Golgi Bodies:

 

Camillo Golgi discovered certain peculiar ultramicroscopic membranous structures in animal cells which showed complex organization. And there were named after him as Golgi complex or Golgi bodies or Golgi membranes. But biologists call such structures found in plant cells as dictyosomes.

Golgi bodies look like a stack of flat membranous sacs of various sizes and dimensions and form a complex. The organelles measure 2 mm to 5 mm in size and consists of 6-20 double membranes. They are found in large numbers particularly in secretory tissues like salivary glands, stigmatic surface cells, tapetal cells and glandular tissues of Drosera and other insectivorous plants. But other cells contain 2 -5 Golgi bodies.

Structures and Biogenesis:

Golgi bodies are specialized membranes designed to perform some specific functions. They are made up of a stack of flat circular or perforated membranous sacs containing lumen of 100 to 200 thickness. The number of membranous sacs vary from 10 to 20 with an inter spaces of 200 to 300 .

Such membrane complexes are associated with SER membranes at the formation face which is slightly convex in its topology called cis Golgi face. The other face is called maturation face Tran Golgi face. At the formation face smaller membranous sacs are is association with numerous vesicles derived from SER from various directions. In fact one can observe a transition from RER to SER and from SER to Golgi vesicles at the formation face. The vesicles thus derived from various SER membranes fuse with one another. So one finds the proteins that are synthesized on RER ultimately find their way into Golgi membranes at the proximal face. Even the lipids that are synthesized in SER and mono or disaccharides accumulated in SER find their way into Golgi membranes. Formation of vesicles from SER and fusion of them into a flat sac is a continuous process, thus many such sacs are stacked one above the other. However, various components like proteins, lipids and carbohydrates are further processed and packed at the peripheral region of membranous sacs which develop into enlarged bulbous structures. Once such packaging process is completed, the membranes pinch off vesicles and the membranes disappear at the maturation face. Thus one each observes various stages of processing and packaging of different components loaded into vesicles in the complex. The membrane flow from SER to Golgi, Golgi to SER and among the golgi membranes and from trans Golgi to other region is performed by a variety of transport proteins such as COPs , Clathrins and Adaptor and adaptin proteins.

Time lapse microphotography clearly shows that the total time required from the assembly of vesicles at the formation face and release of vesicles at the maturation face, is about 20-40 minutes. The formation of Golgi membranes at one face and disappearance of such membranes at the other is a continuous process, but the activity of golgi bodies is very high in certain specialized tissues where they are involved in secretion.

Chemical Composition:

Techniques like equilibrium density gradient centrifugation have come handy in isolating fairly pure Golgi membranes. The chemical analysis of such membrane components reveal that the Golgi membranes are made up of 60 per cent proteins and 40 per cent lipids. Some of the proteins found in Golgi bodies are same as that of endoplasmic reticulum. Nonetheless they differ from ER in other protein components, because the processed and packed proteins vary. But with respect to lipid components, particularly in plant Golgi membranes they have more of phosphotidic acids and phosphotidyl glycerol. Such Golgi membranes are specifically lacking in sialic acids which are predominant components in animal liver Golgi membranes. Plant Golgi membranes also contain carbohydrates like glucosamines, galactose, glucose, mannose, fucose, xylose, arabinose and other sugars. Furthermore NAD dependent enzymes, such as Cyt.C oxidase and Cyt B reductases are found in the membranes. C oxidase, NAD dependent enzymes and Cyt.B are found in Golgi membranes. The most characteristic rather marked enzymatic components are glycosyl transferase and thiamine pyrophosphotase. Acid phosphotase enzymes are also found because primary lysosomal structures are still in the process of formation and release.

Functions:

Various cellular components are loaded into SER, then transported and processed packed into Golgi membranes before they are released into cytosol as vesicles. The problem that still puzzles biologists is that, how different components are sorted out and pack them into asserted groups. One of the possible explanations is that as the incoming products are synthesized on a particular set of polysomal complex on RER the proteins should be same. Then they are processed and packed into a single vesicle. Different components, coming from different SER membranes in from different directions, are separate into different groups at the formation face of the Golgi complex itself; later these are processed and packed into different vesicles. The golgi vesicles do contain receptors for specific epitopes, thus each and every transit proteins are recognized and sorted out.

In plant cells, Golgi bodies process and pack enzymes and carbohydrates into the same vesicles or different vesicles for the synthesis of cell walls. Later such vesicles are transported towards plasma membranes, microtubules direct the transport. Lysosomal enzymes are also packed into different vesicles. Some of the components of micro bodies or glyoxysomes or peroxisomes, mitochondria, nucleus and plastids receive proteins from cytoplasm directly. It is also known that some of the mucilaginous products in plants are produced by Golgi complex.

Golgi membranes are also involved in the aggregation of many enzyme precursors into zymogen granules which get enclosed in the vesicles. Later they are transported to different regions where the precursor proteins are activated. But proteins like insulin in animal cells are processed into active molecules and packed into vesicles which are then released into outer spaces.

Furthermore the membranes of Golgi complexes are involved in lipid accumulation. Such lipid components are either glycosylated or and proteinated to produce glycolipids and lipoproteins respectively. In animal cells, during spermatid maturation, the required acrosomal components are produced by Golgi membranes. They are also responsible for packing a group of hydrolyzing enzymes into primary lysosomal structures.

An important feature of processing, sorting and packing of different components into different vesicles is the requirement of metabolic energy like ATP. Thus Golgi membranes perform functions like receiving, sorting, modifying, packaging and releasing various components into their individual functional but targeted vesicles, which ultimately reach specific destinations to perform their functions. Remarkable organelle indeed!

 

Inbound path: http://users.rcn.com/

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),

Out bound Path:

 

How does a vesicle recognize its correct target?

This involves pairs of complementary integral membrane proteins

v-SNAREs and t-SNAREs bind specifically to each other thanks to the complementary structure of their surface domains.

Binding is followed by fusion of the two membranes.

 

 

 

From SER vesicles containing proteins are budded off and the same join the cist Golgi membranes. There the proteins g through further modification if ne4cessary and as they go through they reach trans Golgi surface they are then budded off and transported to their respective destinations aided by Microtubules.

The trans-Golgi network (TGN) is a tubular network that originates from the last trans-Golgi cisternae. It sorts newly synthesized proteins that arrive from earlier Golgi compartments (I) towards different destinations (15). It also receives input from the endocytic pathway (IIIV) and sends back components to the earlier Golgi compartments (7). The exit routes from the TGN include those towards the apical plasma membrane (1), the basolateral plasma membrane (2), recycling endosomes (3), early endosomes (4), late endosomes (5) and specialized compartments such as secretory granules (6) in secretory cells. These are the main destinations, and for each of them more than one type of carrier might be involved; for example, basolateral transmembrane and soluble trafficking proteins can use different carriers101, 156. Apical transmembrane proteins and soluble, ciliary and GPI-linked proteins appear to use different pathways. Secretory proteins can also use a transendosomal pathway through recycling endosomes to reach the basolateral (3, 3a) or apical surfaces (3, 3b). Golgi-resident proteins (for example, glycosylating enzymes) recycle back to the Golgi stack , as secretion consumes the last trans-Golgi cisternae. Two important morphological features have emerged from the three-dimensional tomographic studies. First, the TGN is composed of tubules emanating from the last two trans-Golgi cisternae (which are smaller and appear to be detaching from the rest of the stack), although only the tubules deriving from the last cisterna are clathrin coated. Second, the endoplasmic reticulum makes close contact with the last two trans-Golgi cisternae (black boxes), through which lipid exchange can occur between these two organelles.; http://www.nature.com/

 

Two models vesicular model and cisternal maturation model; http://greatcourse.cnu.edu.cn/

Protein trafficking via membranes; http://web.bio.utk.edu/

www. micro.magnet.fsu.edu

 

 

jennarever.weebly.com

EM of the cell ( a part) showing Golgi membranes

http://sparkleberrysprings.com/

Putative interactions between Golgi stacks/vesicles and motor proteins; Specific subclasses of myosin XI would be responsible for the long-range transport of Golgi bodies within plant cells. Members of the kinesin superfamily are likely involved in either the anchorage of Golgi stacks to microtubules (such as kinesin-13A) or the short-range transport of Golgi vesicles to defined cell positions (such as AtPAKRP2 and p105). The arrow length indicates the relative velocity as produced by motor proteins. Opposing arrows indicate an anchorage role for the motor. http://journal.frontiersin.org/

Hypothetical model outlining the functions of kinesins during the assembly of cell wall. On the basis of current literature, members of the kinesin-4 subfamily (FRA1 and/or BC12) might be used to organize microtubules beneath the plasma membrane in order to favor either the proper insertion or the activation of cellulose synthase (CesA). The interplay between kinesins and CesA might be also more direct. Members of the kinesin-13 subfamily have been hypothesized either to transport locally or to pause Golgi stacks along microtubules. Once assembled in the Golgi stacks, CesA might move into the so-called SmaCC/MASC compartments, which are known to interact with microtubules, a step required for the insertion of CesA into the plasma membrane. The proteins mediating the interaction between SmaCC/MASC and microtubules are partially known (indicated by question mark) and FRA1/BC12 might putatively be part of such complex. As a further hypothesis, FRA1 and BC12 might be part of the complex that organize the nascent cellulose microfibrils at the plasma membrane interface by delivering specific components that regulate the orientation of cellulose microfibrils (indicated by X). http://journal.frontiersin.org/

 

For More refer: www.grkraj.org