Golgi Apparatus

Golgi apparatus
Golgi apparatus
external image Golgi.gif Many different enzymes (proteins) are present in the Golgi to perform its various synthetic activities. So there must be mechanisms

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Golgi Apparatus
Golgi Apparatus
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Golgi Apparatus

Golgi apparatus
Golgi apparatus
The Golgi apparatus is a membrane-bound structure with a single membrane. It is actually a stack of membrane-bound vesicles that are important in packaging macromolecules for transport elsewhere in the cell. The stack of larger vesicles is surrounded by numerous smaller vesicles containing those packaged macromolecules. The enzymatic or hormonal contents of lysosomes, peroxisomes and secretory vesicles are packaged in membrane-bound vesicles at the periphery of the Golgi apparatus.

-is an organelle
-Makes the vacuoles of a cell
-Also called the golgi body or the golgi complex
It is typically made up of 5 to 8 membrane colored sacs called the cisternae, but can have up to 60 in the golgi body.

The Golgi consists of a stack of membrane-bounded cisternae located between the endoplasmic reticulum and the cell surface.
Many different enzymes (proteins) are present in the Golgi to perform its various synthetic activities. So there must be mechanisms
  • to sort out the processed proteins and send them on to their destinations while
  • reclaiming processing proteins (e.g., glycosylases) for reuse.
All the details are far from worked out, but these are some of the features for which there is considerable experimental evidence.

The Outbound Path

  • Transition vesicles pinch off from the surface of the endoplasmic reticulum carrying
    • integral membrane proteins
    • soluble proteins awaiting processing
    • processing enzymes

  • Pinching off requires that the vesicle be coated with COPII (Coat Protein II)
  • The transition vesicles move toward the cis Golgi on microtubules.
  • As they do so, their COPII coat is removed and they may fuse together forming larger vesicles.
  • These fuse with the cis Golgi
  • Sugars are added to proteins in small packets so many glycoproteins have to undergo a large number of sequential steps of glycosylation, each requiring its own enzymes.
  • These steps take place as shuttle vesicles carry the proteins from cis to medial to the trans Golgi compartments.
  • At the outer face of the trans Golgi, vesicles pinch off and carry their completed products to their various destinations.

The Inbound Path

The movement of cisternal contents through the stack means that essential processing enzymes are also moving away from their proper site of action.
Using a variety of signals, the Golgi separates the products from the processing enzymes that made them and returns the enzymes back to the endoplasmic reticulum.
This transport is also done by pinching off vesicles, but the inbound vesicles are coated with COPI (coat protein I)

The Golgi apparatus (GA), also called Golgi body or Golgi complex and found universally in both plant and animal cells, is typically comprised of a series of five to eight cup-shaped, membrane-covered sacs called cisternae that look something like a stack of deflated balloons. In some unicellular flagellates, however, as many as 60 cisternae may combine to make up the Golgi apparatus. Similarly, the number of Golgi bodies in a cell varies according to its function. Animal cells generally contain between ten and twenty Golgi stacks per cell, which are linked into a single complex by tubular connections between cisternae. This complex is usually located close to the cell nucleus.
Due to its relatively large size, the Golgi apparatus was one of the first organelles ever observed. In 1897, an Italian physician named Camillo Golgi, who was investigating the nervous system by using a new staining technique he developed (and which is still sometimes used today; known as Golgi staining or Golgi impregnation), observed in a sample under his light microscope a cellular structure that he termed the internal reticular apparatus. Soon after he publicly announced his discovery in 1898, the structure was named after him, becoming universally known as the Golgi apparatus. Yet, many scientists did not believe that what Golgi observed was a real organelle present in the cell and instead argued that the apparent body was a visual distortion caused by staining. The invention of the electron microscope in the twentieth century finally confirmed that the Golgi apparatus is a cellular organelle.
The Golgi apparatus is often considered the distribution and shipping department for the cell's chemical products. It modifies proteins and lipids (fats) that have been built in the endoplasmic reticulum and prepares them for export outside of the cell or for transport to other locations in the cell. Proteins and lipids built in the smooth and rough endoplasmic reticulum bud off in tiny bubble-like vesicles that move through the cytoplasm until they reach the Golgi complex. The vesicles fuse with the Golgi membranes and release their internally stored molecules into the organelle. Once inside, the compounds are further processed by the Golgi apparatus, which adds molecules or chops tiny pieces off the ends. When completed, the product is extruded from the GA in a vesicle and directed to its final destination inside or outside the cell. The exported products are secretions of proteins or glycoproteins that are part of the cell's function in the organism. Other products are returned to the endoplasmic reticulum or may undergo maturation to become lysosomes.
The modifications to molecules that take place in the Golgi apparatus occur in an orderly fashion. Each Golgi stack has two distinct ends, or faces. The cis face of a Golgi stack is the end of the organelle where substances enter from the endoplasmic reticulum for processing, while the trans face is where they exit in the form of smaller detached vesicles. Consequently, the cis face is found near the endoplasmic reticulum, from whence most of the material it receives comes, and the trans face is positioned near the plasma membrane of the cell, to where many of the substances it modifies are shipped. The chemical make-up of each face is different and the enzymes contained in the lumens (inner open spaces) of the cisternae between the faces are distinctive. Illustrated in Figure 2 is a fluorescence digital image taken through a microscope of the Golgi apparatus (pseudocolored red) in a typical animal cell. Note the close proximity of the Golgi membranes to the cell nucleus.
Proteins, carbohydrates, phospholipids, and other molecules formed in the endoplasmic reticulum are transported to the Golgi apparatus to be biochemically modified during their transition from the cis to the trans poles of the complex. Enzymes present in the Golgi lumen modify the carbohydrate (or sugar) portion of glycoproteins by adding or subtracting individual sugar monomers. In addition, the Golgi apparatus manufactures a variety of macromolecules on its own, including a variety of polysaccharides. The Golgi complex in plant cells produces pectins and other polysaccharides specifically needed by for plant structure and metabolism. The products exported by the Golgi apparatus through the trans face eventually fuse with the plasma membrane of the cell. Among the most important duties of the Golgi apparatus is to sort the wide variety of macromolecules produced by the cell and target them for distribution to their proper location. Specialized molecular identification labels or tags, such as phosphate groups, are added by the Golgi enzymes to aid in this sorting effort.

animal cell
animal cell

Golgi bodies (also apparatus or complex) store and package cellular secretions for export out of the cell (usually through the use of vacuoles). Salivary, oil, and digestive glands have very active golgi bodies.

The Golgi apparatus (GA), also called Golgi body or Golgi complex and found universally in both plant and animal cells, is typically comprised of a series of five to eight cup-shaped, membrane-covered sacs called cisternae that look something like a stack of deflated balloons. In some unicellular flagellates, however, as many as 60 cisternae may combine to make up the Golgi apparatus. Similarly, the number of Golgi bodies in a cell varies according to its function. Animal cells generally contain between ten and twenty Golgi stacks per cell, which are linked into a single complex by tubular connections between cisternae. This complex is usually located close to the cell nucleus.

The Golgi apparatus
The Golgi apparatus is a cell structure mainly devoted to processing the proteins synthesized in the endoplasmic reticulum (ER).
Link to discussion of the paths taken by proteins when they leave the Golgi.
  • The major processing activity is glycosylation: the adding of sugar molecules to form glycoproteins.
  • In some cells, e.g., mucus-secreting cells in epithelia, the amount of carbohydrate so far exceeds that of the protein that the product is called a mucopolysaccharide (also known as a proteoglycan).
  • In plant cells, the Golgi secretes the cell plate and cell wall.
  • Small peptides, e.g., some hormones and neurotransmitters, are too small to be synthesized directly by ribosomes. Instead, the ribosomes on the ER synthesize a large precursor protein that is later cut up into small peptide fragments as it traverses the Golgi.

external image golgi_apparatus.gifexternal image golgi.gif

Foundation of Vesicles
The Golgi complex gathers simple molecules and combines them to make molecules that are more complex. It then takes those big molecules, packages them in vesicles, and either stores them for later use or sends them out of the cell. It is also the organelle that builds lysosomes (cell digestion machines). Golgi complexes in the plant may also create complex sugars and send them off in secretory vesicles. The vesicles are created in the same way the ER does it. The vesicles are pinched off the membranes and float through the cell.

The Golgi complex is a series of membranes shaped like pancakes. The single membrane is similar to the cell membrane in that it has two layers. The membrane surrounds an area of fluid where the complex molecules (proteins, sugars, enzymes) are stored and changed. Because the Golgi complex absorbs vesicles from the rough ER, you will also find ribosomes in those pancake stacks.

Working with the Rough ER

Process of Golgi forming vesicles
Process of Golgi forming vesicles

The Golgi apparatus (also called the Golgi body, Golgi complex, or dictyosome) is an organelle found in most eukaryotic cells. It was identified in 1898 by the Italian physician Camillo Golgi and was named after him. The primary function of the Golgi apparatus is to process and package the macromolecules such as proteins and lipids that are synthesized by the cell. It is particularly important in the processing of proteins for secretion. The Golgi apparatus forms a part of the endomembrane system of eukaryotic cells.

The Golgi is composed of membrane-bound sacs known as cisternae. Between five and eight are usually present; however, as many as sixty have been observed.[1]

The cisternae stack has five functional regions: the cis-Golgi network, cis-Golgi, medial-Golgi, trans-Golgi, and trans-Golgi network. Vesicles from the endoplasmic reticulum (via the vesicular-tubular cluster) fuse with the cis-Golgi network and subsequently progress through the stack to the trans-Golgi network, where they are packaged and sent to the required destination. Each region contains different enzymes which selectively modify the contents depending on where they are destined to reside.[2].

Cells synthesize a large number of different macromolecules required for life. The Golgi apparatus is integral in modifying, sorting, and packaging these substances for cell secretion (exocytosis) or for use within the cell. It primarily modifies proteins delivered from the rough endoplasmic reticulum, but is also involved in the transport of lipids around the cell, and the creation of lysosomes. In this respect it can be thought of as similar to a post office; it packages and labels items and then sends them to different parts of the cell.

Enzymes within the cisternae are able to modify substances by the addition of carbohydrates (glycosylation) and phosphate (phosphorylation) to them. In order to do so the Golgi transports substances such as nucleotide sugars into the organelle from the cytosol. Proteins are also labelled with a signal sequence of molecules which determine their final destination. For example, the Golgi apparatus adds a mannose-6-phosphate label to proteins destined for lysosomes.

The Golgi also plays an important role in the synthesis of proteoglycans, molecules present in the extracellular matrix of animals, and it is a major site of carbohydrate synthesis.[3]

This includes the productions of glycosaminoglycans or GAGs, long unbranched polysaccharides which the Golgi then attaches to a protein synthesized in the endoplasmic reticulum to form the proteoglycan.[4]Enzymes in the Golgi will polymerize several of these GAGs via a xylose link onto the core protein. Another task of the Golgi involves the sulfation of certain molecules passing through its lumen via sulphotranferases that gain their sulphur molecule from a donor called PAPs. This process occurs on the GAGs of proteoglycans as well as on the core protein. The level of sulfation is very important to the proteoglycans' signalling abilities as well as giving the proteoglycan its overall negative charge.[3]

The Golgi is also capable of phosphorylating molecules. To do so it transports ATP into the lumen.[5] The Golgi itself contains resident kinases, such as casein kinases. One molecule that is phosphorylated in the Golgi is Apolipoprotein, which forms a molecule known as VLDL that is a constitute of blood serum. It is thought that the phosphorylation of these molecules is important to help aid in their sorting of secretion into the blood serum.[6]

The Golgi also has a putative role in apoptosis, with several Bcl-2 family members localised there, as well as to the mitochondria. In addition a newly characterised anti-apoptotic protein, GAAP (Golgi anti-apoptotic protein), which almost exclusively resides in the Golgi, protects cells from apoptosis by an as-yet undefined mechanism (Gubser et al., 2007).

Vesicles which leave the rough endoplasmic reticulum are transported to the cis face of the Golgi apparatus, where they fuse with the Golgi membrane and empty their contents into the lumen. Once inside they are modified, sorted, and shipped towards their final destination. As such, the Golgi apparatus tends to be more prominent and numerous in cells synthesising and secreting many substances: plasma B cells, the antibody-secreting cells of the immune system, have prominent Golgi complexes.

Those proteins destined for areas of the cell other than either the endoplasmic reticulum or Golgi apparatus are moved towards the trans face, to a complex network of membranes and associated vesicles known as the trans-Golgi network (TGN).[2] This area of the Golgi is the point at which proteins are sorted and shipped to their intended destinations by their placement into one of at least three different types of vesicles, depending upon the molecular marker they carry:[2]

Exocytotic vesicles (continuous) Vesicle contains proteins destined for extracellular release. After packaging the vesicles bud off and immediately move towards the plasma membrane, where they fuse and release the contents into the extracellular space in a process known as constitutive secretion. Antibody release by activated plasma B cells
Secretory vesicles (regulated) Vesicle contains proteins destined for extracellular release. After packaging the vesicles bud off and are stored in the cell until a signal is given for their release. When the appropriate signal is received they move towards the membrane and fuse to release their contents. This process is known as regulated secretion. Neurotransmitter release from neurons
Lysosomal vesicles Vesicle contains proteins destined for the lysosome, an organelle of degradation containing many acid hydrolases, or to lysosome-like storage organelles. These proteins include both digestive enzymes and membrane proteins. The vesicle first fuses with the late endosome, and the contents are then transferred to the lysosome via unknown mechanisms. Digestive proteases destined for the lysosome

The transport mechanism which proteins use to progress through the Golgi apparatus is not yet clear; however a number of hypotheses currently exist. Until recently, the vesicular transport mechanism was favoured but now more evidence is coming to light to support cisternal maturation. The two proposed models may actually work in conjunction with each other, rather than being mutually exclusive. This is sometimes referred to as the combined model. [3]

Cisternal maturation model: the cisternae of the Golgi apparatus move by being built at the cis face and destroyed at the trans face. Vesicles from the endoplasmic reticulum fuse with each other to form a cisterna at the cis face, consequently this cisterna would appear to move through the Golgi stack when a new cisterna is formed at the cis face. This model is supported by the fact that structures larger than the transport vesicles, such as collagen rods, were observed microscopically to progress through the Golgi apparatus.[3] This was initially a popular hypothesis, but lost favour in the 1980s. Recently it has made a comeback, as laboratories at the University of Chicago and the University of Tokyo have been able to use new technology to directly observe Golgi compartments maturing.[7]. Additional evidence comes from the fact that COP1 vesicles move in the retrograde direction,. transporting ER proteins back to where they belong by recognizing a signal peptide.[8]
Vesicular transport model: Vesicular transport views the Golgi as a very stable organelle, divided into compartments is the cis to trans direction. Membrane bound carriers transported material between the ER and Golgi and the different compartments of the Golgi.[9] Experimental evidence includes the abundance of small vesicles (known technically as shuttle vesicles) in proximity to the Golgi apparatus. Directionality is achieved by packaging proteins into either forward-moving or backward-moving (retrograde) transport vesicles, or alternatively this directionality may not be necessary as the constant input of proteins from the endoplasmic reticulum on the cis face of the Golgi would ensure flow. Irrespectively, it is likely that the transport vesicles are connected to a membrane via actin filaments to ensure that they fuse with the correct compartment.[3]

The Golgi apparatus is membranous organelle that appears under an electron microscope as a multi-layered stack of flattened sacs. These sacks are referred to as cisternae. Cisternae are not connected, but remain in very close proximity. Their function is to receive and synthesize molecules within the cell. They also prepare molecules for exiting the cell.
    1. Nuclear membrane
    2. Nuclear pore
    3. Rough endoplasmic reticulum (rER)
    4. Smooth endoplasmic reticulum (sER)
    5. Ribosome attached to rER
    6. Macromolecules
    7. Transport vesicles
    8. Golgi apparatus
    9. Cis face of Golgi apparatus
    10. Trans face of Golgi apparatus
    11. Cisternae of Golgi apparatus
Image:Nucleus ER golgi.jpg
Image:Nucleus ER golgi.jpg