Photosynthesis

Photosynthesis is a process used by

plants and other organisms to convert
light energy into chemical energy that,
through cellular respiration, can later be
released to fuel the organism's metabolic
activities. This chemical energy is stored
in carbohydrate molecules, such as
sugars, which are synthesized from
carbon dioxide and water – hence the
name photosynthesis, from the Greek
phōs (φῶς), "light", and sunthesis
(σύνθεσις), "putting together".[1][2][3] In
most cases, oxygen is also released as a
waste product. Most plants, algae, and
cyanobacteria perform photosynthesis;
such organisms are called
photoautotrophs. Photosynthesis is
largely responsible for producing and
maintaining the oxygen content of the
Earth's atmosphere, and supplies most of
the energy necessary for life on Earth.[4]

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
phytoplankton and terrestrial vegetation. Dark red
and blue-green indicate regions of high
photosynthetic activity in the ocean and on land,
respectively.
Although photosynthesis is performed
differently by different species, the
process always begins when energy from
light is absorbed by proteins called
reaction centres that contain green
chlorophyll pigments. In plants, these
proteins are held inside organelles called
chloroplasts, which are most abundant in
leaf cells, while in bacteria they are
embedded in the plasma membrane. In
these light-dependent reactions, some
energy is used to strip electrons from
suitable substances, such as water,
producing oxygen gas. The hydrogen
freed by the splitting of water is used in
the creation of two further compounds
that serve as short-term stores of energy,
enabling its transfer to drive other
reactions: these compounds are reduced
nicotinamide adenine dinucleotide
phosphate (NADPH) and adenosine
triphosphate (ATP), the "energy currency"
of cells.

In plants, algae and cyanobacteria, long-

term energy storage in the form of
sugars is produced by a subsequent
sequence of light-independent reactions
called the Calvin cycle. In the Calvin
cycle, atmospheric carbon dioxide is
incorporated into already existing organic
carbon compounds, such as ribulose
bisphosphate (RuBP).[5] Using the ATP
and NADPH produced by the light-
dependent reactions, the resulting
compounds are then reduced and
removed to form further carbohydrates,
such as glucose. In other bacteria,
different mechanisms such as the
reverse Krebs cycle are used to achieve
the same end.

The first photosynthetic organisms

probably evolved early in the evolutionary
history of life and most likely used
reducing agents such as hydrogen or
hydrogen sulfide, rather than water, as
sources of electrons.[6] Cyanobacteria
appeared later; the excess oxygen they
produced contributed directly to the
oxygenation of the Earth,[7] which
rendered the evolution of complex life
possible. Today, the average rate of
energy capture by photosynthesis
globally is approximately
130 terawatts,[8][9][10] which is about eight
times the current power consumption of
human civilization.[11] Photosynthetic
organisms also convert around 100–115
billion tons (91–104 petagrams) of
carbon into biomass per year.[12][13] The
phenomenon that plants receive some
energy from light – in addition to air, soil,
and water – was first discovered in 1779
by Jan Ingenhousz.

Overview
Photosynthesis changes sunlight into chemical
energy, splits water to liberate O2, and fixes CO2 into
sugar.

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.[4] In plants, algae,
and cyanobacteria, photosynthesis
releases oxygen. This is called oxygenic
photosynthesis and is by far the most
common type of photosynthesis used by
living organisms. Although there are
some differences between oxygenic
photosynthesis in plants, algae, and
cyanobacteria, the overall process is
quite similar in these organisms. There
are also many varieties of anoxygenic
photosynthesis, used mostly by certain
types of bacteria, which consume carbon
dioxide but do not release oxygen.
Carbon dioxide is converted into sugars
in a process called carbon fixation;
photosynthesis captures energy from
sunlight to convert carbon dioxide into
carbohydrate. 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 carbohydrate, cellular
respiration is the oxidation of
carbohydrate or other nutrients to carbon
dioxide. Nutrients used in cellular
respiration include carbohydrates, amino
acids and fatty acids. These nutrients are
oxidized to produce carbon dioxide and
water, and to release chemical energy to
drive the organism's metabolism.
Photosynthesis and cellular respiration
are distinct processes, as they take place
through different sequences of chemical
reactions and in different cellular
compartments.

The general equation for photosynthesis

as first proposed by Cornelis van Niel is
therefore:[14]

CO2 + 2H2A + photons →

carbon electron donor light energy
dioxide

[CH2O] + 2A + H2O
carbohydrate oxidized water
electron
donor

Since water is used as the electron donor

in oxygenic photosynthesis, the equation
for this process is:

CO2 + 2H2O + photons → [CH2O]

carbon water light energy carbohydrate
dioxide

+ O2 + H2O
oxygen water

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:

CO2 + H2O + photons → [CH2O] +

carbon water light energy carbohydrate
dioxide

O2
oxygen
Other processes substitute other
compounds (such as arsenite) for water
in the electron-supply role; for example
some microbes use sunlight to oxidize
arsenite to arsenate:[15] The equation for
this reaction is:

CO2 + (AsO3−) + photons → (AsO3−)

carbon 3 4
light energy
dioxide arsenite arsenate

+ CO (used to build other

carbon
monoxide

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

Most organisms that utilize 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][19]

Photosynthetic membranes
and organelles

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.[20]
However, the membrane may be tightly
folded into cylindrical sheets called
thylakoids,[21] or bunched up into round
vesicles called intracytoplasmic
membranes.[22] 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.[21]

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.[23]
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.[24]

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

Light-dependent reactions

Light-dependent reactions of photosynthesis at the

thylakoid membrane
In the light-dependent reactions, one
molecule of the pigment chlorophyll
absorbs one photon and loses one
electron. This electron is passed to 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 a dioxygen
(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:[25]

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 frees 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 freed electron is
transferred to 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), it initially
functions to generate a chemiosmotic
potential 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 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.
Water photolysis

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,
called P680. The source of electrons for
photosynthesis in green plants and
cyanobacteria is water. Two water
molecules are oxidized by four
successive charge-separation reactions
by 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 then
reduces the oxidized 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 (Dolai's
S-state diagrams). Photosystem II is the
only known biological enzyme that
carries out this oxidation of water. 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 for
cellular respiration, including
photosynthetic organisms.[26][27]

Light-independent reactions
Calvin cycle

In the light-independent (or "dark")

reactions, the enzyme RuBisCO captures
CO2 from the atmosphere and, in a
process called the Calvin cycle, 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[25]: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 into 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 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
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 (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 hexose phosphates,
which ultimately yield sucrose, starch
and cellulose. 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

Overview of C4 carbon fixation

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 and decrease in
carbon fixation. Some plants have
evolved mechanisms to increase the CO2
concentration in the leaves under these
conditions.[28]

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.[29] 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;[30] however,
the evolution of C4 in over 60 plant
lineages makes it a striking example of
convergent evolution.[28]

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. Sixteen
thousand species of plants use CAM.[31]
Calcium oxalate accumulating plants,
such as Amaranthus hybridus and
Colobanthus quitensis, showed 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
cyclereactions. Reactive hydrogen
peroxide (H2O2), the byproduct of oxalate
oxidase reaction, can be neutralized by
catalase. Alarm photosynthesis
represents an unknown photosynthetic
variation to be added to the already
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 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.[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:[13]

Stage Description Time scale

Energy transfer in antenna chlorophyll (thylakoid femtosecond to

1
membranes) picosecond

Transfer of electrons in photochemical reactions (thylakoid picosecond to

2
membranes) nanosecond

Electron transport chain and ATP synthesis (thylakoid microsecond to

3
membranes) millisecond

4 Carbon fixation and export of stable products millisecond to second

Efficiency
Plants usually convert light into chemical
energy with a photosynthetic efficiency
of 3–6%.[36] Absorbed light that is
unconverted is dissipated primarily as
heat, with a small fraction (1–2%)[37] re-
emitted as chlorophyll fluorescence at
longer (redder) wavelengths. This fact
allows measurement of the light reaction
of photosynthesis by using chlorophyll
fluorometers.[37]

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%.[38] 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.

The efficiency of both light and dark

reactions can be measured but the
relationship between the two can be
complex.[39] For example, the ATP and
NADPH energy molecules, created by the
light reaction, can be used for carbon
fixation or for photorespiration in C3
plants.[39] Electrons may also flow to
other electron sinks.[40][41][42] For this
reason, it is not uncommon for authors to
differentiate between work done under
non-photorespiratory conditions and
under photorespiratory
conditions.[43][44][45]
Chlorophyll fluorescence of photosystem
II can measure the light reaction, and
Infrared gas analyzers can measure the
dark reaction.[46] It is also possible to
investigate both at the same time using
an integrated chlorophyll fluorometer and
gas exchange system, or by using two
separate systems together.[47] Infrared
gas analyzers and some moisture
sensors are sensitive enough to measure
the photosynthetic assimilation of CO2,
and of ΔH2O using reliable methods[48]
CO2 is commonly measured in
μmols/(m2/s), parts per million or
volume per million and H2O is commonly
measured in mmol/(m2/s) or in
mbars.[48] By measuring CO2
assimilation, ΔH2O, leaf temperature,
barometric pressure, leaf area, and
photosynthetically active radiation or
PAR, it becomes possible to estimate, "A"
or carbon assimilation, "E" or
transpiration, "gs" or stomatal
conductance, and Ci or intracellular
CO2.[48] However, it is more common to
used chlorophyll fluorescence for plant
stress measurement, where appropriate,
because the most commonly used
measuring parameters FV/FM and Y(II)
or F/FM' can be made in a few seconds,
allowing the measurement of larger plant
populations.[45]
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.[48]

Integrated chlorophyll fluorometer – gas

exchange systems allow a more precise
measure of photosynthetic response and
mechanisms.[46][47] While standard gas
exchange photosynthesis systems can
measure Ci, or substomatal CO2 levels,
the addition of integrated chlorophyll
fluorescence measurements allows a
more precise measurement of CC to
replace Ci.[47][49] The estimation of CO2 at
the site of carboxylation in the
chloroplast, or CC, becomes possible
with the measurement of mesophyll
conductance or gm using an integrated
system.[46][47][50]

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.[51]

A phenomenon known as quantum walk

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
chromophores arranged in an antenna-
shaped structure named a
photocomplex. When a photon is
absorbed by a chromophore, it is
converted into a quasiparticle referred to
as an exciton, which jumps from
chromophore to chromophore towards
the reaction center of the photocomplex,
a collection of molecules that traps its
energy in a chemical form 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.[52][53][54]

Evolution

Life timeline

This box: view talk edit

ges
0 — Flowers Birds Prima ←Earliest apes
Mammals
P Pl t
 –P
h 
Plants tes
a ←Tetrapoda
Dinosaurs    
0 — n Arthropods Molluscs
←Cambrian
e ←Ediacaran
explosion biota
– o r ←Earliest animals
z ←Earliest plants
0 — o Multicellular
i
–
c life ←Sexual
reproduction
P
0 —  r ←Earliest fungi
o
t
–e Eukaryotes
r
0 — o
z
o
–i ←Oxygen crisis
c
0 — ←Atmospheric
oxygen

– Photosynthesis

0 — A
r
–hc 
e
0 — a ←Earliest oxygen
n Single-celled
– f LHB i
 – life ←LHB meteorites
0 — H ←Earliest life
a Water
– d 
e
a ←Earliest water
0 — n ←Earliest Earth
illion years ago) (−4540)

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 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
reducing at that time.[55]

Fossils of what are thought to be

filamentous photosynthetic organisms
have been dated at 3.4 billion years
old.[56][57] More recent studies, reported
in March 2018, also suggest that
photosynthesis may have begun about
3.4 billion years ago.[58][59]

The main source of oxygen in the Earth's

atmosphere derives from oxygenic
photosynthesis, and its first appearance
is sometimes referred to as the oxygen
catastrophe. Geological evidence
suggests that oxygenic photosynthesis,
such as that in cyanobacteria, became
important during the Paleoproterozoic
era around 2 billion years ago. Modern
photosynthesis in plants and most
photosynthetic prokaryotes is oxygenic.
Oxygenic photosynthesis uses water as
an electron donor, which is oxidized to
molecular oxygen (O2) in the
photosynthetic reaction center.

Symbiosis and the origin of

chloroplasts
Plant cells with visible chloroplasts (from a moss,
Plagiomnium affine)

Several groups of animals have formed

symbiotic relationships with
photosynthetic algae. These are most
common in corals, sponges and sea
anemones. It is presumed that this is due
to the particularly simple body plans and
large surface areas of these animals
compared to their volumes.[60] In
addition, a few marine mollusks Elysia
viridis and Elysia chlorotica also maintain
a symbiotic relationship with
chloroplasts they capture from the algae
in their diet and then store in their bodies
(see Kleptoplasty). This allows the
mollusks to survive solely by
photosynthesis for several months at a
time.[61][62] Some of the genes from the
plant cell nucleus have even been
transferred to the slugs, so that the
chloroplasts can be supplied with
proteins that they need to survive.[63]

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.[64][65] 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.[66] 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.[67]

Photosynthetic eukaryotic lineages

Symbiotic and kleptoplastic organisms

excluded:

The glaucophytes and the red and

green algae—clade Archaeplastida
(unicellular and multicellular)
The cryptophytes—clade Cryptista
(unicellular)
The haptophytes—clade Haptista
(unicellular)
 The dinoflagellates and chromerids in
the superphylum Myzozoa—clade
Alveolata (unicellular)
The ochrophytes—clade Heterokonta
(unicellular 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, all of them

belong to the Diaphoretickes.
Archaeplastida and the photosynthetic
Paulinella got their plastids 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". While able to perform
photosynthesis, many of them are
mixotrophs and practice heterotrophy to
various degrees.

Cyanobacteria and the evolution of

photosynthesis

The biochemical capacity to use water as

the source for electrons in
photosynthesis evolved once, in a
common ancestor of extant
cyanobacteria (formerly called blue-
green algae), which are the only
prokaryotes performing oxygenic
photosynthesis. The geological record
indicates that this transforming event
took place early in Earth's history, at least
2450–2320 million years ago (Ma), and,
it is speculated, much earlier.[68][69]
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.[70] 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. Green algae joined
cyanobacteria as the major primary
producers of oxygen on continental
shelves near the end of the Proterozoic,
but it was 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.[71]

Discovery
Although some of the steps in
photosynthesis are still not completely
understood, the overall photosynthetic
equation has been known since the 19th
century.
Portrait of Jan Baptist van Helmont by Mary Beale,

c.1674

Jan van Helmont began the research of

the process in the mid-17th century when
he carefully measured the mass of the
soil used by a plant and the mass of the
plant as it grew. After noticing that the
soil mass changed very little, he
hypothesized that the mass of the
growing plant must come from the water,
the only substance he added to the
potted plant. His hypothesis was partially
accurate – much of the gained mass
also comes from carbon dioxide as well
as water. However, this was a signaling
point to the idea that the bulk of a plant's
biomass comes from the inputs of
photosynthesis, not the soil itself.

Joseph Priestley, a chemist and minister,

discovered that, when he isolated a
volume of air under an inverted jar, and
burned a candle in it (which gave off
CO2), the candle would burn out very
quickly, much before it ran out of wax. He
further discovered that a mouse could
similarly "injure" air. He then showed that
the air that had been "injured" by the
candle and the mouse could be restored
by a plant.[72]

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.[72][73]

In 1796, Jean Senebier, a Swiss pastor,

botanist, and naturalist, demonstrated
that green plants consume carbon
dioxide and release oxygen under the
influence of light. Soon afterward,
Nicolas-Théodore de Saussure showed
that the increase in mass of the plant as
it grows could not be due only to uptake
of CO2 but also to the incorporation of
water. Thus, the basic reaction by which
photosynthesis is used to produce food
(such as glucose) was outlined.[74]

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 –
electron 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 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.[13]

Robert Hill thought that a complex of

reactions consisting of an intermediate
to cytochrome b6 (now a plastoquinone),
another is from cytochrome f to a step in
the carbohydrate-generating
mechanisms. These are linked by
plastoquinone, which does require energy
to reduce cytochrome f for it is a
sufficient reductant. Further experiments
to prove that the oxygen developed
during the photosynthesis of green
plants came from water, were performed
by Hill in 1937 and 1939. He showed that
isolated chloroplasts give off oxygen in
the presence of unnatural reducing
agents like iron oxalate, ferricyanide or
benzoquinone after exposure to light.
The Hill reaction[75] is as follows:

2 H2O + 2 A + (light, chloroplasts) → 2

AH2 + O2

where A is the electron acceptor.

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

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

Nobel Prize-winning scientist Rudolph A.

Marcus was 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 the CO2, activated by
the respiration.[76]

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.[77] In 1954, Daniel I. Arnon et
al. discovered photophosphorylation in
vitro in isolated chloroplasts with the
help of P32.[78][79]

Louis N.M. Duysens and Jan Amesz

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

Development of the concept

In 1893, Charles Reid Barnes proposed

two terms, photosyntax and
photosynthesis, for the biological process
of synthesis of complex carbon
compounds out of carbonic acid, in the
presence of chlorophyll, under the
influence of light. Over time, the term
photosynthesis came into common
usage as the term of choice. Later
discovery of anoxygenic photosynthetic
bacteria and photophosphorylation
necessitated redefinition of the term.[80]

C3 : C4 photosynthesis research

After WWII at late 1940 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.[81] The
pathway of CO2 fixation by the algae
Chlorella in a fraction of a second in light
resulted in a 3 carbon molecule called
phosphoglyceric acid (PGA). For that
original and ground-breaking work, a
Nobel Prize in Chemistry was awarded to
Melvin Calvin in 1961. In parallel, plant
physiologists studied leaf gas exchanges
using the new method of infrared gas
analysis and a leaf chamber where the
net photosynthetic rates ranged from 10
to 13 μmol CO2·m−2·s−1, with the
conclusion that all terrestrial plants
having the same photosynthetic
capacities that were light saturated at
less than 50% of sunlight.[82][83]

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 was not
saturated at near full sunlight.[84][85] This
higher rate in maize was almost double
those observed in other species such as
wheat and soybean, indicating that large
differences in photosynthesis exist
among higher plants. At the University of
Arizona, detailed gas exchange research
on more than 15 species of monocot and
dicot uncovered for the first time that
differences in leaf anatomy are crucial
factors in differentiating photosynthetic
capacities among species.[86][87] 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.[88] 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.[89] The research at Arizona was
designated Citation Classic by the ISI
1986.[87] These species was later termed
C4 plants as the first stable compound of
CO2 fixation in light has 4 carbon as
malate and aspartate.[90][91][92] Other
species that lack Kranz anatomy were
termed C3 type such as cotton and
sunflower, as the first stable carbon
compound is the 3-carbon PGA. At 1000
ppm CO2 in measuring air, both the C3
and C4 plants had similar leaf
photosynthetic rates around 60 μmol
CO2·m−2·s−1 indicating the suppression
of photorespiration in C3 plants.[86][87]

Factors

The leaf is the primary site of photosynthesis in

plants.

There are three main factors affecting

photosynthesis and several corollary
factors. The three main are:
 Light irradiance and wavelength
Carbon dioxide concentration
Temperature.

Total photosynthesis is limited by a

range of environmental factors. These
include the amount of light available, the
amount of leaf area a plant has to
capture light (shading by other plants is a
major limitation of photosynthesis), 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.[93]
Light intensity (irradiance),
wavelength and temperature

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.[94]

The radiation climate within plant

communities is extremely variable, with
both time and space.

In the early 20th century, Frederick

Blackman and Gabrielle Matthaei
investigated the effects of light intensity
(irradiance) and temperature on the rate
of carbon assimilation.

At constant temperature, the rate of

carbon assimilation varies with
irradiance, increasing as the irradiance
increases, but reaching a plateau at
higher irradiance.
 At low irradiance, increasing the
temperature has little influence on the
rate of carbon assimilation. At
constant high irradiance, the rate of
carbon assimilation increases as the
temperature is increased.

These two experiments illustrate several

important points: First, it is known that, in
general, photochemical reactions are not
affected by temperature. However, these
experiments clearly show that
temperature affects the rate of carbon
assimilation, so there must be two sets
of reactions in the full process of carbon
assimilation. These are the light-
dependent 'photochemical' temperature-
independent stage, and the light-
independent, temperature-dependent
stage. Second, Blackman's experiments
illustrate the concept of limiting factors.
Another limiting factor is the wavelength
of light. Cyanobacteria, which reside
several meters underwater, cannot
receive the correct wavelengths required
to cause photoinduced charge
separation in conventional
photosynthetic pigments. To combat this
problem, a series of proteins with
different pigments surround the reaction
center. This unit is called a
phycobilisome.

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
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
Quantum biology
Radiosynthesis
Red edge
Vitamin D

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Further reading

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.
 Govindjee, Beatty JT, Gest H, Allen JF
(2006). Discoveries in Photosynthesis .
Advances in Photosynthesis and
Respiration. 20. Berlin: Springer. ISBN 978-
1-4020-3323-0.
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 12594932 .

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
Photosynthesis Discovery Milestones
– experiments and background
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.
 Photosynthesis – Light Dependent &
Light Independent Stages Archived
2011-09-10 at the Wayback Machine
Khan Academy, video introduction
Ehrenberg R (2017-12-15). "The
photosynthesis fix" . Knowable
Magazine.

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