Chlorophyll

Chlorophyll (also chlorophyl) is any of

several related green pigments found in
cyanobacteria and in the chloroplasts of
algae and plants.[2] Its name is derived
from the Greek words χλωρός, khloros
("pale green") and φύλλον, phyllon
("leaf").[3] Chlorophyll allow plants to
absorb energy from light.

Chlorophyll at different scales

Chlorophyll is responsible for the green color of
many plants and algae.

Seen through a microscope, chlorophyll is

concentrated within organisms in structures called
chloroplasts – shown here grouped inside plant
cells.

Plants are perceived as green because chlorophyll

absorbs mainly the blue and red wavelengths but
green light, reflected by plant structures like cell
walls, is less absorbed.[1]
 There are several types of chlorophyll, but all share
the chlorin magnesium ligand which forms the right
side of this diagram.

Chlorophylls absorb light most strongly in

the blue portion of the electromagnetic
spectrum as well as the red portion.[4]
Conversely, it is a poor absorber of green
and near-green portions of the spectrum.
Hence chlorophyll-containing tissues
appear green because green light,
diffusively reflected by structures like cell
walls, is less absorbed.[1] Two types of
chlorophyll exist in the photosystems of
green plants: chlorophyll a and b.[5]

History

Chlorophyll was first isolated and named

by Joseph Bienaimé Caventou and Pierre
Joseph Pelletier in 1817.[6] The presence
of magnesium in chlorophyll was
discovered in 1906,[7] and was the first
detection of that element in living tissue.[8]

After initial work done by German chemist

Richard Willstätter spanning from 1905 to
1915, the general structure of chlorophyll a
was elucidated by Hans Fischer in 1940.
By 1960, when most of the
stereochemistry of chlorophyll a was
known, Robert Burns Woodward published
a total synthesis of the molecule.[8][9] In
1967, the last remaining stereochemical
elucidation was completed by Ian
Fleming,[10] and in 1990 Woodward and co-
authors published an updated
synthesis.[11] Chlorophyll f was announced
to be present in cyanobacteria and other
oxygenic microorganisms that form
stromatolites in 2010;[12][13] a molecular
formula of C55H70O6N4Mg and a structure
of (2-formyl)-chlorophyll a were deduced
based on NMR, optical and mass
spectra.[14]
Photosynthesis

Absorbance spectra of free

chlorophyll a (blue) and b (red) in a
solvent. The spectra of chlorophyll
molecules are slightly modified in vivo
depending on specific pigment-
protein interactions.
Chlorophyll a
Chlorophyll b

Chlorophyll is vital for photosynthesis,

which allows plants to absorb energy from
light.[15]

Chlorophyll molecules are arranged in and

around photosystems that are embedded
in the thylakoid membranes of
chloroplasts.[16] In these complexes,
chlorophyll serves three functions:

1. The function of the vast majority of

chlorophyll (up to several hundred
molecules per photosystem) is to
absorb light.
2. Having done so, these same centers
execute their second function: The
transfer of that energy by resonance
energy transfer to a specific
chlorophyll pair in the reaction center
of the photosystems.
3. This specific pair performs the final
function of chlorophylls: Charge
separation, which produces the
 unbound protons (H+) and electrons
(e−) that separately propel
biosynthesis.

The two currently accepted photosystem

units are photosystem I and
photosystem II, which have their own
distinct reaction centres, named P700 and
P680, respectively. These centres are
named after the wavelength (in
nanometers) of their red-peak absorption
maximum. The identity, function and
spectral properties of the types of
chlorophyll in each photosystem are
distinct and determined by each other and
the protein structure surrounding them.
The function of the reaction center of
chlorophyll is to absorb light energy and
transfer it to other parts of the
photosystem. The absorbed energy of the
photon is transferred to an electron in a
process called charge separation. The
removal of the electron from the
chlorophyll is an oxidation reaction. The
chlorophyll donates the high energy
electron to a series of molecular
intermediates called an electron transport
chain. The charged reaction center of
chlorophyll (P680+) is then reduced back
to its ground state by accepting an
electron stripped from water. The electron
that reduces P680+ ultimately comes from
the oxidation of water into O2 and H+
through several intermediates. This
reaction is how photosynthetic organisms
such as plants produce O2 gas, and is the
source for practically all the O2 in Earth's
atmosphere. Photosystem I typically
works in series with Photosystem II; thus
the P700+ of Photosystem I is usually
reduced as it accepts the electron, via
many intermediates in the thylakoid
membrane, by electrons coming,
ultimately, from Photosystem II. Electron
transfer reactions in the thylakoid
membranes are complex, however, and the
source of electrons used to reduce P700+
can vary.
The electron flow produced by the reaction
center chlorophyll pigments is used to
pump H+ ions across the thylakoid
membrane, setting up a proton-motive
force a chemiosmotic potential used
mainly in the production of ATP (stored
chemical energy) or to reduce NADP+ to
NADPH. NADPH is a universal agent used
to reduce CO2 into sugars as well as other
biosynthetic reactions.

Reaction center chlorophyll–protein

complexes are capable of directly
absorbing light and performing charge
separation events without the assistance
of other chlorophyll pigments, but the
probability of that happening under a given
light intensity is small. Thus, the other
chlorophylls in the photosystem and
antenna pigment proteins all cooperatively
absorb and funnel light energy to the
reaction center. Besides chlorophyll a,
there are other pigments, called accessory
pigments, which occur in these pigment–
protein antenna complexes.

Chemical structure

Space-filling model of the

chlorophyll a molecule
Several chlorophylls are known. All are
defined as derivatives of the parent chlorin
by the presence of a fifth, ketone-
containing ring beyond the four pyrrole-like
rings. Most chlorophylls are classified as
chlorins, which are reduced relatives of
porphyrins (found in hemoglobin). They
share a common biosynthetic pathway
with porphyrins, including the precursor
uroporphyrinogen III. Unlike hemes, which
contain iron bound to the N4 center, most
chlorophylls bind magnesium. The axial
ligands attached to the Mg2+ center are
often omitted for clarity. Appended to the
chlorin ring are various side chains, usually
including a long phytyl chain (C20H39O).
The most widely distributed form in
terrestrial plants is chlorophyll a. The only
difference between chlorophyll a and
chlorophyll b is that the former has a
methyl group where the latter has a formyl
group. This difference causes a
considerable difference in the absorption
spectrum, allowing plants to absorb a
greater portion of visible light.

The structures of chlorophylls are

summarized below:[17][18]
 Chlorophyll a Chlorophyll b Chlorophyll c1 Chlorophyll c2 Ch

Molecular
C55H72O5N4Mg C55H70O6N4Mg C35H30O5N4Mg C35H28O5N4Mg C54H70
formula

C2 group −CH3 −CH3 −CH3 −CH3 −CH3

C3 group −CH=CH2 −CH=CH2 −CH=CH2 −CH=CH2 −CHO

C7 group −CH3 −CHO −CH3 −CH3 −CH3

C8 group −CH2CH3 −CH2CH3 −CH2CH3 −CH=CH2 −CH2C

C17 group −CH2CH2COO−Phytyl −CH2CH2COO−Phytyl −CH=CHCOOH −CH=CHCOOH −CH2C

C17−C18 Single Single Double Double Single

bond (chlorin) (chlorin) (porphyrin) (porphyrin) (chlori

Occurrence Universal Mostly plants Various algae Various algae Cyano

Structures of chlorophylls
chlorophyll a

chlorophyll b
chlorophyll c1

chlorophyll c2
chlorophyll d

chlorophyll f

Chlorophyll e is reserved for a pigment

that has been extracted from algae in
1966 but not chemically described.
Besides the lettered chlophylls, a wide
variety of sidechain modifications to the
chlorophyll structures are known in the
wild. For example, Prochlorococcus, a
cyanobacterium, uses 8-vinyl Chl a and
b.[19]

Measurement of chlorophyll
content

Chlorophyll forms deep green

solutions in organic solvents.
Chlorophylls can be extracted from the
protein into organic solvents.[20][21][22] In
this way, the concentration of chlorophyll
within a leaf can be estimated.[23] Methods
also exist to separate chlorophyll a and
chlorophyll b.

In diethyl ether, chlorophyll a has

approximate absorbance maxima of
430 nm and 662 nm, while chlorophyll b
has approximate maxima of 453 nm and
642 nm.[24] The absorption peaks of
chlorophyll a are at 465 nm and 665 nm.
Chlorophyll a fluoresces at 673 nm
(maximum) and 726 nm. The peak molar
absorption coefficient of chlorophyll a
exceeds 105 M−1 cm−1, which is among
the highest for small-molecule organic
compounds.[25] In 90% acetone-water, the
peak absorption wavelengths of
chlorophyll a are 430 nm and 664 nm;
peaks for chlorophyll b are 460 nm and
647 nm; peaks for chlorophyll c1 are
442 nm and 630 nm; peaks for chlorophyll
c2 are 444 nm and 630 nm; peaks for
chlorophyll d are 401 nm, 455 nm and
696 nm.[26]

Ratio fluorescence emission can be used

to measure chlorophyll content. By
exciting chlorophyll a fluorescence at a
lower wavelength, the ratio of chlorophyll
fluorescence emission at 705 ± 10 nm and
735 ± 10 nm can provide a linear
relationship of chlorophyll content when
compared with chemical testing. The ratio
F735/F700 provided a correlation value of r2
0.96 compared with chemical testing in
the range from 41 mg m−2 up to
675 mg m−2. Gitelson also developed a
formula for direct readout of chlorophyll
content in mg m−2. The formula provided a
reliable method of measuring chlorophyll
content from 41 mg m−2 up to 675 mg m−2
with a correlation r2 value of 0.95.[27]
Biosynthesis

In some plants, chlorophyll is derived from

glutamate and is synthesised along a
branched biosynthetic pathway that is
shared with heme and
siroheme.[28][29][30]Chlorophyll synthase[31]
is the enzyme that completes the
biosynthesis of chlorophyll a:[32][33]

chlorophyllide a + phytyl diphosphate

chlorophyll a + diphosphate

This converion forms an ester of the

carboxylic acid group in chlorophyllide a
with the 20-carbon diterpene alcohol
phytol. Chlorophyll b is made by the same
enzyme acting on chlorophyllide b. The
same is known for chlorophyll d and f,
both made from corresponding
chlorophyllides ultimately made from
chlorophyllide a.[34]

In Angiosperm plants, the later steps in the

biosynthetic pathway are light-dependent.
Such plants are pale (etiolated) if grown in
darkness. Non-vascular plants and green
algae have an additional light-independent
enzyme and grow green even in
darkness.[35]
Chlorophyll is bound to proteins.
Protochlorophyllide, one of the
biosynthetic intermediates, occurs mostly
in the free form and, under light conditions,
acts as a photosensitizer, forming free
radicals, which can be toxic to the plant.
Hence, plants regulate the amount of this
chlorophyll precursor. In angiosperms, this
regulation is achieved at the step of
aminolevulinic acid (ALA), one of the
intermediate compounds in the
biosynthesis pathway. Plants that are fed
by ALA accumulate high and toxic levels of
protochlorophyllide; so do the mutants
with a damaged regulatory system.[36]
Senescence and the
chlorophyll cycle

The process of plant senescence involves

the degradation of chlorophyll: for
example the enzyme chlorophyllase (EC
3.1.1.14 (https://enzyme.expasy.org/EC/3.
1.1.14) ) hydrolyses the phytyl sidechain
to reverse the reaction in which
chlorophylls are biosynthesised from
chlorophyllide a or b. Since chlorophyllide
a can be converted to chlorophyllide b and
the latter can be re-esterified to chlorophyll
b, these processes allow cycling between
chlorophylls a and b. Moreover, chlorophyll
b can be directly reduced (via
71-hydroxychlorophyll a) back to
chlorophyll a, completing the cycle.[37][38]
In later stages of senescence,
chlorophyllides are converted to a group of
colourless tetrapyrroles known as
nonfluorescent chlorophyll catabolites
(NCC's) with the general structure:

These compounds have also been

identified in ripening fruits and they give
characteristic autumn colours to
deciduous plants.[38][39]

Distribution

The chlorophyll maps show milligrams of

chlorophyll per cubic meter of seawater
each month. Places where chlorophyll
amounts were very low, indicating very low
numbers of phytoplankton, are blue.
Places where chlorophyll concentrations
were high, meaning many phytoplankton
were growing, are yellow. The observations
come from the Moderate Resolution
Imaging Spectroradiometer (MODIS) on
NASA's Aqua satellite. Land is dark gray,
and places where MODIS could not collect
data because of sea ice, polar darkness, or
clouds are light gray. The highest
chlorophyll concentrations, where tiny
surface-dwelling ocean plants are thriving,
are in cold polar waters or in places where
ocean currents bring cold water to the
surface, such as around the equator and
along the shores of continents. It is not the
cold water itself that stimulates the
phytoplankton. Instead, the cool
temperatures are often a sign that the
water has welled up to the surface from
deeper in the ocean, carrying nutrients that
have built up over time. In polar waters,
nutrients accumulate in surface waters
during the dark winter months when plants
cannot grow. When sunlight returns in the
spring and summer, the plants flourish in
high concentrations.[40]

Culinary use

Synthetic chlorophyll is registered as a

food additive colorant, and its E number is
E140. Chefs use chlorophyll to color a
variety of foods and beverages green, such
as pasta and spirits. Absinthe gains its
green color naturally from the chlorophyll
introduced through the large variety of
herbs used in its production.[41]
Chlorophyll is not soluble in water, and it is
first mixed with a small quantity of
vegetable oil to obtain the desired
solution.

Biological use

A 2002 study found that "leaves exposed

to strong light contained degraded major
antenna proteins, unlike those kept in the
dark, which is consistent with studies on
the illumination of isolated proteins". This
appeared to the authors as support for the
hypothesis that "active oxygen species
play a role in vivo" in the short-term
behaviour of plants.[42]
See also

Wikimedia Commons has media related

to Chlorophyll.
Bacteriochlorophyll, related compounds
in phototrophic bacteria
Chlorophyllin, a semi-synthetic
derivative of chlorophyll
Deep chlorophyll maximum
Chlorophyll fluorescence, to measure
plant stress

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