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Figure 2-12

Digestion of substances in pinocytotic or phagocytic vesicles by enzymes derived from lysosomes.

occurs in the uterus after pregnancy, in muscles during long periods of inactivity, and in mammary glands at the end of lactation. Lysosomes are responsible for much of this regression. The mechanism by which lack of activity in a tissue causes the lysosomes to increase their activity is unknown.

Another special role of the lysosomes is removal of damaged cells or damaged portions of cells from tissues. Damage to the cell—caused by heat, cold, trauma, chemicals, or any other factor—induces lyso-somes to rupture. The released hydrolases immediately begin to digest the surrounding organic substances. If the damage is slight, only a portion of the cell is removed, followed by repair of the cell. If the damage is severe, the entire cell is digested, a process called autolysis. In this way, the cell is completely removed, and a new cell of the same type ordinarily is formed by mitotic reproduction of an adjacent cell to take the place of the old one.

The lysosomes also contain bactericidal agents that can kill phagocytized bacteria before they can cause cellular damage. These agents include (1) lysozyme, which dissolves the bacterial cell membrane; (2) lyso-ferrin, which binds iron and other substances before they can promote bacterial growth; and (3) acid at a pH of about 5.0, which activates the hydrolases and inactivates bacterial metabolic systems.

Synthesis and Formation of Cellular Structures by Endoplasmic Reticulum and Golgi Apparatus

Specific Functions of the Endoplasmic Reticulum

The extensiveness of the endoplasmic reticulum and the Golgi apparatus in secretory cells has already been emphasized. These structures are formed primarily of lipid bilayer membranes similar to the cell membrane, and their walls are loaded with protein enzymes that catalyze the synthesis of many substances required by the cell.

Most synthesis begins in the endoplasmic reticulum. The products formed there are then passed on to the Golgi apparatus, where they are further processed before being released into the cytoplasm. But first, let us note the specific products that are synthesized in specific portions of the endoplasmic reticulum and the Golgi apparatus.

Proteins Are Formed by the Granular Endoplasmic Reticulum.

The granular portion of the endoplasmic reticulum is characterized by large numbers of ribosomes attached to the outer surfaces of the endoplasmic reticulum membrane. As we discuss in Chapter 3, protein molecules are synthesized within the structures of the ribo-somes. The ribosomes extrude some of the synthesized protein molecules directly into the cytosol, but they also extrude many more through the wall of the endo-plasmic reticulum to the interior of the endoplasmic vesicles and tubules, that is, into the endoplasmic matrix.

- Pinocytotic or phagocytic vesicle

- Digestive vesicle

■ Residual body

Synthesis of Lipids by the Smooth Endoplasmic Reticulum.

The endoplasmic reticulum also synthesizes lipids, especially phospholipids and cholesterol. These are rapidly incorporated into the lipid bilayer of the endo-plasmic reticulum itself, thus causing the endoplasmic reticulum to grow more extensive. This occurs mainly in the smooth portion of the endoplasmic reticulum.

To keep the endoplasmic reticulum from growing beyond the needs of the cell, small vesicles called ER vesicles or transport vesicles continually break away from the smooth reticulum; most of these vesicles then migrate rapidly to the Golgi apparatus.

Other Functions of the Endoplasmic Reticulum. Other significant functions of the endoplasmic reticulum, especially the smooth reticulum, include the following:

1. It provides the enzymes that control glycogen breakdown when glycogen is to be used for energy.

2. It provides a vast number of enzymes that are capable of detoxifying substances, such as drugs, that might damage the cell. It achieves detoxification by coagulation, oxidation, hydrolysis, conjugation with glycuronic acid, and in other ways.

Specific Functions of the Golgi Apparatus

Synthetic Functions of the Golgi Apparatus. Although the major function of the Golgi apparatus is to provide additional processing of substances already formed in the endoplasmic reticulum, it also has the capability of synthesizing certain carbohydrates that cannot be formed in the endoplasmic reticulum. This is especially true for the formation of large saccharide polymers bound with small amounts of protein; the most important of these are hyaluronic acid and chondroitin sulfate.

A few of the many functions of hyaluronic acid and chondroitin sulfate in the body are as follows: (1) they are the major components of proteoglycans secreted in mucus and other glandular secretions; (2) they are the major components of the ground substance outside the cells in the interstitial spaces, acting as filler between collagen fibers and cells; and (3) they are principal components of the organic matrix in both cartilage and bone.

Processing of Endoplasmic Secretions by the Golgi Apparatus— Formation of Vesicles. Figure 2-13 summarizes the major functions of the endoplasmic reticulum and Golgi apparatus. As substances are formed in the endoplasmic reticulum, especially the proteins, they are transported through the tubules toward portions of the smooth endoplasmic reticulum that lie nearest the Golgi apparatus. At this point, small transport vesicles composed of small envelopes of smooth endo-plasmic reticulum continually break away and diffuse to the deepest layer of the Golgi apparatus. Inside these vesicles are the synthesized proteins and other products from the endoplasmic reticulum.

The transport vesicles instantly fuse with the Golgi apparatus and empty their contained substances into the vesicular spaces of the Golgi apparatus. Here,

Protein Lipid Secretory

Ribosomes formation formation Lysosomes vesicles

Protein Lipid Secretory

Ribosomes formation formation Lysosomes vesicles

endoplasmic endoplasmic apparatus reticulum reticulum

Figure 2-13

endoplasmic endoplasmic apparatus reticulum reticulum

Figure 2-13

Formation of proteins, lipids, and cellular vesicles by the endoplasmic reticulum and Golgi apparatus.

additional carbohydrate moieties are added to the secretions. Also, an important function of the Golgi apparatus is to compact the endoplasmic reticular secretions into highly concentrated packets. As the secretions pass toward the outermost layers of the Golgi apparatus, the compaction and processing proceed. Finally, both small and large vesicles continually break away from the Golgi apparatus, carrying with them the compacted secretory substances, and in turn, the vesicles diffuse throughout the cell.

To give an idea of the timing of these processes: When a glandular cell is bathed in radioactive amino acids, newly formed radioactive protein molecules can be detected in the granular endoplasmic reticulum within 3 to 5 minutes.Within 20 minutes, newly formed proteins are already present in the Golgi apparatus, and within 1 to 2 hours, radioactive proteins are secreted from the surface of the cell.

Types of Vesicles Formed by the Golgi Apparatus—Secretory Vesicles and Lysosomes. In a highly secretory cell, the vesicles formed by the Golgi apparatus are mainly secretory vesicles containing protein substances that are to be secreted through the surface of the cell membrane. These secretory vesicles first diffuse to the cell membrane, then fuse with it and empty their substances to the exterior by the mechanism called exo-cytosis. Exocytosis, in most cases, is stimulated by the entry of calcium ions into the cell; calcium ions interact with the vesicular membrane in some way that is not understood and cause its fusion with the cell membrane, followed by exocytosis—that is, opening of the membrane's outer surface and extrusion of its contents outside the cell.

Some vesicles, however, are destined for intracellu-lar use.

Use of Intracellular Vesicles to Replenish Cellular Membranes.

Some of the intracellular vesicles formed by the Golgi apparatus fuse with the cell membrane or with the membranes of intracellular structures such as the mitochondria and even the endoplasmic reticulum. This increases the expanse of these membranes and thereby replenishes the membranes as they are used up. For instance, the cell membrane loses much of its substance every time it forms a phagocytic or pinocy-totic vesicle, and the vesicular membranes of the Golgi apparatus continually replenish the cell membrane.

In summary, the membranous system of the endo-plasmic reticulum and Golgi apparatus represents a highly metabolic organ capable of forming new intra-cellular structures as well as secretory substances to be extruded from the cell.

Extraction of Energy from Nutrients— Function of the Mitochondria

The principal substances from which cells extract energy are foodstuffs that react chemically with oxygen—carbohydrates, fats, and proteins. In the human body, essentially all carbohydrates are converted into glucose by the digestive tract and liver before they reach the other cells of the body. Similarly, proteins are converted into amino acids and fats into fatty acids. Figure 2-14 shows oxygen and the foodstuffs—glucose, fatty acids, and amino acids—all entering the cell. Inside the cell, the foodstuffs react chemically with oxygen, under the influence of enzymes that control the reactions and channel the energy released in the proper direction. The details of

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