Xyloglucan
Encyclopedia
Xyloglucan is a hemicellulose
Hemicellulose
A hemicellulose is any of several heteropolymers , such as arabinoxylans, present along with cellulose in almost all plant cell walls. While cellulose is crystalline, strong, and resistant to hydrolysis, hemicellulose has a random, amorphous structure with little strength...

 that occurs in the primary cell wall of all vascular plant
Vascular plant
Vascular plants are those plants that have lignified tissues for conducting water, minerals, and photosynthetic products through the plant. Vascular plants include the clubmosses, Equisetum, ferns, gymnosperms and angiosperms...

s, however essential enzymes for xyloglucan metabolism, like XTH and β1→4-Glucan Synthase are found in Charophyceae
Charophyceae
Charophyceae is a taxon of green algae whose exact rank is the matter of some current debate. Some botanists recommend expanding the existing plant kingdom to include charophyceans and chlorophytes while others consider Charophyceae to be a class within either the divisions Chlorophyta,...

 algae. In many dicotyledon
Dicotyledon
The dicotyledons, also known as dicots, are a group of flowering plants whose seed typically has two embryonic leaves or cotyledons. There are around 199,350 species within this group...

ous plants, it is the most abundant hemicellulose in the primary cell wall. Xyloglucan binds to the surface of cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

 microfibril
Microfibril
The microfibril is a very fine fibril, or fiber-like strand, consisting of glycoproteins and cellulose. It is usually, but not always, used as a general term in describing the structure of protein fiber, examples are hair and sperm tail. Its most frequently observed structural pattern is 9+2...

s and may link them together. It is the target of xyloglucan endotransglycosylase
Xyloglucan:xyloglucosyl transferase
In enzymology, a xyloglucan:xyloglucosyl transferase is an enzyme that catalyzes the chemical reaction in which a beta- bond in the backbone of a xyloglucan in broken; the xyloglucanyl segment is then transferred to the O4 of the non-reducing terminal glucose residue of either xyloglucan or an...

, which cuts and ligates xyloglucans, as a means of integrating new xyloglucans into the cell wall. It is also thought to be the target of alpha-expansin
Expansin
Expansin refers to a family of closely related nonenzymatic proteins found in the plant cell wall, with important roles in plant cell growth, fruit softening, abscission, emergence of root hairs, pollen tube invasion of the stigma and style, meristem function, and other developmental processes...

, which promotes cell wall enlargement.

Chemistry

Xyloglucan has a backbone of β1→4-linked glucose
Glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

 residues, most of which substituted with 1-6 linked xylose
Xylose
Xylose is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional group. It is the precursor to hemicellulose, one of the main constituents of biomass...

 sidechains. The xylose residues are often capped with a galactose
Galactose
Galactose , sometimes abbreviated Gal, is a type of sugar that is less sweet than glucose. It is a C-4 epimer of glucose....

 residue sometimes followed by a fucose
Fucose
Fucose is a hexose deoxy sugar with the chemical formula C6H12O5. It is found on N-linked glycans on the mammalian, insect and plant cell surface, and is the fundamental sub-unit of the fucoidan polysaccharide...

 residue. The specific structure of xyloglucan varies among plant families.

Biosynthesis

Xyloglucan is synthesized in Golgi
Golgi
Golgi may refer to:*Camillo Golgi , Italian physician and scientist after which the following terms are named:**Golgi apparatus , an organelle in the eukaryotic cell...

 trans cisternae and in the trans Golgi network (TGN) and is transported to the cell membrane
Cell membrane
The cell membrane or plasma membrane is a biological membrane that separates the interior of all cells from the outside environment. The cell membrane is selectively permeable to ions and organic molecules and controls the movement of substances in and out of cells. It basically protects the cell...

 by vesicles
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

, where it is expelled and adsorbs on nascent cellulosic microfibrils.
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