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biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...
, a light-harvesting complex or LHC is an aggregate consisting of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s bound with chromophores (
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
s and
carotenoid Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
s) that play a key role in
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
. LHCs are arrayed around photosynthetic reaction centers in both
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s and photosynthetic bacteria and collect more of the incoming light than would be captured by the reaction centers alone. The light captured by the chromophores excites molecules from their ground states to (short-lived) higher-energy states, known as the excited states. This energy is then focused toward the reaction centers by
Förster resonance energy transfer Förster resonance energy transfer (FRET), fluorescence resonance energy transfer, resonance energy transfer (RET) or electronic energy transfer (EET) is a mechanism describing energy transfer between two light-sensitive molecules (chromophores). ...
. Light-harvesting complexes are found in a wide variety among the different photosynthetic species, with no homology among the major groups.


Function

Photosynthesis is a process where light is absorbed or harvested by pigment protein complexes which are able to turn sunlight into chemical energy. In this process, a molecule of the pigment protein absorbs a
photon A photon () is an elementary particle that is a quantum of the electromagnetic field, including electromagnetic radiation such as light and radio waves, and the force carrier for the electromagnetic force. Photons are massless particles that can ...
of sunlight, leading to electronic excitation delivered to the reaction centre where the process of charge separation can take place if the energy of the absorbed photon matches that of an electronic transition. The result of such excitation can be a return to the ground state or to another electronic state of the same molecule. When the excited molecule has a nearby neighbour molecule, the excitation energy may also be transferred, through electromagnetic interactions, from one molecule to another. This process is called resonance energy transfer, and the rate depends strongly on the distance between the energy donor and energy acceptor molecules. Before an excited molecule can transition back to its ground state, energy needs to be harvested. This excitation is transferred among chromophores where it is delivered to the reaction centre. Light-harvesting complexes have their pigments specifically positioned to optimize these rates.


In purple bacteria

Purple bacteria are a type of photosynthetic organism with a light harvesting complex consisting of two pigment protein complexes, referred to as LH1 and LH2. Within the photosynthetic membrane, these two complexes differ in their arrangement. The LH1 complexes surround the reaction centre, while the LH2 complexes are arranged peripherally around the LH1 complexes and the reaction centre. Purple bacteria use bacteriochlorophyll and carotenoids to gather light energy. These proteins are arranged in a ring-like fashion, creating a cylinder that spans the membrane.


In green bacteria

The main light harvesting complex in Green bacteria is known as the chlorosome. The chlorosome is equipped with rod-like BChl c aggregates with protein embedded lipids surrounding it. Chlorosomes are found outside of the membrane which covers the reaction centre. Green sulphur bacteria and some Chloroflexia use ellipsoidal complexes known as the chlorosome to capture light. Their form of bacteriochlorophyll is green.


In cyanobacteria and plants

Chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
s and
carotenoids Carotenoids () are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and Fungus, fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips ...
are important in light-harvesting complexes present in plants.
Chlorophyll b } Chlorophyll ''b'' is a form of chlorophyll. Chlorophyll ''b'' helps in photosynthesis by absorbing light energy. It is more soluble than chlorophyll ''a'' in polar solvents because of its carbonyl group. Its color is green, and it primarily ...
is almost identical to chlorophyll a, except it has a formyl group in place of a
methyl group In organic chemistry, a methyl group is an alkyl derived from methane, containing one carbon atom bonded to three hydrogen atoms, having chemical formula (whereas normal methane has the formula ). In formulas, the group is often abbreviated a ...
. This small difference makes chlorophyll b absorb
light Light, visible light, or visible radiation is electromagnetic radiation that can be visual perception, perceived by the human eye. Visible light spans the visible spectrum and is usually defined as having wavelengths in the range of 400– ...
with
wavelengths In physics and mathematics, wavelength or spatial period of a wave or periodic function is the distance over which the wave's shape repeats. In other words, it is the distance between consecutive corresponding points of the same '' phase'' on ...
between 400 and 500 nm more efficiently. Carotenoids are long linear organic molecules that have alternating single and double bonds along their length. Such molecules are called
polyenes In organic chemistry, polyenes are polyunsaturated organic compounds that contain multiple carbon–carbon double bonds (). Some sources consider dienes to be polyenes, whereas others require polyenes to contain at least three carbon–carbon d ...
. Two examples of carotenoids are
lycopene Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene (from the Neo-Latin '' Lycopersicon'', the name of a former tomato genus) is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and ve ...
and β-carotene. These molecules also absorb light most efficiently in the 400 – 500 nm range. Due to their absorption region, carotenoids appear red and yellow and provide most of the red and yellow colours present in
fruits In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
and
flowers Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
. The carotenoid molecules also serve a safeguarding function. Carotenoid molecules suppress damaging photochemical reactions, in particular those including
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
, which exposure to sunlight can cause. Plants that lack carotenoid molecules quickly die upon exposure to oxygen and light.


Phycobilisome

The antenna-shaped light harvesting complex of
cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
, glaucocystophyta, and
red algae Red algae, or Rhodophyta (, ; ), make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest Phylum, phyla of algae, containing over 7,000 recognized species within over 900 Genus, genera amidst ongoing taxon ...
is known as the phycobilisome; it is composed of linear tetrapyrrole pigments. Pigment-protein complexes, referred to as R-phycoerythrin, are rod-like in shape and make up the rods and core of the phycobilisome. Little light reaches algae that reside at a depth of one meter or more in seawater, as light is absorbed by seawater. The pigments, such as phycocyanobilin and
phycoerythrobilin Phycoerythrobilin is a red phycobilin, i.e. an open tetrapyrrole chromophore found in cyanobacteria and in the chloroplasts of red algae, glaucophytes and some cryptomonads. Phycoerythrobilin is present in the phycobiliprotein phycoerythrin, of w ...
, are the chromophores that bind through a covalent
thioether In organic chemistry, a sulfide (British English sulphide) or thioether is an organosulfur functional group with the connectivity as shown on right. Like many other sulfur-containing compounds, Volatile organic compound, volatile sulfides have ...
bond to their apoproteins at cystein residues. The apoprotein with its chromophore is called phycocyanin, phycoerythrin, and allophycocyanin, respectively. They often occur as
hexamer In chemistry and biochemistry, an oligomer () is a molecule that consists of a few repeating units which could be derived, actually or conceptually, from smaller molecules, monomer, monomers.Quote: ''Oligomer molecule: A molecule of intermediate ...
s of α and β subunits (α3β3)2. They enhance the amount and spectral window of light absorption and fill the "green gap", which occurs in higher plants. The geometrical arrangement of a phycobilisome is very elegant and results in 95% efficiency of energy transfer. There is a central core of
allophycocyanin Allophycocyanin ("other algal blue protein"; from Greek language, Greek: '' (allos)'' meaning "other", '' (phykos)'' meaning “alga”, and '' (kyanos)'' meaning "blue") is a protein from the light-harvesting phycobiliprotein family, along with ph ...
, which sits above a photosynthetic reaction center. There are
phycocyanin Phycocyanin is a pigment-protein complex from the light-harvesting phycobiliprotein family, along with allophycocyanin and phycoerythrin. It is an accessory pigment to chlorophyll. All phycobiliproteins are water-soluble, so they cannot exist ...
and
phycoerythrin Phycoerythrin (PE) is a red protein-pigment complex from the light-harvesting phycobiliprotein family, present in cyanobacteria, red algae and Cryptomonad, cryptophytes, accessory to the main chlorophyll pigments responsible for photosynthesis.The ...
subunits that radiate out from this center like thin tubes. This increases the surface area of the absorbing section and helps focus and concentrate light energy down into the reaction center to form chlorophyll. The energy transfer from excited electrons absorbed by pigments in the phycoerythrin subunits at the periphery of these antennas appears at the reaction center in less than 100 ps.Light Harvesting by Phycobilisomes Annual Review of Biophysics and Biophysical Chemistry Vol. 14: 47-77 (Volume publication date June 1985)


See also

*
Photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
* Photosynthetic reaction center * Photosystem II light-harvesting protein * Light harvesting pigment


References


Further reading

* Caffarri (2009)Functional architecture of higher plantphotosystem II supercomplexes. ''The EMBO Journal'' 28: 3052–3063 * Govindjee & Shevela (2011) Adventures with cyanobacteria: a personal perspective. ''Frontiers in Plant Science''. * Liu et al. (2004) Crystal structure of spinach major light-harvesting complex at 2.72A° resolution. ''Nature'' 428: 287–292. * Lokstein (1994)The role of light-harvesting complex II energy dissipation: an in-vivo fluorescence in excess excitation study on the origin of high-energy quenching. Journal of Photochemistry and Photobiology 26: 175-184 * MacColl (1998) Cyanobacterial Phycobilisomes. JOURNAL OF STRUCTURAL BIOLOGY 124(2-3): 311-34.


External links

*
Photosynthesis and all sub categories
{{Carrier proteins Photosynthesis Transmembrane proteins