Sucrose-phosphate synthase (SPS) is a

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 ...

enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...

involved in

sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...

biosynthesis Biosynthesis, i.e., chemical synthesis occurring in biological contexts, is a term most often referring to multi-step, enzyme-Catalysis, catalyzed processes where chemical substances absorbed as nutrients (or previously converted through biosynthe ...

. Specifically, this enzyme

catalyzes Catalysis () is the increase in rate of a chemical reaction due to an added substance known as a catalyst (). Catalysts are not consumed by the reaction and remain unchanged after it. If the reaction is rapid and the catalyst recycles quick ...

the transfer of a hexosyl group from uridine diphosphate glucose ( UDP-glucose) to D-

fructose 6-phosphate Fructose 6-phosphate (sometimes called the Neuberg ester) is a derivative of fructose, which has been phosphorylated at the 6-hydroxy group. It is one of several possible fructosephosphates. The β-D-form of this compound is very common in cells ...

to form UDP and D-sucrose-6-phosphate. This reversible step acts as the key regulatory control point in sucrose biosynthesis, and is an excellent example of various key enzyme regulation strategies such as

allosteric In the fields of biochemistry and pharmacology an allosteric regulator (or allosteric modulator) is a substance that binds to a site on an enzyme or receptor distinct from the active site, resulting in a conformational change that alters the p ...

control and reversible

phosphorylation In biochemistry, phosphorylation is described as the "transfer of a phosphate group" from a donor to an acceptor. A common phosphorylating agent (phosphate donor) is ATP and a common family of acceptor are alcohols: : This equation can be writ ...

. This enzyme participates in

starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diet ...

and sucrose

metabolism Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the co ...

Nomenclature

This enzyme belongs to the family of

glycosyltransferase Glycosyltransferases (GTFs, Gtfs) are enzymes ( EC 2.4) that establish natural glycosidic linkages. They catalyze the transfer of saccharide moieties from an activated nucleotide sugar (also known as the "glycosyl donor") to a nucleophilic gl ...

s, specifically the hexosyltransferases. The

systematic name A systematic name is a name given in a systematic way to one unique group, organism, object or chemical substance, out of a specific population or collection. Systematic names are usually part of a nomenclature. A semisystematic name or semitrivi ...

of this enzyme class is UDP-glucose:D-fructose 6-phosphate 2-alpha-D-glucosyltransferase. Other names in common use include UDP-glucose-fructose-phosphate glucosyltransferase, sucrosephosphate-UDP glucosyltransferase, UDP-glucose-fructose-phosphate glucosyltransferase, SPS, uridine diphosphoglucose-fructose phosphate glucosyltransferase, sucrose 6-phosphate synthase, sucrose phosphate synthetase, and sucrose phosphate-uridine diphosphate glucosyltransferase.

Structure

X-ray diffraction X-ray diffraction is a generic term for phenomena associated with changes in the direction of X-ray beams due to interactions with the electrons around atoms. It occurs due to elastic scattering, when there is no change in the energy of the waves. ...

studies have revealed that the

structure A structure is an arrangement and organization of interrelated elements in a material object or system, or the object or system so organized. Material structures include man-made objects such as buildings and machines and natural objects such as ...

of ''Halothermothrix orenii'' SPS belongs to the GT-B fold family. Like other GT-B proteins, SPS contains two Rossmann fold domains that are named the A domain and the B domain. Generally, the structure of these domains are somewhat similar, as both contain central beta sheets that are surrounded by

alpha helices An alpha helix (or α-helix) is a sequence of amino acids in a protein that are twisted into a coil (a helix). The alpha helix is the most common structural arrangement in the secondary structure of proteins. It is also the most extreme type of l ...

. However, the A domain consists of eight parallel beta strands and seven alpha helices while the B domain contains six parallel beta strands and nine alpha helices. These domains are joined by residue loops to form a

substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...

binding cleft, where the glucosyl group acceptor binds. Although ''H. orenii'' is a non-

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

bacterium, various studies indicate that the structure of its SPS is similar to plant SPS. First,

antibodies An antibody (Ab) or immunoglobulin (Ig) is a large, Y-shaped protein belonging to the immunoglobulin superfamily which is used by the immune system to identify and neutralize antigens such as bacteria and viruses, including those that caus ...

with high specificities for plant SPS also target the bacterial SPS, indicating the structure is conserved enough for the antibody to recognize the enzyme as an antigen. Furthermore,

genomic Genomics is an interdisciplinary field of molecular biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, ...

studies reveal that closely related plant homologues exhibit up to 54% sequence identities.

Mechanism

In the open conformation of ''H. orenii'' SPS, fructose 6-phosphate forms

hydrogen bonds In chemistry, a hydrogen bond (H-bond) is a specific type of molecular interaction that exhibits partial covalent character and cannot be described as a purely electrostatic force. It occurs when a hydrogen (H) atom, covalently bonded to a mo ...

with Gly-33 and Gln-35 residues in the A domain while UDP-glucose interacts with the B-domain. Crystal structures studies reveal that after binding, the two domains twist to narrow the entrance of the substrate binding cleft from 20 Å to 6 Å. In this closed conformation, the Gly-34 residue of the A domain interacts with UDP-glucose and forces the substrate to adapt a folded structure, facilitating its donation of the hexosyl group. After binding, fructose 6-phosphate will interact with UDP via a hydrogen bond, which lowers the

activation energy In the Arrhenius model of reaction rates, activation energy is the minimum amount of energy that must be available to reactants for a chemical reaction to occur. The activation energy (''E''a) of a reaction is measured in kilojoules per mole (k ...

of the

reaction Reaction may refer to a process or to a response to an action, event, or exposure. Physics and chemistry *Chemical reaction *Nuclear reaction *Reaction (physics), as defined by Newton's third law * Chain reaction (disambiguation) Biology and ...

and stabilizes the

transition state In chemistry, the transition state of a chemical reaction is a particular configuration along the reaction coordinate. It is defined as the state corresponding to the highest potential energy along this reaction coordinate. It is often marked w ...

. Finally, the C1 atom of UDP-glucose undergoes

nucleophilic attack In chemistry, a nucleophile is a chemical species that forms bonds by donating an electron pair. All molecules and ions with a free pair of electrons or at least one pi bond can act as nucleophiles. Because nucleophiles donate electrons, they a ...

by an oxygen atom in fructose 6-phosphate, resulting in glucosyl group transfer to fructose 6-phosphate. Whether or not this mechanism requires a divalent ion is currently unclear, but failed attempts to trap and detect the presence of the

magnesium Magnesium is a chemical element; it has Symbol (chemistry), symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 ...

cation An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by convent ...

suggest that this mechanism is metal ion independent.

Regulatory strategies

Phosphorylation

SPS-

kinase In biochemistry, a kinase () is an enzyme that catalyzes the transfer of phosphate groups from high-energy, phosphate-donating molecules to specific substrates. This process is known as phosphorylation, where the high-energy ATP molecule don ...

reversibly phosphorylates a

serine Serine (symbol Ser or S) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α- amino group (which is in the protonated − form under biological conditions), a carboxyl group (which is in the deprotonated − ...

residue and subsequently deactivates SPS, In

spinach Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to Central Asia, Central and Western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common vegetable consumed eit ...

and

maize Maize (; ''Zea mays''), also known as corn in North American English, is a tall stout grass that produces cereal grain. It was domesticated by indigenous peoples in southern Mexico about 9,000 years ago from wild teosinte. Native American ...

, the site of phosphorylation regulation has been identified as Ser158 and Ser162 respectively. While it is currently unclear if this seryl residue homolog in other plant SPSes is phosphorylated to suppress SPS activity, conservation of the neighboring residues has been observed in other plant species. This conserved sequence may potentially aid in recognition of a regulatory SPS-kinase. Once phosphorylated, the inactivated enzyme can be dephosphorylated and reactivated by SPS-phosphatase. Aside from controlling the levels of sucrose in the cell, regulation via phosphorylation can help the cell adapt to hyperosmotic conditions; in times of

osmotic stress Osmotic shock or osmotic stress is physiologic dysfunction caused by a sudden change in the solute concentration around a cell, which causes a rapid change in the movement of water across its cell membrane. Under hypertonic conditions - conditi ...

, the seryl residue is phosphorylated and enzyme activity decreases. This regulation strategy also controls

carbon Carbon () is a chemical element; it has chemical symbol, symbol C and atomic number 6. It is nonmetallic and tetravalence, tetravalent—meaning that its atoms are able to form up to four covalent bonds due to its valence shell exhibiting 4 ...

flux from photosynthesis, as studies indicate the

signal transduction pathway A signal is both the process and the result of transmission of data over some media accomplished by embedding some variation. Signals are important in multiple subject fields including signal processing, information theory and biology. In ...

responsible for SPS activation responds to

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– ...

stimulus.

Allostery

Glucose 6-phosphate binds to an allosteric site, resulting in conformational changes to SPS that increase the enzyme's

affinity Affinity may refer to: Commerce, finance and law * Affinity (law), kinship by marriage * Affinity analysis, a market research and business management technique * Affinity Credit Union, a Saskatchewan-based credit union * Affinity Equity Pa ...

for the glucosyl accepting substrate. Inorganic phosphate can also bind to this allosteric site, preventing glucose 6-phosphate activation of SPS. Like regulation via phosphorylation, this regulation strategy is also closely related to photosynthesis, as high rates of photosynthesis will deplete levels of inorganic phosphate and increase concentrations of glucose 6-phosphate in the

chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...

. Overall, increased rates of photosynthesis will increase SPS activity.

Function

SPS plays a major role in partitioning

between sucrose and

in photosynthetic and non-photosynthetic tissues, affecting the growth and development of the plant. In ripening fruits, SPS is responsible for converting starch to sucrose and other soluble sugars. Additionally, SPS is also active in cells that mostly degrade sucrose, participating in futile cycles that allow for large, rapid changes in sucrose

flux Flux describes any effect that appears to pass or travel (whether it actually moves or not) through a surface or substance. Flux is a concept in applied mathematics and vector calculus which has many applications in physics. For transport phe ...

. At low temperature, SPS activity and sucrose biosynthesis rates are increased. Sucrose accumulation is advantageous at low temperature, as sucrose is a form of energy storage that can be rapidly metabolized for respiratory purposes. Furthermore, increased amounts of sucrose can help the plant withstand freezing.

References

{{Portal bar, Biology, border=no EC 2.4.1 Enzymes of unknown structure