Photosystems are functional and structural units of

protein complex A protein complex or multiprotein complex is a group of two or more associated polypeptide chains. Protein complexes are distinct from multienzyme complexes, in which multiple catalytic domains are found in a single polypeptide chain. Protein ...

es involved in

photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...

. Together they carry out the primary

photochemistry Photochemistry is the branch of chemistry concerned with the chemical effects of light. Generally, this term is used to describe a chemical reaction caused by absorption of ultraviolet ( wavelength from 100 to 400 nm), visible light (400� ...

: the absorption of light and the transfer of

energy In physics, energy (from Ancient Greek: ἐνέργεια, ''enérgeia'', “activity”) is the quantitative property that is transferred to a body or to a physical system, recognizable in the performance of work and in the form of ...

and

electrons The electron ( or ) is a subatomic particle with a negative one elementary electric charge. Electrons belong to the first generation of the lepton particle family, and are generally thought to be elementary particles because they have n ...

. Photosystems are found in the

thylakoid membrane Thylakoids are membrane-bound compartments inside chloroplasts and cyanobacteria. They are the site of the light-dependent reactions of photosynthesis. Thylakoids consist of a thylakoid membrane surrounding a thylakoid lumen. Chloroplast thyla ...

s of plants, algae, and cyanobacteria. These membranes are located inside the

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it i ...

s of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII. PSII will absorb red light, and PSI will absorb far-red light. Although photosynthetic activity will be detected when the photosystems are exposed to either red or far-red light, the photosynthetic activity will be the greatest when plants are exposed to both

wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tr ...

s of light. Studies have actually demonstrated that the two wavelengths together have a synergistic effect on the photosynthetic activity, rather than an additive one. Each photosystem has two parts: a reaction center, where the photochemistry occurs, and an

antenna complex The light-harvesting complex (or antenna complex; LH or LHC) is an array of protein and chlorophyll molecules embedded in the thylakoid membrane of plants and cyanobacteria, which transfer light energy to one chlorophyll ''a'' molecule at the reac ...

, which surrounds the reaction center. The antenna complex contains hundreds of

chlorophyll Chlorophyll (also chlorophyl) 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 allow plants to ...

molecules which funnel the excitation energy to the center of the photosystem. At the reaction center, the energy will be trapped and transferred to produce a high energy molecule. The main function of PSII is to efficiently split water into oxygen molecules and protons. PSII will provide a steady stream of electrons to PSI, which will boost these in energy and transfer them to NADP and H to make

NADPH Nicotinamide adenine dinucleotide phosphate, abbreviated NADP or, in older notation, TPN (triphosphopyridine nucleotide), is a cofactor used in anabolic reactions, such as the Calvin cycle and lipid and nucleic acid syntheses, which require NA ...

. The hydrogen from this NADPH can then be used in a number of different processes within the plant.

Reaction centers

Reaction centers are multi-protein complexes found within the thylakoid membrane. At the heart of a photosystem lies the

reaction center A photosynthetic reaction center is a complex of several proteins, pigments and other co-factors that together execute the primary energy conversion reactions of photosynthesis. Molecular excitations, either originating directly from sunlight or ...

, which is an

enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...

that uses light to reduce and oxidize molecules (give off and take up electrons). This reaction center is surrounded by light-harvesting complexes that enhance the absorption of light. In addition, surrounding the reaction center are

pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...

s which will absorb light. The pigments which absorb light at the highest energy level are found furthest from the reaction center. On the other hand, the pigments with the lowest energy level are more closely associated with the reaction center. Energy will be efficiently transferred from the outer part of the antenna complex to the inner part. This funneling of energy is performed via resonance transfer, which occurs when energy from an excited molecule is transferred to a molecule in the ground state. This ground state molecule will be excited, and the process will continue between molecules all the way to the reaction center. At the reaction center, the electrons on the special chlorophyll molecule will be excited and ultimately transferred away by electron carriers. (If the electrons were not transferred away after excitation to a high energy state, they would lose energy by fluorescence back to the ground state, which would not allow plants to drive photosynthesis.) The reaction center will drive photosynthesis by taking light and turning it into chemical energy that can then be used by the chloroplast. Two families of reaction centers in photosystems can be distinguished: type I reaction centers (such as

photosystem I Photosystem I (PSI, or plastocyanin–ferredoxin oxidoreductase) is one of two photosystems in the photosynthetic light reactions of algae, plants, and cyanobacteria. Photosystem I is an integral membrane protein complex that us ...

(

P700 P700, or photosystem I primary donor, is the reaction-center chlorophyll ''a'' molecular dimer associated with photosystem I in plants, algae, and cyanobacteria. Etymology Its name is derived from the word “pigment” (P) and the presence ...

) in chloroplasts and in green-sulfur bacteria) and type II reaction centers (such as

photosystem II Photosystem II (or water-plastoquinone oxidoreductase) is the first protein complex in the light-dependent reactions of oxygenic photosynthesis. It is located in the thylakoid membrane of plants, algae, and cyanobacteria. Within the photosyst ...

(

P680 P680, or photosystem II primary donor, is the reaction-center chlorophyll ''a'' molecular dimer associated with photosystem II in plants, algae, and cyanobacteria, and central to oxygenic photosynthesis. Etymology Its name is derived from th ...

) in chloroplasts and in non-sulfur purple bacteria). The two photosystems originated from a common ancestor, but have since diversified. Each of the photosystem can be identified by the

of light to which it is most reactive (700

nanometer 330px, Different lengths as in respect to the molecular scale. The nanometre (international spelling as used by the International Bureau of Weights and Measures; SI symbol: nm) or nanometer (American and British English spelling differences#-re, ...

s for PSI and 680 nanometers for PSII in chloroplasts), the amount and type of light-harvesting complex present, and the type of terminal electron acceptor used. Type I photosystems use

ferredoxin Ferredoxins (from Latin ''ferrum'': iron + redox, often abbreviated "fd") are iron–sulfur proteins that mediate electron transfer in a range of metabolic reactions. The term "ferredoxin" was coined by D.C. Wharton of the DuPont Co. and applied t ...

-like iron-sulfur cluster proteins as terminal electron acceptors, while type II photosystems ultimately shuttle electrons to a

quinone The quinones are a class of organic compounds that are formally "derived from aromatic compounds uch as benzene or naphthalene">benzene.html" ;"title="uch as benzene">uch as benzene or naphthalene] by conversion of an even number of –CH= group ...

terminal electron acceptor. Both reaction center types are present in chloroplasts and cyanobacteria, and work together to form a unique photosynthetic chain able to extract electrons from water, creating oxygen as a byproduct.

Structure of PSI and PSII

A reaction center comprises several (about 25-30) protein subunits, which provide a scaffold for a series of cofactors. The cofactors can be pigments (like

pheophytin Pheophytin or phaeophytin is a chemical compound that serves as the first electron carrier intermediate in the electron transfer pathway of Photosystem II (PS II) in plants, and the type II photosynthetic reaction center (RC P870) found in pur ...

carotenoids Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, co ...

), quinones, or

iron-sulfur clusters Iron–sulfur proteins (or iron–sulphur proteins in British spelling) are proteins characterized by the presence of iron–sulfur clusters containing sulfide-linked di-, tri-, and tetrairon centers in variable oxidation states. Iron–sulfur cl ...

. Each photosystem has two main subunits: an

(a light harvesting complex or LHC) and a reaction center. The antenna complex is where light is captured, while the reaction center is where this light energy is transformed into chemical energy. At the reaction center, there are many polypeptides that are surrounded by pigment proteins. At the center of the reaction center is a special pair of chlorophyll molecules. Each PSII has about 8 LHCII. These contain about 14 chlorophyll ''a'' and chlorophyll ''b'' molecules, as well as about four

carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, ...

s. In the reaction center of PSII of plants and cyanobacteria, the light energy is used to split water into oxygen, protons, and electrons. The protons will be used in proton pumping to fuel the ATP synthase at the end of an

electron transport chain An electron transport chain (ETC) is a series of protein complexes and other molecules that transfer electrons from electron donors to electron acceptors via redox reactions (both reduction and oxidation occurring simultaneously) and couples ...

. A majority of the reactions occur at the D1 and D2 subunits of PSII.

In oxygenic photosynthesis

Both

and II are required for oxygenic photosynthesis. Oxygenic photosynthesis can be performed by plants and cyanobacteria; cyanobacteria are believed to be the progenitors of the photosystem-containing chloroplasts of

eukaryotes Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...

. Photosynthetic bacteria that cannot produce oxygen have only one photosystem, which is similar to either PSI or PSII. At the core of photosystem II is P680, a special chlorophyll to which incoming excitation energy from the antenna complex is funneled. One of the electrons of excited P680* will be transferred to a non-

fluorescent Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, ...

molecule, which ionizes the chlorophyll and boosts its energy further, enough that it can split water in the oxygen evolving complex (OEC) of PSI and recover its electron. At the heart of the OEC are 4 Mn atoms, each of which can trap one electron. The electrons harvested from the splitting of two waters fill the OEC complex in its highest-energy state, which holds 4 excess electrons. Electrons travel through the cytochrome ''b6f'' complex to photosystem I via an electron transport chain within the

. Energy from PSI drives this process and is harnessed (the whole process is termed

chemiosmosis Chemiosmosis is the movement of ions across a semipermeable membrane bound structure, down their electrochemical gradient. An important example is the formation of adenosine triphosphate (ATP) by the movement of hydrogen ions (H+) across a memb ...

) to pump protons across the membrane, into the thylakoid lumen space from the chloroplast stroma. This will provide a potential energy difference between lumen and stroma, which amounts to a proton-motive force that can be utilized by the proton-driven

ATP synthase ATP synthase is a protein that catalyzes the formation of the energy storage molecule adenosine triphosphate (ATP) using adenosine diphosphate (ADP) and inorganic phosphate (Pi). It is classified under ligases as it changes ADP by the formation ...

to generate ATP. If electrons only pass through once, the process is termed noncyclic photophosphorylation, but if they pass through PSI and the proton pump multiple times it is called cyclic photophosphorylation. When the electron reaches photosystem I, it fills the electron deficit of light-excited reaction-center chlorophyll P700 of PSI. The electron may either continue to go through cyclic electron transport around PSI or pass, via ferredoxin, to the enzyme NADP reductase. Electrons and protons are added to NADP to form NADPH. This reducing (hydrogenation) agent is transported to the Calvin cycle to react with glycerate 3-phosphate, along with ATP to form

glyceraldehyde 3-phosphate Glyceraldehyde 3-phosphate, also known as triose phosphate or 3-phosphoglyceraldehyde and abbreviated as G3P, GA3P, GADP, GAP, TP, GALP or PGAL, is a metabolite that occurs as an intermediate in several central pathways of all organisms.Nelson, D ...

, the basic building block from which plants can make a variety of substances.

Photosystem repair

In intense light, plants use various mechanisms to prevent damage to their photosystems. They are able to release some light energy as heat, but the excess light can also produce

reactive oxygen species In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen. The reduction of molecular oxygen () p ...

. While some of these can be detoxified by

antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...

s, the remaining oxygen species will be detrimental to the photosystems of the plant. More specifically, the D1 subunit in the reaction center of PSII can be damaged. Studies have found that

deg1 tRNA pseudouridine38/39 synthase (, ''Deg1'', ''Pus3p'', ''pseudouridine synthase 3'') is an enzyme with systematic name ''tRNA-uridine38/39 uracil mutase''. This enzyme catalyses the following chemical reaction : tRNA uridine38/39 \rightleftharp ...

proteins are involved in the degradation of these damaged D1 subunits. New D1 subunits can then replace these damaged D1 subunits in order to allow PSII to function properly again.

References

External links