Photosystems are functional and structural units of protein complexes involved in photosynthesis that together carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. They are found in the thylakoid membranes of plants, algae and cyanobacteria (in plants and algae these are located in the chloroplasts), or in the cytoplasmic membrane of photosynthetic bacteria.
At the heart of a photosystem lies the reaction center, which is an enzyme that uses light to reduce molecules. In a photosystem, this reaction center is surrounded by light-harvesting complexes that enhance the absorption of light and transfer the energy to the reaction centers. Light-harvesting and reaction center complexes are membrane protein complexes that are made of several protein-subunits and contain numerous cofactors.
In the photosynthetic membranes, reaction centers provide the driving force for the bioenergetic electron and proton transfer chain. When light is absorbed by a reaction center (either directly or passed by neighbouring pigment-antennae), a series of oxido-reduction reactions is initiated, leading to the reduction of a terminal acceptor.
Two families of reaction centers in photosystems exist: type I reaction centers (such as photosystem I (P700) in chloroplasts and in green-sulphur bacteria) and type II reaction centers (such as photosystem II (P680) in chloroplasts and in non-sulphur purple bacteria).
Each photosystem can be identified by the wavelength of light to which it is most reactive (700 and 680 nanometers, respectively for PSI and PSII in chloroplasts), the amount and type of light-harvesting complexes present and the type of terminal electron acceptor used.
Type I photosystems use ferredoxin-like iron-sulfur cluster proteins as terminal electron acceptors, while type II photosystems ultimately shuttle electrons to a quinone terminal electron acceptor. One has to note that both reaction center types are present in chloroplasts and cyanobacteria, working together to form a unique photosynthetic chain able to extract electrons from water, creating oxygen as a byproduct.
A reaction center comprises several (>10 or >11) protein subunits, providing a scaffold for a series of cofactors. The latter can be pigments (like chlorophyll, pheophytin, carotenoids), quinones, or iron-sulfur clusters.
Relationship between Photosystems I and II
For oxygenic photosynthesis, both photosystems I and II are required. Oxygenic photosynthesis can be performed by plants and cyanobacteria; cyanobacteria are believed to be the progenitors of the photosystem-containing chloroplasts of eukaryotes. Photosynthetic bacteria that cannot produce oxygen have a single photosystem called BRC, bacterial reaction center.
The photosystem I was named "I" since it was discovered before photosystem II, but this does not represent the order of the electron flow.
When photosystem II absorbs light, electrons in the reaction-center chlorophyll are excited to a higher energy level and are trapped by the primary electron acceptors. To replenish the deficit of electrons, electrons are extracted from water by a cluster of four Manganese ions in photosystem II and supplied to the chlorophyll via a redox-active tyrosine.
Photoexcited electrons travel through the cytochrome b6f complex to photosystem I via an electron transport chain set in the thylakoid membrane. This energy fall is harnessed, (the whole process termed chemiosmosis), to transport hydrogen (H+) through the membrane, to the lumen, to provide a proton-motive force to generate ATP. The protons are transported by the plastoquinone. If electrons only pass through once, the process is termed noncyclic photophosphorylation.
When the electron reaches photosystem I, it fills the electron deficit of the reaction-center chlorophyll of photosystem I. The deficit is due to photo-excitation of electrons that are again trapped in an electron acceptor molecule, this time that of photosystem I.
ATP is generated when the ATP synthase transports the protons present in the lumen to the stroma, through the membrane. The electrons may either continue to go through cyclic electron transport around PS I or pass, via ferredoxin, to the enzyme NADP+ reductase. Electrons and hydrogen ions are added to NADP+ to form NADPH. This reducing agent is transported to the Calvin cycle to react with glycerate 3-phosphate, along with ATP to form glyceraldehyde 3-phosphate, the basic building-block from which plants can make a variety of substances.
- Photosynthetic reaction centre
- Light reaction
- Photosystem I
- Photosystem II
- Photosystems I + II: Imperial College, Barber Group
- Photosystem I: Molecule of the Month in the Protein Data Bank
- Photosystem II: Molecule of the Month in the Protein Data Bank
- Photosystem II: ANU
- UMich Orientation of Proteins in Membranes families/superfamily-2 - Calculated spatial positions of photosynthetic reaction centers and photosystems in membrane
- Rutherford AW, Faller P (January 2003). "Photosystem II: evolutionary perspectives". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 358 (1429): 245–53. doi:10.1098/rstb.2002.1186. PMC 1693113. PMID 12594932.