Photosynthesis
Composite image showing the global distribution of photosyn-
Schematic of photosynthesis in plants. The carbohydrates pro-
duced are stored in or used by the plant.
Light
6CO2 6H2O C 6H12O6 6O2
Carbon dioxide Water Sugar Oxygen
Overall equation for the type of photosynthesis that occurs in
plants
Photosynthesis is a process used by plants and other
organisms to convert light energy into chemical energy
that can later be released to fuel the organisms activities
(energy transformation). This chemical energy is stored
in carbohydrate molecules, such as sugars, which are syn-
thesized from carbon dioxide and water hence the name
photosynthesis, from the Greek , phs, light, and
, synthesis, putting together.[1][2][3] In most
cases, oxygen is also released as a waste product. Most
plants, most algae, and cyanobacteria perform photosyn-
thesis; such organisms are called photoautotrophs. Pho-
tosynthesis is largely responsible for producing and main-
taining the oxygen content of the Earths atmosphere, and
supplies all of the organic compounds and most of the en-
ergy necessary for life on Earth.[4]
thesis, 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 dierently by dif-
ferent 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, produc-
ing oxygen gas. The hydrogen freed by the splitting of
water is used in the creation of two further compounds
that act as an immediate energy storage means: reduced
nicotinamide adenine dinucleotide phosphate (NADPH)
and adenosine triphosphate (ATP), the energy currency
of cells.
In plants, algae and cyanobacteria, long-term energy stor-
age in the form of sugars is produced by a subsequent se-
quence of light-independent reactions called the Calvin
cycle; some bacteria use dierent mechanisms, such as
the reverse Krebs cycle, to achieve the same end. 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.
The rst photosynthetic organisms probably evolved early
in the evolutionary history of life and most likely used
reducing agents such as hydrogen or hydrogen sulde,
rather than water, as sources of electrons.[6] Cyanobacte-
ria appeared later; the excess oxygen they produced con-
tributed directly to the oxygenation of the Earth,[7] which
1
2 1
rendered the evolution of complex life possible. To-
day, the average rate of energy capture by photosynthesis
globally is approximately 130 terawatts,[8][9][10] which is
about three times the current power consumption of hu-
man civilization.[11] Photosynthetic organisms also con-
vert around 100115 thousand million metric tonnes of
carbon into biomass per year.[12][13]
1 Overview
Light
H 2O O2
Light reactions
+
NAD
Pi
P
AT
CO2
NA
P
P
PH
AD
Calvin
Cycle
sugar
Photosynthesis changes sunlight into chemical energy, splits water
to liberate O2 , and xes 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. How-
ever, not all organisms that use light as a source of energy
carry out photosynthesis; photoheterotrophs use organic
compounds, rather than carbon dioxide, as a source of
carbon.[4] In plants, algae, and cyanobacteria, photosyn-
thesis releases oxygen. This is called oxygenic photosyn-
thesis and is by far the most common type of photosyn-
thesis used by living organisms. Although there are some
dierences between oxygenic photosynthesis in plants,
algae, and cyanobacteria, the overall process is quite sim-
ilar 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 re-
lease oxygen.
Carbon dioxide is converted into sugars in a process
called carbon xation. Photosynthesis provides the en-
ergy in the form of free electrons that are used to split
carbon from carbon dioxide that is then used to x that
OVERVIEW
carbon once again as carbohydrate. Carbon xation is an
endothermic redox reaction, so photosynthesis supplies
the energy that drives both process. In general outline,
photosynthesis is the opposite of cellular respiration, in
which glucose and other compounds are oxidized to pro-
duce carbon dioxide and water, and to release chemi-
cal energy (an exothermic reaction) to drive the organ-
isms metabolism. The two processes, reduction of car-
bon dioxide to carbohydrate and then later oxidation of
the carbohydrate, are distinct: photosynthesis and cel-
lular respiration take place through a dierent sequence
of chemical reactions and in dierent cellular compart-
ments.
The general equation for photosynthesis as rst proposed
by Cornelius van Niel is therefore:[14]
CO2 + 2H2 A + photons [CH2 O] + 2A +
H2 O
carbon dioxide + electron donor + light energy
carbohydrate + oxidized electron donor +
water
Since water is used as the electron donor in oxygenic pho-
tosynthesis, the equation for this process is:
CO2 + 2H2 O + photons [CH2 O] + O2 +
H2 O
carbon dioxide + water + light energy car-
bohydrate + 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 + H2 O + photons [CH2 O] + O2
carbon dioxide + water + light energy car-
bohydrate + oxygen
Other processes substitute other compounds (such as
arsenite) for water in the electron-supply role; for ex-
ample some microbes use sunlight to oxidize arsenite to
arsenate:[15] The equation for this reaction is:
CO2 + (AsO3
3) + photons (AsO3
4) + CO[16]
carbon dioxide + arsenite + light energy ar-
senate + carbon monoxide (used to build other
compounds in subsequent reactions)
Photosynthesis occurs in two stages. In the rst 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,
 3

the light-independent reactions use these products to cap- In plants and algae, photosynthesis takes place in
ture and reduce carbon dioxide. organelles called chloroplasts. A typical plant cell con-
Most organisms that utilize oxygenic photosynthesis use tains about 10 to 100 chloroplasts. The chloroplast is
visible light for the light-dependent reactions, although at enclosed by a membrane. This membrane is composed
least three use shortwave infrared or, more specically, of a phospholipid inner membrane, a phospholipid outer
far-red radiation. [17] membrane, and an intermembrane space. Enclosed by
the membrane is an aqueous uid called the stroma. Em-
Some organisms employ even more radical variants of bedded within the stroma are stacks of thylakoids (grana),
photosynthesis. Some archea use a simpler method that which are the site of photosynthesis. The thylakoids ap-
employs a pigment similar to those used for vision in ani- pear as attened disks. The thylakoid itself is enclosed by
mals. The bacteriorhodopsin changes its conguration in the thylakoid membrane, and within the enclosed volume
response to sunlight, acting as a proton pump. This pro- is a lumen or thylakoid space. Embedded in the thylakoid
duces a proton gradient more directly, which is then con- membrane are integral and peripheral membrane protein
verted to chemical energy. The process does not involve complexes of the photosynthetic system.
carbon dioxide xation and does not release oxygen, and
seems to have evolved separately from the more common Plants absorb light primarily using the pigment
types of photosynthesis.[18][19] chlorophyll. The green part of the light spectrum
is not absorbed but is reected which is the reason
that most plants have a green color. Besides chloro-
phyll, plants also use pigments such as carotenes and
2 Photosynthetic membranes and xanthophylls.[23] Algae also use chlorophyll, but vari-
organelles ous other pigments are present, such as phycocyanin,
carotenes, and xanthophylls in green algae, phycoerythrin
in red algae (rhodophytes) and fucoxanthin in brown
7 algae and diatoms resulting in a wide variety of colors.
1
8
2
3
These pigments are embedded in plants and algae in com-
plexes called antenna proteins. In such proteins, the pig-
ments are arranged to work together. Such a combination
9
of proteins is also called a light-harvesting complex.
10 Although all cells in the green parts of a plant have
4
5
11 chloroplasts, the majority of those are found in specially
6
12
adapted structures called leaves. Certain species adapted
to conditions of strong sunlight and aridity, such as many
Chloroplast ultrastructure: Euphorbia and cactus species, have their main photosyn-
1. outer membrane thetic organs in their stems. The cells in the interior tis-
2. intermembrane space sues of a leaf, called the mesophyll, can contain between
3. inner membrane (1+2+3: envelope) 450,000 and 800,000 chloroplasts for every square mil-
4. stroma (aqueous uid) limeter of leaf. The surface of the leaf is coated with a
5. thylakoid lumen (inside of thylakoid)
water-resistant waxy cuticle that protects the leaf from ex-
6. thylakoid membrane
cessive evaporation of water and decreases the absorption
7. granum (stack of thylakoids)
8. thylakoid (lamella) of ultraviolet or blue light to reduce heating. The trans-
9. starch parent epidermis layer allows light to pass through to the
10. ribosome palisade mesophyll cells where most of the photosynthe-
11. plastidial DNA sis takes place.
12. plastoglobule (drop of lipids)

Main articles: Chloroplast and Thylakoid

In photosynthetic bacteria, the proteins that gather light
for photosynthesis are embedded in cell membranes. In
its simplest form, this involves the membrane surround-
ing 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 ll most of the inte-
rior of a cell, giving the membrane a very large surface
area and therefore increasing the amount of light that the
bacteria can absorb.[21]
3 Light-dependent reactions
Main article: Light-dependent reactions
In the light-dependent reactions, one molecule of the
pigment chlorophyll absorbs one photon and loses one
electron. This electron is passed to a modied form of
chlorophyll called pheophytin, which passes the electron
to a quinone molecule, starting the ow of electrons down
an electron transport chain that leads to the ultimate re-
duction of NADP to NADPH. In addition, this creates a
4 3 LIGHT-DEPENDENT REACTIONS

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 com-
plex 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 func-
Light-dependent reactions of photosynthesis at the thylakoid tions to generate a chemiosmotic potential by pumping
membrane proton cations (H+ ) across the membrane and into the
thylakoid space. An ATP synthase enzyme uses that
proton gradient (energy gradient) across the chloroplast chemiosmotic potential to make ATP during photophos-
membrane, which is used by ATP synthase in the syn- phorylation, whereas NADPH is a product of the termi-
thesis of ATP. The chlorophyll molecule ultimately re- nal redox reaction in the Z-scheme. The electron enters a
gains the electron it lost when a water molecule is split chlorophyll molecule in Photosystem I. There it is further
in a process called photolysis, which releases a dioxygen excited by the light absorbed by that photosystem. The
(O2 ) molecule as a waste product. electron is then passed along a chain of electron accep-
tors to which it transfers some of its energy. The energy
The overall equation for the light-dependent reactions un- delivered to the electron acceptors is used to move hydro-
der the conditions of non-cyclic electron ow in green gen ions across the thylakoid membrane into the lumen.
plants is:[24] The electron is eventually used to reduce the co-enzyme
NADP with a H+ to NADPH (which has functions in the
2 H2 O + 2 NADP+ + 3 ADP + 3 P + light light-independent reaction); at that point, the path of that
2 NADPH + 2 H+ + 3 ATP + O2 electron ends.
The cyclic reaction is similar to that of the non-cyclic,
Not all wavelengths of light can support photosynthesis. but diers in that it generates only ATP, and no reduced
The photosynthetic action spectrum depends on the type NADP (NADPH) is created. The cyclic reaction takes
of accessory pigments present. For example, in green place only at photosystem I. Once the electron is dis-
plants, the action spectrum resembles the absorption placed from the photosystem, the electron is passed down
spectrum for chlorophylls and carotenoids with absorp- the electron acceptor molecules and returns to photosys-
tion peaks in violet-blue and red light. In red algae, the ac- tem I, from where it was emitted, hence the name cyclic
tion spectrum is blue-green light, which allows these algae reaction.
to use the blue end of the spectrum to grow in the deeper
waters that lter out the longer wavelengths (red light)
used by above ground green plants. The non-absorbed 3.2 Water photolysis
part of the light spectrum is what gives photosynthetic
organisms their color (e.g., green plants, red algae, pur- Main articles: Photodissociation and Oxygen evolution
ple bacteria) and is the least eective for photosynthesis
in the respective organisms.
The NADPH is the main reducing agent produced by
chloroplasts, which then goes on to provide a source of
3.1 Z scheme energetic electrons in other cellular reactions. Its pro-
duction leaves chlorophyll in photosystem I with a decit
of electrons (chlorophyll has been oxidized), which must
be balanced by some other reducing agent that will sup-
ply the missing electron. The excited electrons lost from
chlorophyll from photosystem I are supplied from the
electron transport chain by plastocyanin. However, since
photosystem II is the rst step of the Z-scheme, an ex-
ternal source of electrons is required to reduce its oxi-
The Z scheme dized chlorophyll a molecules. The source of electrons
in green-plant and cyanobacterial photosynthesis is wa-
In plants, light-dependent reactions occur in the thylakoid ter. Two water molecules are oxidized by four successive
membranes of the chloroplasts where they drive the syn- charge-separation reactions by photosystem II to yield
thesis of ATP and NADPH. The light-dependent reac- a molecule of diatomic oxygen and four hydrogen ions;
tions are of two forms: cyclic and non-cyclic. the electrons yielded are transferred to a redox-active
4.2 Carbon concentrating mechanisms 5

tyrosine residue that then reduces the oxidized chloro- ADP

phyll a (called P680) that serves as the primary light-

driven electron donor in the photosystem II reaction cen-
ATP
ter. That photo receptor is in eect reset and is then able
to repeat the absorption of another photon and the re-
lease of another photo-dissociated electron. The oxida-
tion of water is catalyzed in photosystem II by a redox-
ATP
active structure that contains four manganese ions and a
calcium ion; this oxygen-evolving complex binds two wa-
ter molecules and contains the four oxidizing equivalents ADP

that are used to drive the water-oxidizing reaction. Photo-

system II is the only known biological enzyme that carries
NADPH
out this oxidation of water. The hydrogen ions released
contribute to the transmembrane chemiosmotic potential
that leads to ATP synthesis. Oxygen is a waste product NADP+

of light-dependent reactions, but the majority of organ-

isms on Earth use oxygen for cellular respiration, includ-
ing photosynthetic organisms.[25][26] Overview of the Calvin cycle and carbon xation

triose phosphate. Most (5 out of 6 molecules) of the

4 Light-independent reactions glyceraldehyde 3-phosphate produced is used to regen-
erate ribulose 1,5-bisphosphate so the process can con-
4.1 Calvin cycle tinue. The triose phosphates not thus recycled often
condense to form hexose phosphates, which ultimately
Main articles: Calvin cycle, Carbon xation, and Light- yield sucrose, starch and cellulose. The sugars produced
independent reactions during carbon metabolism yield carbon skeletons that can
be used for other metabolic reactions like the production
In the light-independent (or dark) reactions, the enzyme of amino acids and lipids.
RuBisCO captures CO2 from the atmosphere and, in a
process called the Calvin-Benson cycle, it uses the newly
formed NADPH and releases three-carbon sugars, which 4.2 Carbon concentrating mechanisms
are later combined to form sucrose and starch. The over-
all equation for the light-independent reactions in green 4.2.1 On land
plants is[24]:128
In hot and dry conditions, plants close their stomata to
3 CO2 + 9 ATP + 6 NADPH + 6 H + prevent water loss. Under these conditions, CO2 will de-
C3 H6 O3 -phosphate + 9 ADP + 8 P + 6 crease and oxygen gas, produced by the light reactions
+
NADP + 3 H2 O of photosynthesis, will increase, causing an increase of
photorespiration by the oxygenase activity of ribulose-
Carbon xation produces the intermediate three-carbon 1,5-bisphosphate carboxylase/oxygenase and decrease in
carbon xation. Some plants have evolved mechanisms to
sugar product, which is then converted to the nal carbo-
hydrate products. The simple carbon sugars produced by increase the[27] CO2 concentration in the leaves under these
conditions.
photosynthesis are then used in the forming of other or-
ganic compounds, such as the building material cellulose, Main article: C4 carbon xation
the precursors for lipid and amino acid biosynthesis, or as
a fuel in cellular respiration. The latter occurs not only in Plants that use the C carbon xation process chemically
4
plants but also in animals when the energy from plants is x carbon dioxide in the cells of the mesophyll by adding
passed through a food chain. it to the three-carbon molecule phosphoenolpyruvate
The xation or reduction of carbon dioxide is a pro- (PEP), a reaction catalyzed by an enzyme called
cess in which carbon dioxide combines with a ve-carbon PEP carboxylase, creating the four-carbon organic acid
sugar, ribulose 1,5-bisphosphate, to yield two molecules oxaloacetic acid. Oxaloacetic acid or malate synthe-
of a three-carbon compound, glycerate 3-phosphate, also sized by this process is then translocated to specialized
known as 3-phosphoglycerate. Glycerate 3-phosphate, bundle sheath cells where the enzyme RuBisCO and other
in the presence of ATP and NADPH produced during Calvin cycle enzymes are located, and where CO2 re-
the light-dependent stages, is reduced to glyceraldehyde leased by decarboxylation of the four-carbon acids is
3-phosphate. This product is also referred to as 3- then xed by RuBisCO activity to the three-carbon 3-
phosphoglyceraldehyde (PGAL) or, more generically, as phosphoglyceric acids. The physical separation of Ru-
6 6
Overview of C4 carbon xation
BisCO from the oxygen-generating light reactions re-
duces photorespiration and increases CO2 xation 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 car-
bon xation 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
xation, compared to 3% that use C4 carbon xation;[29]
however, the evolution of C4 in over 60 plant lineages
makes it a striking example of convergent evolution.[27]
Main article: CAM photosynthesis
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 x-
ation to PEP from the Calvin cycle, CAM temporally
separates these two processes. CAM plants have a dif-
ferent leaf anatomy from C3 plants, and x the CO2 at
night, when their stomata are open. CAM plants store
the CO2 mostly in the form of malic acid via carboxy-
lation of phosphoenolpyruvate to oxaloacetate, which is
then reduced to malate. Decarboxylation of malate dur-
ing the day releases CO2 inside the leaves, thus allowing
carbon xation to 3-phosphoglycerate by RuBisCO. Six-
teen thousand species of plants use CAM.[30]
EFFICIENCY
4.2.2 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 (HCO
3). Before the CO2 diuses out it is quickly sponged
up by RuBisCO, which is concentrated within the car-
boxysomes. HCO
3 ions are made from CO2 outside the cell by another car-
bonic 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 anhy-
drase. This causes the HCO
3 ions to accumulate within the cell from where they dif-
fuse into the carboxysomes.[31] Pyrenoids in algae and
hornworts also act to concentrate CO2 around rubisco.[32]
5 Order and kinetics
The overall process of photosynthesis takes place in four
stages:[13]
6 Eciency
Probability distribution resulting from one-dimensional discrete
time random walks. The quantum walk created using the
Hadamard coin is plotted (blue) vs a classical walk (red) after
50 time steps.
Main article: Photosynthetic eciency
Plants usually convert light into chemical energy with
a photosynthetic eciency of 36%.[33] Absorbed light
that is unconverted is dissipated primarily as heat, with a
small fraction (12%)[34] re-emitted as chlorophyll uo-
rescence at longer (redder) wavelengths. A fact that al-
lows measurement of the light reaction of photosynthesis
by using chlorophyll uorometers.[35]

Actual plants photosynthetic eciency varies with the
frequency of the light being converted, light intensity,
temperature and proportion of carbon dioxide in the at-
mosphere, and can vary from 0.1% to 8%.[36] By com-
parison, solar panels convert light into electric energy at
an eciency of approximately 620% for mass-produced
panels, and above 40% in laboratory devices.
The eciency of both light and dark reactions can be
measured but the relationship between the two can be
complex.[37] For example, the ATP and NADPH en-
ergy molecules, created by the light reaction, can be
used for carbon xation or for photorespiration in C3
plants.[37] Electrons may also ow to other electron
sinks.[38][39][40] For this reason, it is not uncommon for
authors to dierentiate between work done under non-
photorespiratory conditions and under photorespiratory
conditions.[41][42][43]
Chlorophyll uorescence of photosystem II can measure
the light reaction, and Infrared gas analyzers can mea-
sure the dark reaction.[44] It is also possible to investi-
gate both at the same time using an integrated chlorophyll
uorometer and gas exchange system, or by using two
separate systems together.[45] Infrared gas analyzers and
some moisture sensors are sensitive enough to measure
the photosynthetic assimilation of CO2 , and of H2 O
using reliable methods[46] CO2 is commonly measured
in mols/m2 /s1 , parts per million or volume per million
and H2 0 is commonly measured in mmol/m2 /s1 or in
mbars.[46] By measuring CO2 assimilation, H2 O, leaf
temperature, barometric pressure, leaf area, and photo-
synthetically active radiation or PAR, it becomes possible
to estimate, A or carbon assimilation, E or transpira-
tion, gs or stomatal conductance, and Ci or intracellular
CO2 .[46] However, it is more common to used chlorophyll
uorescence for plant stress measurement, where appro- 7 Evolution
priate, because the most commonly used measuring pa-
rameters FV/FM and Y(II) or F/FM can be made in a
few seconds, allowing the measurement of larger plant Life timeline
populations.[43]
Gas exchange systems that oer control of CO2 levels,
above and below ambient, allow the common practice of 4000
measurement of A/Ci curves, at dierent CO2 levels, to
characterize a plants photosynthetic response.[46]
Integrated chlorophyll uorometer gas exchange sys-
tems allow a more precise measure of photosynthetic re- 3000
sponse and mechanisms.[44][47] While standard gas ex-
change photosynthesis systems can measure Ci, or sub- 2500
stomatal CO2 levels, the addition of integrated chloro-
phyll uorescence measurements allows a more precise 2000
measurement of CC to replace Ci.[45][48] The estima-
tion of CO2 at the site of carboxylation in the chloro- 1500
plast, or CC, becomes possible with the measurement
of mesophyll conductance or g using an integrated 1000
system.[44][45][49]
Photosynthesis measurement systems are not designed to
directly measure the amount of light absorbed by the
7
leaf. But analysis of chlorophyll-uorescence, P700- and
P515-absorbance and gas exchange measurements reveal
detailed information about e.g. the photosystems, quan-
tum eciency and the CO2 assimilation rates. With some
instruments even wavelength-dependency of the photo-
synthetic eciency can be analyzed.[50]
A phenomenon known as quantum walk increases the ef-
ciency of the energy transport of light signicantly. 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 photo-
complex. 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 col-
lection of molecules that traps its energy in a chemical
form that makes it accessible for the cells metabolism.
The excitons wave properties enable it to cover a wider
area and try out several possible paths simultaneously, al-
lowing it to instantaneously choose the most ecient
route, where it will have the highest probability of arriv-
ing at its destination in the minimum possible time. Be-
cause 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 proper-
ties 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.[51][52][53]
view discuss
4500
3500
500
0
8 7 EVOLUTION

water A
Single-celled r
life c
photosynthesis h
Eukaryotes e
Multicellular a
life n
Land life
Dinosaurs
H
Mammals
a
Flowers
d
e
Earliest Earth (4540) a
n
Earliest water

Earliest life Pongola

LHB meteorites Huronian

Earliest oxygen
Cryogenian
Atmospheric oxygen
Andean
Oxygen crisis

Karoo
Earliest sexual reproduction

Ediacara biota Quaternary

Cambrian explosion Axis scale: millions of years.

Orange labels: known ice ages.
Earliest humans Also see: Human timeline and Nature timeline
Main article: Evolution of photosynthesis

P
Early photosynthetic systems, such as those in green and
h
purple sulfur and green and purple nonsulfur bacteria,
a
are thought to have been anoxygenic, and used various
n
other molecules as electron donors rather than water.
e
Green and purple sulfur bacteria are thought to have used
r
hydrogen and sulfur as electron donors. Green nonsul-
o
fur bacteria used various amino and other organic acids
z
as an electron donor. Purple nonsulfur bacteria used a
o
variety of nonspecic organic molecules. The use of
i
these molecules is consistent with the geological evidence
c
that Earths early atmosphere was highly reducing at that
time.[54]
P
r Fossils of what are thought to be lamentous photosyn-
o thetic organisms have been dated at 3.4 billion years
t old.[55][56]
e The main source of oxygen in the Earths atmosphere de-
r rives from oxygenic photosynthesis, and its rst appear-
o ance is sometimes referred to as the oxygen catastrophe.
z Geological evidence suggests that oxygenic photosynthe-
o sis, such as that in cyanobacteria, became important dur-
i ing the Paleoproterozoic era around 2 billion years ago.
c Modern photosynthesis in plants and most photosynthetic
7.2 Cyanobacteria and the evolution of photosynthesis 9

prokaryotes is oxygenic. Oxygenic photosynthesis uses 7.2 Cyanobacteria and the evolution of
water as an electron donor, which is oxidized to molecu- photosynthesis
lar oxygen (O
2) in the photosynthetic reaction center. The biochemical capacity to use water as the source for
electrons in photosynthesis evolved once, in a common
ancestor of extant cyanobacteria. The geological record
7.1 Symbiosis and the origin of chloro- indicates that this transforming event took place early
plasts in Earths history, at least 24502320 million years ago
(Ma), and, it is speculated, much earlier.[64][65] Because
the Earths atmosphere contained almost no oxygen dur-
ing the estimated development of photosynthesis, it is
believed that the rst photosynthetic cyanobacteria did
not generate oxygen.[66] Available evidence from geo-
biological studies of Archean (>2500 Ma) sedimentary
rocks indicates that life existed 3500 Ma, but the question
of when oxygenic photosynthesis evolved is still unan-
swered. A clear paleontological window on cyanobac-
terial evolution opened about 2000 Ma, revealing an
already-diverse biota of blue-green algae. Cyanobacteria
remained the principal primary producers of oxygen
throughout the Proterozoic Eon (2500543 Ma), in part
because the redox structure of the oceans favored pho-
toautotrophs capable of nitrogen xation. Green algae
joined blue-green algae as the major primary produc-
Plant cells with visible chloroplasts (from a moss, Plagiomnium ers of oxygen on continental shelves near the end of the
ane) Proterozoic, but it was only with the Mesozoic (25165
Ma) radiations of dinoagellates, coccolithophorids, and
Several groups of animals have formed symbiotic rela- diatoms did the primary production of oxygen in marine
tionships with photosynthetic algae. These are most com- shelf waters take modern form. Cyanobacteria remain
mon in corals, sponges and sea anemones. It is presumed critical to marine ecosystems as primary producers of
that this is due to the particularly simple body plans and oxygen in oceanic gyres, as agents of biological nitrogen
large surface areas of these animals compared to their xation, and, in modied form, as the plastids of marine
volumes.[57] In addition, a few marine mollusks Elysia algae.[67]
viridis and Elysia chlorotica also maintain a symbiotic re-
lationship with chloroplasts they capture from the algae
in their diet and then store in their bodies. This allows 8 Discovery
the mollusks to survive solely by photosynthesis for sev-
eral months at a time.[58][59] Some of the genes from the
plant cell nucleus have even been transferred to the slugs, Although some of the steps in photosynthesis are still not
completely understood, the overall photosynthetic equa-
so that the chloroplasts can be supplied with proteins that
they need to survive. [60] tion has been known since the 19th century.

An even closer form of symbiosis may explain the ori- Jan van Helmont began the research of the process in the
gin of chloroplasts. Chloroplasts have many similar- mid-17th century when he carefully measured the mass
ities with photosynthetic bacteria, including a circular of the soil used by a plant and the mass of the plant as
chromosome, prokaryotic-type ribosome, and similar it grew. After noticing that the soil mass changed very
proteins in the photosynthetic reaction center.[61][62] The little, he hypothesized that the mass of the growing plant
endosymbiotic theory suggests that photosynthetic bac- must come from the water, the only substance he added to
teria were acquired (by endocytosis) by early eukaryotic the potted plant. His hypothesis was partially accurate
cells to form the rst plant cells. Therefore, chloroplasts much of the gained mass also comes from carbon dioxide
may be photosynthetic bacteria that adapted to life in- as well as water. However, this was a signaling point to
side plant cells. Like mitochondria, chloroplasts possess the idea that the bulk of a plants biomass comes from the
their own DNA, separate from the nuclear DNA of their inputs of photosynthesis, not the soil itself.
plant host cells and the genes in this chloroplast DNA re- Joseph Priestley, a chemist and minister, discovered that,
semble those found in cyanobacteria.[63] DNA in chloro- when he isolated a volume of air under an inverted jar,
plasts codes for redox proteins such as those found in and burned a candle in it, the candle would burn out very
the photosynthetic reaction centers. The CoRR Hypoth- quickly, much before it ran out of wax. He further dis-
esis proposes that this Co-location is required for Redox covered that a mouse could similarly injure air. He then
Regulation. showed that the air that had been injured by the candle
10 8 DISCOVERY

and the mouse could be restored by a plant.

In 1778, Jan Ingenhousz, repeated Priestleys experi-
ments. He discovered that it was the inuence of sun-
light on the plant that could cause it to revive a mouse in
a matter of hours.
In 1796, Jean Senebier, a Swiss pastor, botanist, and nat-
uralist, demonstrated that green plants consume carbon
dioxide and release oxygen under the inuence of light.
Soon afterward, Nicolas-Thodore 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 incorpo-
ration of water. Thus, the basic reaction by which pho-
tosynthesis is used to produce food (such as glucose) was
outlined.
Cornelis Van Niel made key discoveries explaining the
chemistry of photosynthesis. By studying purple sulfur
bacteria and green bacteria he was the rst to demonstrate
that photosynthesis is a light-dependent redox reaction, in
which hydrogen reduces carbon dioxide.
Robert Emerson discovered two light reactions by testing
plant productivity using dierent wavelengths of light.
With the red alone, the light reactions were suppressed.
When blue and red were combined, the output was much Melvin Calvin works in his photosynthesis laboratory.
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 topes to determine that the oxygen liberated in photosyn-
PSI. PSI contains only chlorophyll a, PSII contains pri- thesis came from the water.
marily chlorophyll a with most of the available chloro- Melvin Calvin and Andrew Benson, along with James
phyll b, among other pigment. These include phyco- Bassham, elucidated the path of carbon assimilation (the
bilins, which are the red and blue pigments of red and photosynthetic carbon reduction cycle) in plants. The
blue algae respectively, and fucoxanthol for brown algae carbon reduction cycle is known as the Calvin cycle,
and diatoms. The process is most productive when the which ignores the contribution of Bassham and Benson.
absorption of quanta are equal in both the PSII and PSI, Many scientists refer to the cycle as the Calvin-Benson
assuring that input energy from the antenna complex is Cycle, Benson-Calvin, and some even call it the Calvin-
divided between the PSI and PSII system, which in turn Benson-Bassham (or CBB) Cycle.
powers the photochemistry.[13]
Nobel Prize-winning scientist Rudolph A. Marcus was
Robert Hill thought that a complex of reactions consist- able to discover the function and signicance of the elec-
ing of an intermediate to cytochrome b6 (now a plasto- tron transport chain.
quinone), another is from cytochrome f to a step in the
carbohydrate-generating mechanisms. These are linked Otto Heinrich Warburg and Dean Burk discovered the I-
by plastoquinone, which does require energy to reduce quantum photosynthesis reaction that splits the CO2 , ac-
cytochrome f for it is a sucient reductant. Further tivated by the respiration.[69]
experiments to prove that the oxygen developed dur- Louis N.M. Duysens and Jan Amesz discovered that
ing the photosynthesis of green plants came from wa- chlorophyll a will absorb one light, oxidize cytochrome
ter, were performed by Hill in 1937 and 1939. He f, chlorophyll a (and other pigments) will absorb another
showed that isolated chloroplasts give o oxygen in the light, but will reduce this same oxidized cytochrome, stat-
presence of unnatural reducing agents like iron oxalate, ing the two light reactions are in series.
ferricyanide or benzoquinone after exposure to light. The
Hill reaction[68] is as follows:
8.1 Development of the concept
2 H2 O + 2 A + (light, chloroplasts) 2 AH2
+ O2
In 1893, Charles Reid Barnes proposed two terms, pho-
tosyntax and photosynthesis, for the biological process of
where A is the electron acceptor. Therefore, in light, the synthesis of complex carbon compounds out of carbonic
electron acceptor is reduced and oxygen is evolved. acid, in the presence of chlorophyll, under the inuence
Samuel Ruben and Martin Kamen used radioactive iso- of light. Over time, the term photosynthesis came into
 11

common usage as the term of choice. Later discovery of carbon PGA acid. At 1000 ppm CO2 in measuring air,
anoxygenic photosynthetic bacteria and photophosphory- both the C3 and C4 plants had similar leaf photosynthetic
lation necessitated redenition of the term.[70] rates around 60 u mole CO2/square meter.sec. indicating
the suppression of phototorespiration in C3 plants.[76][77]

8.2 C3 : C4 photosynthesis research

9 Factors
After WWII at late 1940 at the University of Cali-
fornia, 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 iso-
tope and paper chromatography techniques.[71] The path-
way of CO2 xation by the algae Chlorella in a frac-
tion 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 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 u
mole CO2/square metere.sec., with the conclusion that
all terrestrial plants having the same photosynthetic ca-
pacities that were light saturated at less than 50% of The leaf is the primary site of photosynthesis in plants.
sunlight.[72][73] These rates were determined in potted
plants grown indoors under low light intensity. There are three main factors aecting photosynthesis and
Later in 1958-1963 at Cornell University, eld grown several corollary factors. The three main are:
maize was reported to have much greater leaf photo-
synthetic rates of 40 u mol CO2/square meter.sec and Light irradiance and wavelength
was not saturated at near full sunlight.[74][75] This higher
Carbon dioxide concentration
rate in maize was almost double those observed in other
species such as wheat and soybean, indicating that large Temperature.
dierences in photosynthesis exist among higher plants.
At the University of Arizona, detailed gas exchange re-
search on more than 15 species of monocot and dicot un- 9.1 Light intensity (irradiance), wave-
covered for the rst time that dierences in leaf anatomy length and temperature
are crucial factors in dierentiating photosynthetic ca-
pacities among species.[76][77] In tropical grasses, includ- See also: PI (photosynthesis-irradiance) curve
ing maize, sorghum, sugarcane, Bermuda grass and in the The process of photosynthesis provides the main input
dicot amaranthus, leaf photosynthetic rates were around of free energy into the biosphere, and is one of four main
3840 u mol CO2/square meter.sec., and the leaves have ways in which radiation is important for plant life.[83]
two types of green cells, i. e. outer layer of mes-
ophyll cells surrounding a tightly packed cholorophyl- The radiation climate within plant communities is ex-
lous vascular bundle sheath cells. This type of anatomy tremely variable, with both time and space.
was termed Kranz anatomy in the 19th century by the In the early 20th century, Frederick Blackman and
botanist Gottlieb Haberlandt while studying leaf anatomy Gabrielle Matthaei investigated the eects of light inten-
of sugarcane.[78] Plant species with the greatest photo- sity (irradiance) and temperature on the rate of carbon
synthetic rates and Kranz anatomy showed no appar- assimilation.
ent photorespiration, very low CO2 compensation point,
high optimum temperature, high stomatal resistances and At constant temperature, the rate of carbon assim-
lower mesophyll resistances for gas diusion and rates ilation varies with irradiance, increasing as the ir-
never saturated at full sun light.[79] The research at Ari- radiance increases, but reaching a plateau at higher
zona was designated Citation Classic by the ISI 1986.[77] irradiance.
These species was later termed C4 plants as the rst sta-
ble compound of CO2 xation in light has 4 carbon as At low irradiance, increasing the temperature has lit-
malate and aspartate.[80][81][82] Other species that lack tle inuence on the rate of carbon assimilation. At
Kranz anatomy were termed C3 type such as cotton and constant high irradiance, the rate of carbon assimi-
sunower, as the rst stable carbon compound is the 3- lation increases as the temperature is increased.
12
Chlorophyll b
Chlorophyll a
Absorbance
400 500 600 700
Wavelength [nm]
400 500 600
Absorbance spectra of free chlorophyll a (green) and b (red) in a
solvent. The action spectra of chlorophyll molecules are slightly
modied in vivo depending on specic pigment-protein interac-
tions.
These two experiments illustrate several important
points: First, it is known that, in general, photochemical
reactions are not aected by temperature. However,
these experiments clearly show that temperature af-
fects the rate of carbon assimilation, so there must be
two sets of reactions in the full process of carbon as-
similation. These are, of course, the light-dependent
'photochemical' temperature-independent stage, and the
light-independent, temperature-dependent stage. Sec-
ond, Blackmans experiments illustrate the concept of
limiting factors. Another limiting factor is the wavelength
of light. Cyanobacteria, which reside several meters un-
derwater, cannot receive the correct wavelengths required
to cause photoinduced charge separation in conventional
photosynthetic pigments. To combat this problem, a se-
ries of proteins with dierent pigments surround the re-
action center. This unit is called a phycobilisome.
9.2 Carbon dioxide levels and photorespi- oxygenase activity is more commonly known as
ration photorespiration, since it is characterized by light-
Photorespiration
As carbon dioxide concentrations rise, the rate at which
10 SEE ALSO
sugars are made by the light-independent reactions in-
creases until limited by other factors. RuBisCO,
the enzyme that captures carbon dioxide in the light-
independent reactions, has a binding anity for both car-
bon dioxide and oxygen. When the concentration of car-
bon dioxide is high, RuBisCO will x carbon dioxide.
However, if the carbon dioxide concentration is low, Ru-
BisCO 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 phosphoglyco-
700 late (2 carbon) instead of 3-phosphoglycerate (3 car-
bon). Phosphoglycolate cannot be metabolized by
the Calvin-Benson cycle and represents carbon lost
from the cycle. A high oxygenase activity, there-
fore, 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 glyco-
late 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 pro-
duce ammonia (NH3 ), which is able to diuse out of
the plant, leading to a loss of nitrogen.
A highly simplied summary is:
2 glycolate + ATP 3-
phosphoglycerate + car-
bon dioxide + ADP +
NH3
The salvaging pathway for the products of RuBisCO
dependent oxygen consumption and the release of
carbon dioxide.
10 See also
Jan Anderson (scientist)
Articial photosynthesis
Calvin-Benson cycle
Carbon xation
Cellular respiration
 13

Chemosynthesis [9] Whitmarsh J, Govindjee (1999). The photosynthetic

process. In Singhal GS, Renger G, Sopory SK, Irrgang
Integrated uorometer KD, Govindjee. Concepts in photobiology: photosynthe-
sis and photomorphogenesis. Boston: Kluwer Academic
Light-dependent reaction Publishers. pp. 1151. ISBN 0-7923-5519-9. 1001015
grams of carbon/year xed by photosynthetic organisms,
Organic reaction
which is equivalent to 41018 kJ/yr = 41021 J/yr of free
Photobiology energy stored as reduced carbon.

Photoinhibition [10] Steger U, Achterberg W, Blok K, Bode H, Frenz W,

Gather C, Hanekamp G, Imboden D, Jahnke M, Kost M,
Photosynthetic reaction center Kurz R, Nutzinger HG, Ziesemer T (2005). Sustainable
development and innovation in the energy sector. Berlin:
Photosynthetically active radiation Springer. p. 32. ISBN 3-540-23103-X. The average
global rate of photosynthesis is 130 TW.
Photosystem
[11] World Consumption of Primary Energy by Energy Type
Photosystem I and Selected Country Groups, 19802004. Energy In-
formation Administration. July 31, 2006. Archived from
Photosystem II the original (XLS) on November 9, 2006. Retrieved
2007-01-20.
Quantum biology
[12] Field CB, Behrenfeld MJ, Randerson JT, Falkowski P (Jul
Red edge 1998). Primary production of the biosphere: integrating
terrestrial and oceanic components. Science. 281 (5374):
Vitamin D
23740. Bibcode:1998Sci...281..237F. PMID 9657713.
Hill reaction doi:10.1126/science.281.5374.237.

[13] Photosynthesis. McGraw-Hill Encyclopedia of Science

& Technology. 13. New York: McGraw-Hill. 2007.
11 References ISBN 0-07-144143-3.

[14] Whitmarsh J, Govindjee (1999). Chapter 2: The Ba-

[1] photosynthesis. Online Etymology Dictionary.
[2] . Liddell, Henry George; Scott, Robert; A Greek
English Lexicon at the Perseus Project
[3] . Liddell, Henry George; Scott, Robert; A
GreekEnglish Lexicon at the Perseus Project
[4] Bryant DA, Frigaard NU (Nov 2006). Prokaryotic
photosynthesis and phototrophy illuminated. Trends
in Microbiology. 14 (11): 48896. PMID 16997562.
doi:10.1016/j.tim.2006.09.001.
[5] Reece J, Urry L, Cain M, Wasserman S, Minorsky P,
Jackson R. Biology (International ed.). Upper Saddle
River, NJ: Pearson Education. pp. 235, 244. ISBN 0-
321-73975-2. This initial incorporation of carbon into
organic compounds is known as carbon xation.
[6] Olson JM (May 2006). Photosynthesis in the Archean
era. Photosynthesis Research. 88 (2): 10917. PMID
16453059. doi:10.1007/s11120-006-9040-5.
[7] Buick R (Aug 2008). When did oxygenic pho-
tosynthesis evolve?". Philosophical Transactions of
the Royal Society of London, Series B. 363 (1504):
273143. PMC 2606769 . PMID 18468984.
doi:10.1098/rstb.2008.0041.
[8] Nealson KH, Conrad PG (Dec 1999). Life: past,
present and future. Philosophical Transactions of
the Royal Society of London, Series B. 354 (1392):
192339. PMC 1692713 . PMID 10670014.
doi:10.1098/rstb.1999.0532.
sic Photosynthetic Process. In Singhal GS, Renger G,
Sopory SK, Irrgang KD, Govindjee. Concepts in Photo-
biology: Photosynthesis and Photomorphogenesis. Boston:
Kluwer Academic Publishers. p. 13. ISBN 978-0-7923-
5519-9.
[15] Anaerobic Photosynthesis, Chemical & Engineering
News, 86, 33, August 18, 2008, p. 36
[16] Kulp TR, Hoeft SE, Asao M, Madigan MT, Hol-
libaugh JT, Fisher JC, Stolz JF, Culbertson CW, Miller
LG, Oremland RS (Aug 2008). Arsenic(III) fuels
anoxygenic photosynthesis in hot spring biolms from
Mono Lake, California. Science. 321 (5891): 967
70. Bibcode:2008Sci...321..967K. PMID 18703741.
doi:10.1126/science.1160799.
[17] Scientists discover unique microbe in Californias largest
lake. Retrieved 2009-07-20.
[18] Plants: Diversity and Evolution, page 14, Martin In-
grouille, Bill Eddie
[19] Evolution of Photosynthesis
[20] Tavano CL, Donohue TJ (Dec 2006). Development of
the bacterial photosynthetic apparatus. Current Opinion
in Microbiology. 9 (6): 62531. PMC 2765710 . PMID
17055774. doi:10.1016/j.mib.2006.10.005.
[21] Mullineaux CW (1999). The thylakoid membranes
of cyanobacteria: structure, dynamics and function.
Australian Journal of Plant Physiology. 26 (7): 671677.
doi:10.1071/PP99027.
14
[22] Sener MK, Olsen JD, Hunter CN, Schulten K (Oct
2007). Atomic-level structural and functional model
of a bacterial photosynthetic membrane vesicle.
Proceedings of the National Academy of Sciences of
the United States of America. 104 (40): 157238.
Bibcode:2007PNAS..10415723S. PMC 2000399 .
PMID 17895378. doi:10.1073/pnas.0706861104.
[23] Campbell NA, Williamson B, Heyden RJ (2006). Biology
Exploring Life. Upper Saddle River, NJ: Prentice Hall.
ISBN 0-13-250882-6.
[24] Raven PH, Evert RF, Eichhorn SE (2005). Biology of
Plants, (7th ed.). New York: W. H. Freeman and Com-
pany. pp. 124127. ISBN 0-7167-1007-2.
[25] Yachandra Group Home page.
[26] Pushkar Y, Yano J, Sauer K, Boussac A, Yachandra VK
(Feb 2008). Structural changes in the Mn4Ca clus-
ter and the mechanism of photosynthetic water split-
ting. Proceedings of the National Academy of Sci-
ences of the United States of America. 105 (6): 1879
84. Bibcode:2008PNAS..105.1879P. PMC 2542863 .
PMID 18250316. doi:10.1073/pnas.0707092105.
[27] Williams BP, Johnston IG, Covsho S, Hibberd JM
(September 2013). Phenotypic landscape inference re-
veals multiple evolutionary paths to C4 photosynthesis.
eLife. 2: e00961. PMC 3786385 . PMID 24082995.
doi:10.7554/eLife.00961.
[28] Taiz L, Geiger E (2006). Plant Physiology (4th ed.).
Sinauer Associates. ISBN 978-0-87893-856-8.
[29] Monson RK, Sage RF (1999). The Taxonomic Distri-
bution of C4 Photosynthesis. C plant biology. Boston:
Academic Press. pp. 551580. ISBN 0-12-614440-0.
[30] Dodd AN, Borland AM, Haslam RP, Griths
H, Maxwell K (Apr 2002). Crassulacean acid
metabolism: plastic, fantastic. Journal of Experi-
mental Botany. 53 (369): 56980. PMID 11886877.
doi:10.1093/jexbot/53.369.569.
[31] Badger MR, Price GD (Feb 2003). CO2 concentrat-
ing mechanisms in cyanobacteria: molecular compo-
nents, their diversity and evolution. Journal of Exper-
imental Botany. 54 (383): 60922. PMID 12554704.
doi:10.1093/jxb/erg076.
[32] Badger MR, Andrews JT, Whitney SM, Ludwig M, Yel-
lowlees DC, Leggat W, Price GD (1998). The diver-
sity and coevolution of Rubisco, plastids, pyrenoids, and
chloroplast-based CO2 -concentrating mechanisms in al-
gae. Canadian Journal of Botany. 76 (6): 10521071.
doi:10.1139/b98-074.
[33] Miyamoto K. Chapter 1 Biological energy production.
Renewable biological systems for alternative sustainable
energy production (FAO Agricultural Services Bulletin
128). Food and Agriculture Organization of the United
Nations. Retrieved 2009-01-04.
[34] Maxwell K, Johnson GN (Apr 2000). Chlorophyll
uorescence--a practical guide. Journal of Experimen-
tal Botany. 51 (345): 65968. PMID 10938857.
doi:10.1093/jexbot/51.345.659.
11 REFERENCES
[35] Maxwell K, Johnson GN (2000). Chloro-
phyll uorescence a practical guide. Jour-
nal of Experimental Botany. 51 (345): 659668.
doi:10.1093/jxb/51.345.659.
[36] Govindjee R. What is Photosynthesis. Biology at Illi-
nois.
[37] Rosenqvist E., van Kooten O., (2006) Chlorophyll Flu-
orescence: A General Description and Nomenclature.
From Chapter 2 Practical Applications of Chlorophyll
Fluorescence in Plant Biology. by Jennifer R. DeEll
(Editor), Peter M.A. Toivonen (Editor) Kluwer Academic
Publishers group, P.O Box 322, 3300 A.H. Dordrecht, the
Netherlands, pages 39-78
[38] Baker N. R., Oxborough K., (2004) Chlorophyll uores-
cence as a probe of photosynthetic productivity. From
Chapter 3, Chlorophylla Fluorescence a Signature of
Photosynthesis, edited by George Papaqeorgiou and
Govindjee, published by Springer 2004, PO Box17, 3300
AA Dordrecht, The Netherlands, pages 66-79
[39] Flexas 1999 Water stress induces dierent levels
of photosynthesis and electron transport rate regulation
in grapevines"J. FLEXAS, J. M. ESCALONA & H.
MEDRANO Plant, Cell & Environment Volume 22 Is-
sue 1 Page 39-48, January 1999
[40] Fryer M. J., Andrews J.R., Oxborough K., Blowers D.
A., Baker N.E. (1998) Relationship between CO2 assim-
ilation, photosynthetic electron transport and active 02
metabolism in leaves of maize in the eld during periods
of low temperature
[41] Earl H, Said Ennahli S (2004). Estimating photosyn-
thetic electron transport via chlorophyll uorometry with-
out Photosystem II light saturation. Photosynthesis Re-
search. 82 (177186): 2004.
[42] Genty B, Briantais J. M. & Baker N. R. (1989) The re-
lationship between the quantum yield of photosynthetic
electron transport and quenching of chlorophyll uores-
cence. Biochimica et Biophysica Acta. 990: 8792.
doi:10.1016/s0304-4165(89)80016-9.
[43] Baker NR (2008). Chlorophyll Fluorescence: A
Probe of Photosynthesis In Vivo. Annu. Rev.
Plant Biol. 59: 89113. PMID 18444897.
doi:10.1146/annurev.arplant.59.032607.092759.
[44] Bernacchi CJ, Portis A. R., Nakano H., von Caemmerer
S., and Long S.P. (2002) Temperature Response of Mes-
ophyll Conductance. Implications for the Determination
of Rubisco Enzyme Kinetics and for Limitations to Pho-
tosynthesis in Vivo. Plant Physiology. 130: 19921998.
doi:10.1104/pp.008250.
[45] Ribas-Carbo M., Flexas J., Robinson S.A., Tcherkez G.
G. B., (2010) In vivo measurement of plant respiration
University of Wollongong Research Online
[46] Long S.P., and Bernacchi C.J. (2003) Gas exchange mea-
surements, what can they tell us about the underlying lim-
itations to photosynthesis? Procedures and sources of er-
ror.

[47] Ribas-Carbo M., Flexas J., Robinson S.A., Tcherkez G.
G. B., (2010) In vivo measurement of plant respiration
University of Wollongong Research Online
[48] Bernacchi CJ, Portis A. R., Nakano H., von Caemmerer
S., and Long S.P. (2002) Temperature Response of Mes-
ophyll Conductance. Implications for the Determination
of Rubisco Enzyme Kinetics and for Limitations to Pho-
tosynthesis in Vivo. Plant Physiology. 130: 19921998.
doi:10.1104/pp.008250.
[49] YIN X, Struik PC (2009). Theoretical reconsidera-
tions when estimating the mesophyll conductanceto CO2
diusion in leaves of C3 plants by analysis of com-
bined gas exchange and chlorophyll uorescence mea-
surements pce_2016 1513..1. Plant, Cell and Envi-
ronment. 32: 15131524 [1524]. doi:10.1111/j.1365-
3040.2009.02016.x.
[50] Schreiber U, Klughammer C, Kolbowski J (2012).
Assessment of wavelength-dependent parameters of
photosynthetic electron transport with a new type of
multi-color PAM chlorophyll uorometer. Photosynthe-
sis research. 113 (13): 127144. doi:10.1007/s11120-
012-9758-1.
[51] Palmer J (21 June 2013). Plants 'seen doing quantum
physics". BBC News.
[52] Lloyd S (10 March 2014). Quantum Biology: Better Liv-
ing Through Quantum Mechanics - The Nature of Real-
ity. Nova: PBS Online, WGBH Boston.
[53] Hildner R, Brinks D, Nieder JB, Cogdell RJ, van Hulst
NF (Jun 2013). Quantum coherent energy transfer over
varying pathways in single light-harvesting complexes.
Science. 340 (6139): 144851. PMID 23788794.
doi:10.1126/science.1235820.
[54] Gale, Joseph (2009). Astrobiology of Earth : The emer-
gence, evolution and future of life on a planet in turmoil.
OUP Oxford. pp. 112113. ISBN 9780191548352.
[55] Photosynthesis got a really early start, New Scientist, 2
October 2004
[56] Revealing the dawn of photosynthesis, New Scientist, 19
August 2006
[57] Venn AA, Loram JE, Douglas AE (2008). Photo-
synthetic symbioses in animals. Journal of Experi-
mental Botany. 59 (5): 106980. PMID 18267943.
doi:10.1093/jxb/erm328.
[58] Rumpho ME, Summer EJ, Manhart JR (May 2000).
Solar-powered sea slugs. Mollusc/algal chloroplast sym-
biosis. Plant Physiology. 123 (1): 2938. PMC 1539252
. PMID 10806222. doi:10.1104/pp.123.1.29.
[59] Muscatine L, Greene RW (1973). Chloroplasts and
algae as symbionts in molluscs. International Review
of Cytology. International Review of Cytology. 36:
13769. ISBN 9780123643360. PMID 4587388.
doi:10.1016/S0074-7696(08)60217-X.
15
[60] Rumpho ME, Worful JM, Lee J, Kannan K, Tyler
MS, Bhattacharya D, Moustafa A, Manhart JR (Nov
2008). Horizontal gene transfer of the algal nuclear
gene psbO to the photosynthetic sea slug Elysia chlorot-
ica. Proceedings of the National Academy of Sciences
of the United States of America. 105 (46): 17867
71. Bibcode:2008PNAS..10517867R. PMC 2584685 .
PMID 19004808. doi:10.1073/pnas.0804968105.
[61] Douglas SE (Dec 1998). Plastid evolution: ori-
gins, diversity, trends. Current Opinion in Genetics
& Development. 8 (6): 65561. PMID 9914199.
doi:10.1016/S0959-437X(98)80033-6.
[62] Reyes-Prieto A, Weber AP, Bhattacharya D
(2007). The origin and establishment of the
plastid in algae and plants. Annual Review
of Genetics. 41: 14768. PMID 17600460.
doi:10.1146/annurev.genet.41.110306.130134.
[63] Raven JA, Allen JF (2003). Genomics and chloro-
plast evolution: what did cyanobacteria do for plants?".
Genome Biology. 4 (3): 209. PMC 153454 . PMID
12620099. doi:10.1186/gb-2003-4-3-209.
[64] Tomitani A, Knoll AH, Cavanaugh CM, Ohno T (Apr
2006). The evolutionary diversication of cyanobac-
teria: molecular-phylogenetic and paleontological per-
spectives. Proceedings of the National Academy of
Sciences of the United States of America. 103 (14):
54427. PMC 1459374 . PMID 16569695.
doi:10.1073/pnas.0600999103.
[65] Cyanobacteria: Fossil Record. Ucmp.berkeley.edu.
Retrieved 2010-08-26.
[66] Smith, Alison (2010). Plant biology. New York, NY:
Garland Science. p. 5. ISBN 0815340257.
[67] Herrero A, Flores E (2008). The Cyanobacteria: Molec-
ular Biology, Genomics and Evolution (1st ed.). Caister
Academic Press. ISBN 978-1-904455-15-8.
[68] Walker DA (2002). "'And whose bright presence' - an
appreciation of Robert Hill and his reaction (PDF). Pho-
tosynthesis Research. 73 (13): 514. PMID 16245102.
doi:10.1023/A:1020479620680.
[69] Otto Warburg Biography. Nobelprize.org (1970-08-
01). Retrieved on 2011-11-03.
[70] Gest, Howard (2002). History of the word photosynthe-
sis and evolution of its denition.. Photosynthesis Re-
search. 73 (13): 710. doi:10.1023/A:1020419417954.
[71] Calvin, Melvin (July 1989). Forty years of photosynthe-
sis and related activities. Photosynthesis Research. 21
(1). doi:10.1007/BF00047170.
[72] Verduin, J. 1953. A table of photosynthesis rates under
optimal, near natural conditions. Am.J. Bot. 40:675-679.
[73] Verduin, J., Whitwer, E. E., and Cowell, B.C. 1959. Max-
imal photosynthetic rates in nature, Science 130:268-269.
[74] Hesketh, J. D., and Musgrave, R.B. 1962. Photosynthesis
under eld conditions. IV. Light studies with individual
corn leaves. Crop Sci. 2:311-315.
16 13 EXTERNAL LINKS

[75] Hesketh, J.D., and Moss, D. N. 1963.Variation in the re-
sponse of photosynthesis to light. Crop Sci. 3:107-110.
[76] El-Sharkawy, M. A., and Hesketh, J. D. 1965. Photosyn-
thesis among species in relation to characteristics of leaf
anatomy and CO2 diusion resistances. Crop Sci. 5:517-
521.
[77] El-Sharkawy, M. A., and Hesketh, J. D. 1986.Ci-
tation Classic-Photosynthesis among species in rela-
tion to characteristics of leaf anatomy and CO2 diu-
sion resistances. Curr. Cont./Agr.Biol.Environ.27:14-
14. Online http://www.library.upenn/edu/classics1986/
A1986C691300001.pdf.
[78] Haberlandt, G. 1904.Physiologische Panzanatomie. En-
gelmann, Leipzig.
[79] El-Sharkawy, M. A. 1965.Factors Limiting Photosyn-
thetic Rates of Dierent Plant Species. Ph.D. Disserta-
tion, The University of Arizona, Tucson, USA.
[80] Karpilov YS (1960). The distribution of radioactvity
in carbon-14 among the products of photosynthesis in
maize. Proc. Kazan Agric. Inst. 14: 1524.
12.2 Papers
Gupta RS, Mukhtar T, Singh B (Jun 1999).
Evolutionary relationships among photosynthetic
prokaryotes (Heliobacterium chlorum, Chloroexus
aurantiacus, cyanobacteria, Chlorobium tepidum
and proteobacteria): implications regarding the
origin of photosynthesis. Molecular Microbi-
ology. 32 (5): 893906. PMID 10361294.
doi:10.1046/j.1365-2958.1999.01417.x.
Rutherford AW, Faller P (Jan 2003). Photosystem
II: evolutionary perspectives. Philosophical Trans-
actions of the Royal Society of London. Se-
ries B, Biological Sciences. 358 (1429): 245
53. PMC 1693113 . PMID 12594932.
doi:10.1098/rstb.2002.1186.
13 External links
A collection of photosynthesis pages for all levels
from a renowned expert (Govindjee)

In depth, advanced treatment of photosynthesis, also

[81] Kortschak HP, Hart CE, Burr GO (1965). Carbon diox-
ide xation in sugarcane leaves. Plant Physiol. 40: 209
213. doi:10.1104/pp.40.2.209.
[82] Hatch MD, Slack CR (1966). Photosynthesis by sugar-
cane leaves. A new carboxylation reaction and the path-
way of sugar formation. Biochem. J. 101: 103111.
doi:10.1042/bj1010103.
[83] Jones HG (2014). Plants and Microclimate: a Quantitative
Approach to Environmental Plant Physiology (Third ed.).
Cambridge: Cambridge University Press. ISBN 978-0-
521-27959-8.
from Govindjee
Science Aid: Photosynthesis Article appropriate for
high school science
Metabolism, Cellular Respiration and Photosynthe-
sis The Virtual Library of Biochemistry and Cell
Biology
Overall examination of Photosynthesis at an inter-
mediate level
Overall Energetics of Photosynthesis

Photosynthesis Discovery Milestones experiments

and background
12 Further reading
The source of oxygen produced by photosynthesis
Interactive animation, a textbook tutorial
12.1 Books
Marshall J (2011-03-29). First practical articial
Bidlack JE, Stern KR, Jansky S (2003). Introductory leaf makes debut. Discovery News.
plant biology. New York: McGraw-Hill. ISBN 0-
Photosynthesis Light Dependent & Light Indepen-
07-290941-2.
dent Stages
Blankenship RE (2014). Molecular Mechanisms of Khan Academy, video introduction
Photosynthesis (2nd ed.). John Wiley & Sons. ISBN
978-1-4051-8975-0.

Govindjee, Beatty JT, Gest H, Allen JF (2006). Dis-

coveries in Photosynthesis. Advances in Photosyn-
thesis and Respiration. 20. Berlin: Springer. ISBN
1-4020-3323-0.

Reece JB, et al. (2013). Campbell Biology.

Benjamin Cummings. ISBN 978-0321775658.
 17

14 Text and image sources, contributors, and licenses

14.1 Text
Photosynthesis Source: https://en.wikipedia.org/wiki/Photosynthesis?oldid=800249060 Contributors: AxelBoldt, Magnus Manske, Marj
Tiefert, Mav, Bryan Derksen, Tarquin, Malcolm Farmer, -- April, Josh Grosse, Rgamble, William Avery, Chlor~enwiki, Ben-Zin~enwiki,
Heron, Jaknouse, Ram-Man, Spi~enwiki, Dwmyers, Michael Hardy, Modster, Jtdirl, Ixfd64, Cyde, Sannse, GTBacchus, Dori, Sebastian-
Helm, Looxix~enwiki, Ahoerstemeier, DavidWBrooks, Angela, JWSchmidt, Glenn, Ciphergoth, Tristanb, Evercat, Rob Hooft, Ghewgill,
Selket, Hao2lian, Tpbradbury, Tero~enwiki, Nv8200pa, Larryorr, Samsara, Traroth, Jecar, Elwoz, Fvw, AnonMoos, Jusjih, Shafei, An-
thonyQBachler, Jeq, Jni, Robbot, Fredrik, Modulatum, Postdlf, Academic Challenger, Jondel, Hadal, Fuelbottle, Cyrius, Pengo, Dina,
ManuelGR, Alan Liefting, Giftlite, Dbenbenn, Ryanrs, MPF, Mat-C, Kim Bruning, Lupin, Peruvianllama, Everyking, NeoJustin, Michael
Devore, Andris, Ans, Patrickdavidson, Sundar, Mboverload, Delta G, SonicAD, Adenosine, Gadum, Dullhunk, Antandrus, Onco p53,
OverlordQ, PDH, Bodnotbod, Zfr, Feniouk, Talrias, Mrrhum, Creidieki, Chao, Fermion, Sonett72, Kevyn, Demiurge, Adashiel, Aut1221,
Chepry, Canterbury Tail, Clubjuggle, Shotwell, Atrian, Perey, Freakofnurture, ClockworkTroll, DanielCD, RedWordSmith, Discospin-
ster, Rich Farmbrough, Cacycle, Vsmith, Wk muriithi, Xezbeth, Mani1, Pavel Vozenilek, Paul August, Bender235, ESkog, Kbh3rd,
Danny B-), Neko-chan, RJHall, Germen, *drew, Pinzo, Joanjoc~enwiki, Phoenix Hacker, Art LaPella, RoyBoy, Plociam, Guettarda,
Causa sui, Bobo192, Whosyourjudas, Shenme, Cmdrjameson, Vortexrealm, Cohesion, Dungodung, Arcadian, Cayte, SpeedyGonsales,
La goutte de pluie, Xoddam, Hi112233, Rje, Burnhamd, Brainy J, Haham hanuka, Iolar~enwiki, HasharBot~enwiki, Tra, Red Winged
Duck, Alansohn, Gary, JYolkowski, Etxrge, Nik42, Eric Kvaalen, Valrie75, JoaoRicardo, R Calvete, SlimVirgin, Mlessard, Hgrenbor,
Malo, Sir Joseph, PaePae, BanyanTree, ClockworkSoul, Dbrett480, Evil Monkey, Amorymeltzer, ComCat, Gene Nygaard, Bookand-
coee, Kazvorpal, Michael Ward, Ceyockey, Tchaika, Feezo, Nuno Tavares, Angr, OwenX, Woohookitty, Henrik, Cimex, TigerShark,
Etacar11, Camw, LOL, Consequencefree, Dandv, PoccilScript, TheArmadillo, Schlice, Mooinglemur, Username314, Fenteany, Bbatsell,
Optichan, Sengkang, Jugger90, Xiong Chiamiov, MassGalactusUniversum, Magister Mathematicae, BD2412, Li-sung, Tradnor, Chun-
hian, Lanoitarus, FreplySpang, Swmcd, Lord.lucan, Pentawing, Search4Lancer, Drbogdan, Sjakkalle, Rjwilmsi, Commander, Vary, Quid-
dity, Sdornan, Tawker, CQJ, The wub, Bhadani, Dar-Ape, GregAsche, Sango123, Molybdenumblue, Yamamoto Ichiro, Red Herring,
FlaBot, Ian Pitchford, Nivix, Elmer Clark, RexNL, Jrtayloriv, KFP, Miketam, Srleer, Daycd, Procrastinator~enwiki, Nsteinberg, King
of Hearts, Nicholasink, Chobot, DaGizza, Mhking, Korg, Dj Capricorn, WriterHound, Gwernol, Cassius987, YurikBot, Wavelength, Tex-
asAndroid, Hawaiian717, JDnCoke, Vuvar1, Sceptre, Jlittlet, Petiatil, Crazytales, Arado, Postglock, Acefox, Bergsten, SpuriousQ, Oke-
dem, Gaius Cornelius, Eleassar, Ihope127, Salsb, Wimt, GeeJo, Herbertxu, Jrbouldin, Shanel, Wiki alf, Jaxl, Johann Wolfgang, Dugosz,
Robchurch, Chakazul, Apokryltaros, Irishguy, Nick, Raven4x4x, Epipelagic, Pierpontpaul2351, FF2010, Lt-wiki-bot, Bayerischermann,
U.S.Vevek, Closedmouth, Studentne, KGasso, Fram, Katieh5584, Kungfuadam, The Catsh, DVD R W, Quadpus, David Wahler, Crys-
tallina, SmackBot, FocalPoint, MattieTK, Dweller, Malkinann, Tarret, Slashme, Prodego, TestPilot, Hydrogen Iodide, Melchoir, K-UNIT,
Unyoyega, Pgk, AndyZ, Jfg284, Davewild, Matthuxtable, Alksub, EncycloPetey, Hardyplants, Frymaster, RobotJcb, HalfShadow, Ga,
Yamaguchi , ITOD, Samballance, Gilliam, Ohnoitsjamie, Hmains, Skizzik, Rmosler2100, Anachronist, Master Jay, Persian Poet Gal,
Rmstock, Tito4000, Dreg743, Tree Biting Conspiracy, SchftyThree, Tetraglot, Funper, EScribe, Kungming2, Munyanah, Gruzd, Sbharris,
Polyhedron, Sunholm, Reaper X, Simpsons contributor, Veggies, Myrryam, Xchbla423, Lawrenceuniversity1, Can't sleep, clown will eat
me, OrphanBot, Init~enwiki, Rrburke, Kcordina, Wine Guy, Celarnor, SundarBot, Stevenmitchell, DinosaursLoveExistence, Iapetus, De-
cltype, Makemi, Bowlhover, Nakon, TedE, Nick125, Dreadstar, Smokefoot, Crittens, Drphilharmonic, Jklin, ATMarsden, Mion, InNuce,
Clicketyclack, SashatoBot, Nisadaninisa, Kuru, John, AmiDaniel, Soumyasch, JoshuaZ, Chodorkovskiy, JorisvS, Tim Q. Wells, Minna
Sora no Shita, Teh wise one, Mgiganteus1, Zarniwoot, Tikoo, Ocatecir, Eaglestrike117, Saseigel, Ckatz, White.matthew.09, Gwendy,
Smith609, Munita Prasad, Beetstra, Bendzh, Waggers, ASKingquestions, Funnybunny, Ryulong, Condem, Gregturn, Zepheus, Hu12,
CerealKiller~enwiki, Ginkgo100, Ramcy~enwiki, Vanished user, Nehrams2020, Iridescent, Pipedreambomb, Cheesy Yeast, Melnicki,
NativeForeigner, Mrg024, Hipporoo, Cbrown1023, SweetNeo85, Blehfu, Marysunshine, Color probe, Daharde, Courcelles, Tawkerbot2,
Daniel5127, Ouishoebean, Vices, Meltzerju, Rozzychan~enwiki, JForget, Toadams, Mattbr, Van helsing, Agathman, Shorespirit, Scohoust,
Woudloper, Riader0, CWY2190, Hometown, GHe, Pajast, TheTito, Z 153, Fox6453, Kupirijo, MC10, 678910, BGFMSM, Govindjee,
Pontifex101, Anonymi, B820, Chasingsol, Michael C Price, DumbBOT, Chrislk02, Robertinventor, Narayanese, NMChico24, HBNayr,
GeeOh, Thijs!bot, Epbr123, Dsinghs, Pajz, Kamal Chid, Kablammo, Angam~enwiki, Nonagonal Spider, Headbomb, John254, Kathovo,
Tellyaddict, Uiteoi, GregMinton, Dfrg.msc, Pkpat2011, Wikidenizen, Dawnseeker2000, Natalie Erin, Drdaveng, Escarbot, Darekun, David
D., AntiVandalBot, Kylemcinnes, Abu-Fool Danyal ibn Amir al-Makhiri, Luna Santin, Prince Godfather, EarthPerson, Quintote, Pro
crast in a tor, TimVickers, Peipei, Smartse, Darklilac, Malcolm, Zedla, Imogen89, Storkk, Ed, Incredio, Trappist, Deective, MER-C,
Kaabi, Sanchom, Hello32020, Db099221, Mwarren us, Johna100, Bencherlite, Robsavoie, Magioladitis, Gekedo, Connormah, Bongwar-
rior, VoABot II, VolcomRUXC, Wikidudeman, Aeioun, SparrowsWing, Avicennasis, Qwerqwerqwer, Indon, 28421u2232nfenfcenc, User
A1, Laur2ro, DerHexer, Stephenchou0722, Hdt83, MartinBot, Unclepea, Arjun01, MNAdam, Anaxial, Sarin123, CommonsDelinker,
Leyo, Joshjoshjosh1, Jamib0y, Pekaje, WelshMatt, Smokizzy, J.delanoy, Pkoden, Blahed768, Ali, Bogey97, Nbauman, Name b, Hans
Dunkelberg, Boghog, Usdi, Uncle Dick, Interplanet Janet, Icseaturtles, NinaStarry, McSly, RTBoyce, Mikael Hggstrm, L'Aquatique,
(jarbarf), Exdejesus, NewEnglandYankee, Domminico, Fullmetal123321, Rhysboy, Dienamight, Dacrycarpus, DorganBot, Treisijs, King
Toadsworth, Bonadea, JamesR777, Goatman12, Ali salter, Darklich14, P a k 2, AJRM, Cdorman2, Iron Dragon91, Bertiethecat, Cardi-
nalDan, RJASE1, Idioma-bot, Nonluddite, Lights, Deor, VolkovBot, Clyde10race, OMGHeHasAGun, Indubitably, AlnoktaBOT, Philip
Trueman, TXiKiBoT, Rambatino, Kww, Tameeria, Malljaja, Norrello, SueHay, Hpfreak26, Qxz, Someguy1221, HLHJ, Steven J. An-
derson, Martin451, JhsBot, Juhwade, Raymondwinn, Ilyushka88, Duncan.Hull, Maxim, Coching, Synthebot, @pple, Turgan, Likeabird,
Bateman1234, Thanatos666, PoeticX, WatermelonPotion, Lily15, Delgolo, AlleborgoBot, Imi2012, Logan, Pcruce, Rock2e, EmxBot,
Neyne, Bailey654, Lord 1284, SieBot, Jwray, Primal400, Scarian, ToePeu.bot, Caltas, Randomnotice, Hunrizzo8, Keilana, Yon Yon-
son, Collegedegree, Momo san, Samtux~enwiki, Oiws, Yerpo, Mimihitam, Oxymoron83, Jasonphollinger, Steven Crossin, Lightmouse,
Towerofthunder, Dr. Mandellez, Sunrise, Maelgwnbot, Viper777~enwiki, Spartan-James, Kopid03, Thatotherdude, Mygerardromance,
WikiLaurent, Pinkadelica, Hordaland, Troy 07, WikipedianMarlith, Touchstone42, Bfahome, Faithlessthewonderboy, ClueBot, Snigbrook,
Fox, Stoic Squirrel, The Thing That Should Not Be, Jan1nad, Yikrazuul, Nsk92, Shark96z, Wutsje, Skpperd, Boing! said Zebedee,
Niceguyedc, Malomaboy06, Dasyornis, Proguitarboy, DragonBot, Excirial, Eeekster, Estirabot, Silasmellor, Mdavies 965, Dekisugi, Muro
Bot, Ottawa4ever, ChrisHodgesUK, Thingg, Chirpy7, Aitias, SoxBot III, Tdslk, DumZiBoT, RMFan1, Jamesscottbrown, Eik Corell, Nnew-
ton, Jovianeye, WikHead, BaroqueFox, Alexius08, Jbeans, Torahjerus14, ZooFari, Gggh, Lemchesvej, Surtsicna, CalumH93, Gacha1,
Addbot, CurtisSwain, DOI bot, Guoguo12, Oliverc1991, Theleftorium, Upeksha112, C3r4, Nuvitauy07, Tattoo275, Ronhjones, Cana-
dianLinuxUser, Jpoelma13, Download, Morning277, Chamal N, EconoPhysicist, Demi s96, Debresser, Doniago, LinkFA-Bot, 5 albert
square, Imanoob69, Brainmachine, Numbo3-bot, Bluedeed, Tide rolls, Ettrig, Math Champion, Luckas-bot, Yobot, TaBOT-zerem, II Mus-
LiM HyBRiD II, The Earwig, THEN WHO WAS PHONE?, , Szajci, YouAreNotReadingThis, AnomieBOT, 1exec1, Accuruss,
18 14 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

JackieBot, Oggie boggie in the loo, 9258fahskh917fas, Piano non troppo, Fahadsadah, Kingpin13, Nibls, Ulric1313, EryZ, Materialsci-
entist, Kasper90, Citation bot, ArthurBot, Xqbot, The sock that should not be, Engineering Guy, Gigemag76, Corrigen, Tomaschwutz,
Call me Bubba, Twirligig, EqualMusic, Tackyleprechaun, Antony2, Dinsdalea, Me big shy, Jazzy211, Eumeng.chong, Prunesqualer,
Oxnora2012, Mrmonsterman, Ryantoh wiki, Shadowjams, Racknack7824, Samwb123, GliderMaven, FrescoBot, DavidDidwin, Lucien-
BOT, Paine Ellsworth, Tobby72, Wikipe-tan, Oldlaptop321, Staccatoque, D'ohBot, Dirty MF Harry, Crazychinchilla, PJsg1011, Fgbhrgth-
nrtgnb, Citation bot 1, Citation bot 4, Shiningstarr1997, Daniel233, Intelligentsium, Pinethicket, P00100p, HRoestBot, Edderso, Watplay,
Abductive, Jonesey95, Sdittami, RedBot, DotComCairney, Thinking of England, SpaceFlight89, NarSakSasLee, Barras, Wejer, SW3 5DL,
Jauhienij, Mj455972007, TobeBot, Trappist the monk, Sweet xx, NYMFan69-86, Lotje, NMFA Northern Eagle, Vrenator, Zvn, LilyKitty,
Jackgeddes1234, Eddturtle, Amkilpatrick, Weedwhacker128, Amercer09, Tbhotch, Stroppolo, Reach Out to the Truth, Minimac, Sd card,
Tastycheeze, Mikimickz, DARTH SIDIOUS 2, RjwilmsiBot, Bento00, DrMtl, Gould363, Billare, Grondemar, DASHBot, EmausBot, John
of Reading, O le gend, Ohyeahbabe, Klbrain, Tommy2010, Platoface, Dcirovic, Rachelpurdon, Sergey WereWolf, Starzjake, Stevead-
cuk, Nickresch, Pvgetsome, Wiitennisxd, Qniemiec, Zap Rowsdower, Jay-Sebastos, L Kensington, ChuispastonBot, Cman12345657578,
Saif11, Iam angryrawr, Staticd, Mrwarriorman, LikeLakers2, JonRicheld, ClueBot NG, Cwmhiraeth, Gilderien, Zxoxm, Devil42400,
ScottSteiner, Helpful Pixie Bot, Calabe1992, Bibcode Bot, BG19bot, CatPath, AvocatoBot, Ghart717, Davidiad, Weblars, Neuraxs,
Tom Pippens, Exercisephys, Aranea Mortem, Glevum, Robert Thyder, Derekrhu, Jacob635, Aidentbrown, Zedshort, At09kg, Minsbot,
Rhinopias, Kc kennylau, Jack No1, ChrisGualtieri, Tn4196, JYBot, BrightStarSky, Dexbot, Mogism, Me, Myself, and I are Here, Ab-
otan, Kylcon2, FallingGravity, Zorahia, FamAD123, JPaestpreornJeolhlna, NHCLS, Eunkyoon, Csutric, Mahusha, PaulGWiki, Monkbot,
Somepics, D. Cordoba-Bahle, Trackteur, Manaiak31, Plant Stress Doctor, Nmcke1, Agc990, Felipe Jo, IMcBeats, VeenM64, KasparBot,
Baljeet Bilaspur, SeemaaSaleh, InternetArchiveBot, KATMAKROFAN, KolbertBot, Kalli navya and Anonymous: 1376

14.2 Images
File:Aegopodium_podagraria1_ies.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/bf/Aegopodium_podagraria1_ies.
jpg License: CC-BY-SA-3.0 Contributors: Own work Original artist: Frank Vincentz
File:Calvin-cycle4.svg Source: https://upload.wikimedia.org/wikipedia/commons/8/8e/Calvin-cycle4.svg License: CC BY-SA 3.0 Con-
tributors: Own work Original artist: Mike Jones
File:Chlorophyll_ab_spectra-en.svg Source: https://upload.wikimedia.org/wikipedia/commons/2/23/Chlorophyll_ab_spectra-en.svg
License: CC BY-SA 3.0 Contributors: This le was derived from Chlorophyll ab spectra2.PNG: <a href='//commons.wikimedia.org/wiki/
File:Chlorophyll_ab_spectra2.PNG' class='image'><img alt='Chlorophyll ab spectra2.PNG' src='https://upload.wikimedia.org/wikipedia/
commons/thumb/6/68/Chlorophyll_ab_spectra2.PNG/50px-Chlorophyll_ab_spectra2.PNG' width='50' height='47' srcset='https:
//upload.wikimedia.org/wikipedia/commons/thumb/6/68/Chlorophyll_ab_spectra2.PNG/75px-Chlorophyll_ab_spectra2.PNG 1.5x,
https://upload.wikimedia.org/wikipedia/commons/thumb/6/68/Chlorophyll_ab_spectra2.PNG/100px-Chlorophyll_ab_spectra2.PNG 2x'
data-le-width='664' data-le-height='629' /></a>
Original artist: Chlorophyll_ab_spectra2.PNG: Daniele Pugliesi
File:Chloroplast.svg Source: https://upload.wikimedia.org/wikipedia/commons/2/26/Chloroplast.svg License: CC BY-SA 3.0 Contribu-
tors: own work based on Chloroplaste-schema.gif Original artist: SuperManu
File:Commons-logo.svg Source: https://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: PD Contributors: ? Origi-
nal artist: ?
File:Earth_Day_Flag.png Source: https://upload.wikimedia.org/wikipedia/commons/6/6a/Earth_Day_Flag.png License: Public domain
Contributors: File:Earth ag PD.jpg, File:The Earth seen from Apollo 17 with transparent background.png Original artist: NASA (Earth
photograph)
SiBr4 (ag image)
File:Folder_Hexagonal_Icon.svg Source: https://upload.wikimedia.org/wikipedia/en/4/48/Folder_Hexagonal_Icon.svg License: Cc-by-
sa-3.0 Contributors: ? Original artist: ?
File:HatchSlackpathway2.svg Source: https://upload.wikimedia.org/wikipedia/commons/4/47/HatchSlackpathway2.svg License: CC
BY-SA 2.5 Contributors:
HatchSlackpathway.svg Original artist: HatchSlackpathway.svg: *HatchSlackpathway.png: Adenosine
File:Interactive_icon.svg Source: https://upload.wikimedia.org/wikipedia/commons/e/e6/Interactive_icon.svg License: CC BY 4.0 Con-
tributors: Own work Original artist: Thomas Shafee
File:Leaf_1_web.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/f4/Leaf_1_web.jpg License: Public domain Contribu-
tors: PdPhoto Original artist: Jon Sullivan
File:Lock-green.svg Source: https://upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg License: CC0 Contributors: en:File:
Free-to-read_lock_75.svg Original artist: User:Trappist the monk
File:Melvin_Calvin.jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/04/Melvin_Calvin.jpg License: Public domain
Contributors: LBL Collection Original artist: Photolab
File:Metabolic_Metro_Map_(no_legends).svg Source: https://upload.wikimedia.org/wikipedia/commons/4/41/Metabolic_Metro_
Map_%28no_legends%29.svg License: CC BY-SA 4.0 Contributors: Own work Original artist: Chakazul
File:Metabolic_metro_blue.svg Source: https://upload.wikimedia.org/wikipedia/commons/1/1b/Metabolic_metro_blue.svg License:
CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_brown.svg Source: https://upload.wikimedia.org/wikipedia/commons/9/99/Metabolic_metro_brown.svg Li-
cense: CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_green.svg Source: https://upload.wikimedia.org/wikipedia/commons/9/97/Metabolic_metro_green.svg License:
CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_grey.svg Source: https://upload.wikimedia.org/wikipedia/commons/c/c8/Metabolic_metro_grey.svg License:
CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
14.3 Content license 19

File:Metabolic_metro_orange.svg Source: https://upload.wikimedia.org/wikipedia/commons/a/a0/Metabolic_metro_orange.svg Li-

cense: CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_pink.svg Source: https://upload.wikimedia.org/wikipedia/commons/d/db/Metabolic_metro_pink.svg License:
CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_purple.svg Source: https://upload.wikimedia.org/wikipedia/commons/e/e7/Metabolic_metro_purple.svg Li-
cense: CC BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:Metabolic_metro_red.svg Source: https://upload.wikimedia.org/wikipedia/commons/b/ba/Metabolic_metro_red.svg License: CC
BY-SA 4.0 Contributors: Own work Original artist: Thomas Shafee
File:One_dimensional_quantum_random_walk.png Source: https://upload.wikimedia.org/wikipedia/commons/4/4b/One_
dimensional_quantum_random_walk.png License: CC BY-SA 3.0 Contributors: Own work Original artist: shoyer
File:People_icon.svg Source: https://upload.wikimedia.org/wikipedia/commons/3/37/People_icon.svg License: CC0 Contributors: Open-
Clipart Original artist: OpenClipart
File:Photorespiration.svg Source: https://upload.wikimedia.org/wikipedia/commons/f/f8/Photorespiration.svg License: CC BY-SA 3.0
Contributors: Own work Original artist: Yikrazuul
File:Photosynthesis.gif Source: https://upload.wikimedia.org/wikipedia/commons/d/db/Photosynthesis.gif License: CC BY-SA 3.0 Con-
tributors: Own work Original artist: At09kg
File:Photosynthesis_equation.svg Source: https://upload.wikimedia.org/wikipedia/commons/f/fd/Photosynthesis_equation.svg License:
Public domain Contributors: Own work Original artist: ZooFari
File:Plagiomnium_affine_laminazellen.jpeg Source: https://upload.wikimedia.org/wikipedia/commons/4/49/Plagiomnium_affine_
laminazellen.jpeg License: CC-BY-SA-3.0 Contributors: Self-photographed Original artist: Kristian Peters -- Fabelfroh
File:Portal-puzzle.svg Source: https://upload.wikimedia.org/wikipedia/en/f/fd/Portal-puzzle.svg License: Public domain Contributors: ?
Original artist: ?
File:Seawifs_global_biosphere.jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/44/Seawifs_global_biosphere.jpg Li-
cense: Public domain Contributors: http://oceancolor.gsfc.nasa.gov/SeaWiFS/BACKGROUND/Gallery/index.html and from en:Image:
Seawifs global biosphere.jpg Original artist: Provided by the SeaWiFS Project, Goddard Space Flight Center and ORBIMAGE
File:Simple_photosynthesis_overview.svg Source: https://upload.wikimedia.org/wikipedia/commons/0/0c/Simple_photosynthesis_
overview.svg License: GFDL Contributors: Own work Original artist: Daniel Mayer (mav) - original image

File:The_Earth_seen_from_Apollo_17_with_transparent_background.png Source: https://upload.wikimedia.org/wikipedia/

commons/4/43/The_Earth_seen_from_Apollo_17_with_transparent_background.png License: Public domain Contributors:
http://nssdc.gsfc.nasa.gov/imgcat/html/object_page/a17_h_148_22727.html Original artist: NASA
File:Thylakoid_membrane_3.svg Source: https://upload.wikimedia.org/wikipedia/commons/4/49/Thylakoid_membrane_3.svg License:
CC BY-SA 4.0 Contributors: Own work Original artist: Somepics
File:Z-scheme.png Source: https://upload.wikimedia.org/wikipedia/commons/c/c7/Z-scheme.png License: CC-BY-SA-3.0 Contributors:
http://en.wikipedia.org/wiki/Image:Z-scheme.png Original artist: w:User:Bensaccount

14.3 Content license

Creative Commons Attribution-Share Alike 3.0