Thylakoid

A thylakoid is a membrane-bound compartment 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 (singular: granum). Grana are connected by intergranal or stroma thylakoids, which join granum stacks together as a single functional compartment. The word Thylakoid comes from the Greek word thylakos meaning 'sac' or 'pouch'. Thus, thylakoid means 'sac-like' or 'pouch-like'. Thylakoids are membrane-bound structures embedded in the chloroplast stroma. A stack of thylakoids is called a granum and resembles a stack of coins. The thylakoid membrane is the site of the light-dependent reactions of photosynthesis with the photosynthetic pigments embedded directly in the membrane. It is an alternating pattern of dark and light bands measuring each 1 nanometre. The thylakoid lipid bilayer shares characteristic features with prokaryotic membranes and the inner chloroplast membrane. For example, acidic lipids can be found in thylakoid membranes, cyanobacteria and other photosynthetic bacteria and are involved in the functional integrity of the photosystems. The thylakoid membranes of higher plants are composed primarily of phospholipids and galactolipids that are asymmetrically arranged along and across the membranes. Thylakoid membranes are richer in galactolipids rather than phospholipids; also they predominantly consist of hexagonal phase II forming monogalacotosyl diglyceride lipid. Despite this unique composition, plant thylakoid membranes have been shown to assume largely lipid-bilayer dynamic organization. Lipids forming the thylakoid membranes, richest in high-fluidity linolenic acid are synthesized in a complex pathway involving exchange of lipid precursors between the endoplasmic reticulum and inner membrane of the plastid envelope and transported from the inner membrane to the thylakoids via vesicles. The thylakoid lumen is a continuous aqueous phase enclosed by the thylakoid membrane. It plays an important role for photophosphorylation during photosynthesis. During the light-dependent reaction, protons are pumped across the thylakoid membrane into the lumen making it acidic down to pH 4. In higher plants thylakoids are organized into a granum-stroma membrane assembly. A granum (plural grana) is a stack of thylakoid discs. Chloroplasts can have from 10 to 100 grana. Grana are connected by stroma thylakoids, also called intergranal thylakoids or lamellae. Grana thylakoids and stroma thylakoids can be distinguished by their different protein composition. Grana contribute to chloroplasts' large surface area to volume ratio. Different interpretations of electron tomography imaging of thylakoid membranes has resulted in two models for grana structure. Both posit that lamellae intersect grana stacks in parallel sheets, though whether these sheets intersect in planes perpendicular to the grana stack axis, or are arranged in a right-handed helix is debated. Chloroplasts develop from proplastids when seedlings emerge from the ground. Thylakoid formation requires light. In the plant embryo and in the absence of light, proplastids develop into etioplasts that contain semicrystalline membrane structures called prolamellar bodies. When exposed to light, these prolamellar bodies develop into thylakoids. This does not happen in seedlings grown in the dark, which undergo etiolation. An underexposure to light can cause the thylakoids to fail. This causes the chloroplasts to fail resulting in the death of the plant. Thylakoid formation requires the action of vesicle-inducing protein in plastids 1 (VIPP1). Plants cannot survive without this protein, and reduced VIPP1 levels lead to slower growth and paler plants with reduced ability to photosynthesize. VIPP1 appears to be required for basic thylakoid membrane formation, but not for the assembly of protein complexes of the thylakoid membrane. It is conserved in all organisms containing thylakoids, including cyanobacteria, green algae, such as Chlamydomonas, and higher plants, such as Arabidopsis thaliana.