants had similar leaf photosynthetic rates around
60 μmol CO2·m−2·s−1 indicating the suppression of photorespiration
in C3 plants.[100][101]
Factors

The leaf is the primary site of

photosynthesis in plants.
There are four main factors influencing photosynthesis[clarification
needed]
and several corollary factors. The four main are:[107]

 Light irradiance and wavelength

 Water absorption
 Carbon dioxide concentration
  Temperature.
Total photosynthesis is limited by a range of environmental factors.
These include the amount of light available, the amount
of leaf area a plant has to capture light (shading by other plants is
a major limitation of photosynthesis), the rate at which carbon
dioxide can be supplied to the chloroplasts to support
photosynthesis, the availability of water, and the availability of
suitable temperatures for carrying out photosynthesis.[108]
Light intensity (irradiance), wavelength and
temperature
See also: PI (photosynthesis-irradiance) curve

Absorbance spectra of free

chlorophyll a (blue) and b (red) in a solvent. The action spectra of
chlorophyll molecules are slightly modified in vivo depending on
specific pigment–protein interactions.
The process of photosynthesis provides the main input of free
energy into the biosphere, and is one of four main ways in which
radiation is important for plant life.[109]
The radiation climate within plant communities is extremely
variable, in both time and space.
In the early 20th century, Frederick Blackman and Gabrielle
Matthaei investigated the effects of light intensity (irradiance) and
temperature on the rate of carbon assimilation.

 At constant temperature, the rate of carbon assimilation

varies with irradiance, increasing as the irradiance
increases, but reaching a plateau at higher irradiance.
 At low irradiance, increasing the temperature has little
influence on the rate of carbon assimilation. At constant
high irradiance, the rate of carbon assimilation
increases as the temperature is increased.
These two experiments illustrate several important points: First, it
is known that, in general, photochemical reactions are not affected
by temperature. However, these experiments clearly show that
temperature affects the rate of carbon assimilation, so there must
be two sets of reactions in the full process of carbon assimilation.
These are the light-dependent 'photochemical' temperature-
independent stage, and the light-independent, temperature-
dependent stage. Second, Blackman's experiments illustrate the
concept of limiting factors. Another limiting factor is the wavelength
of light. Cyanobacteria, which reside several meters underwater,
cannot receive the correct wavelengths required to cause
photoinduced charge separation in conventional photosynthetic
pigments. To combat this problem, Cyanobacteria have a light-
harvesting complex called Phycobilisome.[110] This complex is made
up of a series of proteins with different pigments which surround
the reaction center.
Carbon dioxide levels and photorespiration

Photorespiration
As carbon dioxide concentrations rise, the rate at which sugars are
made by the light-independent reactions increases until limited by
other factors. RuBisCO, the enzyme that captures carbon dioxide
in the light-independent reactions, has a binding affinity for both
carbon dioxide and oxygen. When the concentration of carbon
dioxide is high, RuBisCO will fix carbon dioxide. However, if the
carbon dioxide concentration is low, RuBisCO will bind oxygen
instead of carbon dioxide. This process, called photorespiration,
uses energy, but does not produce sugars.
RuBisCO oxygenase activity is disadvantageous to plants for
several reasons:

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 (NH3), which is able to diffuse out
of the plant, leading to a loss of nitrogen.
A highly simplified summary is:
2 glycolate + ATP → 3-phosphoglycerate + carbon dioxide + ADP + NH3
The salvaging pathway for the products of RuBisCO
oxygenase activity is more commonly known as
photorespiration, since it is characterized by light-
dependent oxygen consumption and the release of carbon
dioxide.
See also

 Environment portal

 Ecology portal

 Earth sciences portal

 Jan Anderson (scientist)

 Artificial photosynthesis
 Calvin-Benson cycle
 Carbon fixation
 Cellular respiration
 Chemosynthesis
 Daily light integral
 Hill reaction
 Integrated fluorometer
 Light-dependent reaction
 Organic reaction
 Photobiology
 Photoinhibition
 Photosynthetic reaction center
 Photosynthetically active radiation
 Photosystem
 Photosystem I
 Photosystem II
 Quantasome
 Quantum biology
 Radiosynthesis
 Red edge
 Vitamin D
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Further reading
Library resources about
Photosynthesis

 Resources in your library

Books
 Bidlack JE, Stern KR, Jansky S (2003). Introductory Plant
Biology. New York: McGraw-Hill. ISBN 978-0-07-290941-8.
 Blankenship RE (2014). Molecular Mechanisms of
Photosynthesis (2nd ed.). John Wiley & Sons. ISBN 978-1-
4051-8975-0. Archived from the original on 2023-01-19.
Retrieved 2019-04-17.
 Govindjee, Beatty JT, Gest H, Allen JF (2006). Discoveries in
Photosynthesis. Advances in Photosynthesis and Respiration.
Vol. 20. Berlin: Springer. ISBN 978-1-4020-3323-
0. Archived from the original on 2023-01-19. Retrieved 2019-
04-17.
 Reece JB, et al. (2013). Campbell Biology. Benjamin
Cummings. ISBN 978-0-321-77565-8.

Papers
 Gupta RS, Mukhtar T, Singh B (Jun 1999). "Evolutionary
relationships among photosynthetic prokaryotes
(Heliobacterium chlorum, Chloroflexus aurantiacus,
cyanobacteria, Chlorobium tepidum and proteobacteria):
implications regarding the origin of photosynthesis". Molecular
Microbiology. 32 (5): 893–906. doi:10.1046/j.1365-
2958.1999.01417.x. PMID 10361294. S2CID 33477550.
 Rutherford AW, Faller P (Jan 2003). "Photosystem II:
evolutionary perspectives". Philosophical Transactions of the
Royal Society of London. Series B, Biological
Sciences. 358 (1429): 245–
253. doi:10.1098/rstb.2002.1186. PMC 1693113. PMID 12594
932.

External links
 A collection of photosynthesis pages for all levels
from a renowned expert (Govindjee)
 In depth, advanced treatment of photosynthesis,
also from Govindjee
 Science Aid: Photosynthesis Article appropriate
for high school science
 Metabolism, Cellular Respiration and
Photosynthesis – The Virtual Library of
Biochemistry and Cell Biology
 Overall examination of Photosynthesis at an
intermediate level
 Overall Energetics of Photosynthesis
  The source of oxygen produced by
photosynthesis Interactive animation, a textbook
tutorial
 Marshall J (2011-03-29). "First practical artificial
leaf makes debut". Discovery News. Archived
from the original on 2012-03-22. Retrieved 2011-
03-29.
 Photosynthesis – Light Dependent & Light
Independent Stages Archived 2011-09-10 at
the Wayback Machine
 Khan Academy, video introduction
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Botany

 History

 Outline

 Archaeobotany

 Astrobotany

 Bryology

 Dendrology

 Ethnobotany

 Paleobotany

Subdisciplines  Phycology
 Phytochemistry

 Phytogeography
o Geobotany

 Plant anatomy

 Plant ecology

 Plant pathology

Plant groups  Algae

 Archaeplastida

 Bryophyte

 Non-vascular plants

 Vascular plants

 Fern

 Lycophyte

 Spermatophytes
  Gymnosperm

 Angiosperm

Plant anatomy  Cell wall

 Plant  Phragmoplast
morphology Plant cells  Plastid
(glossary)  Plasmodesma

 Vacuole

 Cork

 Ground tissue
o Mesophyll

 Meristem
Tissues
 Storage organs

 Vascular tissue
o Vascular bundle

 Wood

 Bulb

 Root

 Rhizoid

 Rhizome

 Shoot

Vegetative o Bud

o Leaf

 Cataphyll

 Petiole
o Sessility

o Stem

Reproductive  Archegonium

(incl. Flower)  Antheridium

 Androecium
o Pollen

o Stamen

o Staminode

o Tapetum

 Flower
o Aestivation

o Flower development
 o Floral diagram

o Floral formula

o Floral symmetry

o Whorl

 Fruit
o Anatomy

o Berry

o Capsule

o Nut

o Pyrena

o Seed

 Dispersal

 Endosperm
 Gametophyte

 Gynandrium

 Gynoecium
o Ovary

 Locule

 Ovule
o Stigma

 Hypanthium (Floral cup)

 Inflorescence
o Bract

o Pedicellate

o Raceme

o Umbel

 Perianth
o Tepal

o Petal

o Sepal

 Plant embryo

 Receptacle

 Sporophyll

 Sporophyte

Surface  Cuticle

structures  Epicuticular wax

 Epidermis
  Nectar

 Stoma

 Thorns, spines, and prickles

 Trichome

 Aleurone

 Apical dominance

 Bulk flow

 Cellulose

 Nutrition

 Photosynthesis
o Chlorophyll

 Phytomelanin
Plant physiology
 Plant hormones
Materials
 Respiration
o Gas Exchange

o Cellular respiration

 Sap

 Starch

 Sugar

 Transpiration

 Turgor pressure

 Habit
o Cushion plants

o Rosettes

o Shrubs

 Prostrate shrubs

 Subshrubs
Plant growth
o Succulent plants
and habit
o Trees

o Vines

 Lianas
 Herbaceous plants

 Secondary growth

 Woody plants

Reproduction  Alternation of generations

 Evolution  Double fertilization

  Evolutionary development

 Evolutionary history
o timeline

 Flora

 Germination

 Pollination
o Artificial

o Pollinators
 Ecology
o Pollen tube

o Self

 Sporangium
o Microsporangia

 Microspore
o Megasporangium

 Megaspore
o Spore

 Biological classification

 Botanical nomenclature
o Botanical name

o Correct name

o Author citation

o International Code of Nomenclature (ICN)

o ICN for Cultivated Plants (ICNCP)

 Cultivated plant taxonomy

o Citrus taxonomy
Plant taxonomy
o Cultigen

 Cultivar

 Group

 Grex
 History of plant systematics

 Herbarium

 International Association for Plant Taxonomy (IAPT)

 Plant taxonomy systems

 Taxonomic rank

Practice  Agronomy

 Floriculture

 Forestry
  Horticulture

 Botanical terms

 Botanists
 Lists
o by author abbreviation
 Related topics
 Botanical expeditions

 Individual trees

 Category

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Metabolism map

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Ecology: Modelling ecosystems: Trophic components

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Ecology: Modelling ecosystems: Other components

 France

 BnF data

 Germany

Authority control databases: National  Israel

 United States

 Japan

 Czech Republic
Categories:
 Photosynthesis
 Agronomy
 Biological processes
 Botany
 Cellular respiration
 Ecosystems
 Metabolism
  Plant nutrition
 Plant physiology
 Quantum biology
 This page was last edited on 30 April 2024, at 00:19 (UTC).
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