Photosynthetic state transition

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In photosynthesis, state transitions are rearrangements of the photosynthetic apparatus which occur on short time-scales (seconds to minutes). The effect is prominent in cyanobacteria, whereby the phycobilisome light-harvesting antenna complexes alter their preference for transfer of excitation energy between the two reaction centers, PS I and PS II. [1] This shift helps to minimize photodamage caused by reactive oxygen species (ROS) under stressful conditions such as high light, but may also be used to offset imbalances between the rates of generating reductant and ATP.

The phenomenon was first discovered in unicellular green algae, [2] and may also occur in plants. [3] However, in these organisms it occurs by a different mechanism, which is not as well understood. The plant/algal mechanism is considered functionally analogous to the cyanobacterial mechanism but involves completely different components. The foremost difference is the presence of fundamentally different types of light-harvesting antenna complexes: plants and green algae use an intrinsically-bound membrane complex of chlorophyll a/b binding proteins for their antenna, instead of the soluble phycobilisome complexes used by cyanobacteria (and certain algae).

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Photosynthesis Biological process to convert light into chemical energy

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 carbohydrate molecules, such as sugars and starches, which are synthesized from carbon dioxide and water – hence the name photosynthesis, from the Greek phōs, "light", and sunthesis, "putting together". In most cases, oxygen is also released as a waste product that stores three times more chemical energy than the carbohydrates. Most plants, algae, and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies most of the energy necessary for life on Earth.

Cyanobacteria Phylum of photosynthesising prokaryotes

Cyanobacteria, also known as Cyanophyta, are a phylum of Gram-negative bacteria that obtain energy via photosynthesis. The name cyanobacteria refers to their color, giving them their other name, "blue-green algae", though modern botanists restrict the term algae to eukaryotes and do not apply it to cyanobacteria, which are prokaryotes. They appear to have originated in freshwater or a terrestrial environment. Sericytochromatia, the proposed name of the paraphyletic and most basal group, is the ancestor of both the non-photosynthetic group Melainabacteria and the photosynthetic cyanobacteria, also called Oxyphotobacteria.

Plastid Plant cell organelles that perform photosynthesis and store starch

The plastid is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyanobacteria. Examples include chloroplasts, chromoplasts, and leucoplasts.

Thylakoid Membrane enclosed compartments in chloroplasts and cyanobacteria

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 thylakoids frequently form stacks of disks referred to as grana. Grana are connected by intergranal/stromal thylakoids, which join granum stacks together as a single functional compartment.

<i>Chloroflexus aurantiacus</i> Species of bacterium

Chloroflexus aurantiacus is a photosynthetic bacterium isolated from hot springs, belonging to the green non-sulfur bacteria. This organism is thermophilic and can grow at temperatures from 35 °C to 70 °C. Chloroflexus aurantiacus can survive in the dark if oxygen is available. When grown in the dark, Chloroflexus aurantiacus has a dark orange color. When grown in sunlight it is dark green. The individual bacteria tend to form filamentous colonies enclosed in sheaths, which are known as trichomes.

Photosystem

Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.

Photosystem I Second protein complex in photosynthetic light reactions

Photosystem I is one of two photosystems in the photosynthetic light reactions of algae, plants, and cyanobacteria. Photosystem I is an integral membrane protein complex that uses light energy to catalyze the transfer of electrons across the thylakoid membrane from plastocyanin to ferredoxin. Ultimately, the electrons that are transferred by Photosystem I are used to produce the moderate-energy hydrogen carrier NADPH. The photon energy absorbed by Photosystem I also produces a proton-motive force that is used to generate ATP. PSI is composed of more than 110 cofactors, significantly more than Photosystem II.

Phycobilisome Light-energy harvesting structure in cyanobacteria and red algae

Phycobilisomes are light harvesting antennae of photosystem II in cyanobacteria, red algae and glaucophytes. It was lost in the plastids of green algae/ plants (chloroplasts).

Photosynthetic reaction centre

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 transferred as excitation energy via light-harvesting antenna systems, give rise to electron transfer reactions along the path of a series of protein-bound co-factors. These co-factors are light-absorbing molecules (also named chromophores or pigments) such as chlorophyll and pheophytin, as well as quinones. The energy of the photon is used to excite an electron of a pigment. The free energy created is then used, via a chain of nearby electron acceptors, for a transfer of hydrogen atoms (as protons and electrons) from H2O or hydrogen sulfide towards carbon dioxide, eventually producing glucose. These electron transfer steps ultimately result in the conversion of the energy of photons to energy stored in relatively weak chemical bonds.

A light-harvesting complex consists of a number of chromophores which are complex subunit proteins that may be part of a larger super complex of a photosystem, the functional unit in photosynthesis. It is used by plants and photosynthetic bacteria to collect more of the incoming light than would be captured by the photosynthetic reaction center alone. The light which is captured by the chromophores is capable of exciting molecules from their ground state to a higher energy state, known as the excited state. This excited state does not last very long and is known to be short-lived. Light-harvesting complexes are found in a wide variety among the different photosynthetic species. The complexes consist of proteins and photosynthetic pigments and surround a photosynthetic reaction center to focus energy, attained from photons absorbed by the pigment, toward the reaction center using Förster resonance energy transfer.

Photoprotection is the biochemical process that helps organisms cope with molecular damage caused by sunlight. Plants and other oxygenic phototrophs have developed a suite of photoprotective mechanisms to prevent photoinhibition and oxidative stress caused by excess or fluctuating light conditions. Humans and other animals have also developed photoprotective mechanisms to avoid UV photodamage to the skin, prevent DNA damage, and minimize the downstream effects of oxidative stress.

P700, or photosystem I primary donor, is the reaction-center chlorophyll a molecular dimer associated with photosystem I in plants, algae, and cyanobacteria.

Biohydrogen

Biohydrogen is H2 that is produced biologically. Interest is high in this technology because H2 is a clean fuel and can be readily produced from certain kinds of biomass.

Photoinhibition

Photoinhibition is light-induced reduction in the photosynthetic capacity of a plant, alga, or cyanobacterium. Photosystem II (PSII) is more sensitive to light than the rest of the photosynthetic machinery, and most researchers define the term as light-induced damage to PSII. In living organisms, photoinhibited PSII centres are continuously repaired via degradation and synthesis of the D1 protein of the photosynthetic reaction center of PSII. Photoinhibition is also used in a wider sense, as dynamic photoinhibition, to describe all reactions that decrease the efficiency of photosynthesis when plants are exposed to light.

Bacterial antenna complex

Bacterial antenna complex proteins are the main light-absorbing components in photosynthetic bacteria. Also known as a light-harvesting complex/system, the bacterial antenna complex is responsible for the transfer of solar energy to the photosynthetic reaction centre.

Non-photochemical quenching (NPQ) is a mechanism employed by plants and algae to protect themselves from the adverse effects of high light intensity. It involves the quenching of singlet excited state chlorophylls (Chl) via enhanced internal conversion to the ground state, thus harmlessly dissipating excess excitation energy as heat through molecular vibrations. NPQ occurs in almost all photosynthetic eukaryotes, and helps to regulate and protect photosynthesis in environments where light energy absorption exceeds the capacity for light utilization in photosynthesis.

Light-dependent reactions Photosynthetic reactions

In photosynthesis, the light-dependent reactions take place on the thylakoid membranes. The inside of the thylakoid membrane is called the lumen, and outside the thylakoid membrane is the stroma, where the light-independent reactions take place. The thylakoid membrane contains some integral membrane protein complexes that catalyze the light reactions. There are four major protein complexes in the thylakoid membrane: Photosystem II (PSII), cytochrome b6f complex, Photosystem I (PSI), and ATP synthase. These four complexes work together to ultimately produce ATP and NADPH.

Orange carotenoid protein

Orange carotenoid protein (OCP) is a water-soluble protein which plays a role in photoprotection in diverse cyanobacteria. It is the only photoactive protein known to use a carotenoid as the photoresponsive chromophore. The protein consists of two domains, with a single keto-carotenoid molecule non-covalently bound between the two domains. It is a very efficient quencher of excitation energy absorbed by the primary light-harvesting antenna complexes of cyanobacteria, the phycobilisomes. The quenching is induced by blue-green light. It is also capable of preventing oxidative damage by directly scavenging singlet oxygen (1O2).

Photoautotrophs are organisms that use light energy and inorganic carbon to produce organic materials. Eukaryotic photoautotrophs absorb energy through the chlorophyll molecules in their chloroplasts while prokaryotic photoautotrophs use chlorophylls and bacteriochlorophylls present in their cytoplasm. All known photoautotrophs perform photosynthesis. Examples include plants, algae, and cyanobacteria.

Alexander Glazer was a Professor of the Graduate School in the Department of Molecular and Cell Biology at the University of California, Berkeley. He had a passion for protein chemistry and structure function relationships. He also had a longstanding interest in light-harvesting complexes in cyanobacteria and red algae called phycobilisomes. He had also spent more than 10 years working on the human genome project where he has investigated methods for DNA detection and sequencing which most notably includes the development of fluorescent reagents involved in cell labeling. Most recently, he had focused his studies on issues in environmental sciences. He died on July 18th 2021 in Orinda, California

References

  1. Biggins, John; Bruce, Doug (1989). "Regulation of excitation energy transfer in organisms containing phycobilins". Photosynthesis Research. 20 (1): 1–34. doi:10.1007/BF00028620. ISSN   0166-8595. PMID   24425462.
  2. Bonaventura, Celia; Myers, Jack (1969). "Fluorescence and oxygen evolution from Chlorella pyrenoidosa". Biochimica et Biophysica Acta (BBA) - Bioenergetics. 189 (3): 366–383. doi:10.1016/0005-2728(69)90168-6. ISSN   0005-2728. PMID   5370012.
  3. Ruban, Alexander V.; Johnson, Matthew P. (2008). "Dynamics of higher plant photosystem cross-section associated with state transitions". Photosynthesis Research. 99 (3): 173–183. doi:10.1007/s11120-008-9387-x. ISSN   0166-8595. PMID   19037743.