~ 1 ~ Shanes notes on Cell Biology and Biomolecules
CELL ORGANELLES Microbodies: Lysosomes, Peroxisomes and glyoxisomes The term microbody was used by cell biologists and cytologists for many years to describe a variety of different small cellular components. More recently, it has usually been restricted to organelles possessing flavin oxidases and catalase. Microbodies are a variety of small, ovaoid, rounded or spherical subcellular organelles having oxidase, peroxidase or catalase enzymes, found scattered in the cytoplasm. These were first described by J. Rhodin (1954) in the cells of mouse kidney. Organelles possessing these activities are typically spherical or ovoid structures having a single bounding membrane, a diameter of about 0.5-1.5 m, and containing an amorphous gran- ular matrix, occasionally with crystalloid inclusions. The organelles vary somewhat in structure, appearance, and function from one tissue to another and from species to species. Certain micro- bodies exhibit specific biochemical characteristics as well as specific distributions among animal, plant, and microbial cells. Included here are the peroxisomes and glyoxysomes.
LYSOSOMES (G. lysis: dissolution; soma: body) (The Suicide Bags of Cell) Introduction: Lysosomes are membranous bag-like cell organelles containing hydrolytic enzymes whose main function is the controlled intracellular digestion of macromolecules and degrade these polymers into their monomeric subunits. History: Lysosomes were discovered accidently by Nobel laureates Christian DeDuve (1955) in the liver cells because of their acid phosphate activity. These were named lysosomes because these contained digestive or tissue dissolving enzymes. Novikoff examined these acid-phosphatase containing subcellular fractions under electron microscope about a decade after they had been first described. Occurrence: Lysosomes are present in all animal cells but only in a few plant cells. These are most abundant in macrophages which perform digestive functions and in the cells of those organs which are associated with enzymatic actions such as liver, pancreas, thyroid, spleen, intestine and kidney. In plants, these are found in the meristematic cells.
Table: cells and tissues containing lysosomes
Cell organelles
~ 2 ~ Shanes notes on Cell Biology and Biomolecules Structure: Lysosomes are a structurally heterogeneous group of organelles and vary dramatically in size and morphology. As a result, it is difficult to identify lysosomes strictly on the basis of morphological criteria. When lysosome- rich fractions were initially isolated centrifugally by de Duve and Novikoff and examined with the electron microscope, it was found that the suspected lysosomes were generally about the same size as small mitochondria. The heterogenicity of lysosomal morphology reflects the wide variety of digestive funtionsi.e., digestion of extra and intracellular debris and the digestion of phagocytosed microorganisms. Typically, they varied in diameter from about 0.04 to 0.8 m, were bounded by a single membrane and enclose a dense solid matrix. Identification of lysosomes in sections of whole cells is considerably more difficult because other small, dense organelles are also bounded by a single membrane. The matrix may appear to be almost solid and differentiated into very dense outer layer one and a less dense central mass. The central mass may enclose cavities or vacuoles with granular substance which enclose large amount of hydrolytic enzymes.
Types of lysosomes or polymorphism of lysosomes They are of four types: 1. Primary lysosome (storage granules): It is a small sac-like body whose enzymatic contents are synthesized by ribosomes and accumulated in ER. From there, they enter the Golgi region, where acid phosphatase reaction takes place. The GERL region, i.e., acid phosphatase rich region of Golgi maturing face is thought to be involved in the production of lysosomes. The primary lysosome comprises only one type of enzyme or another. 2. Secondary lysosomes (digestive vacuole or heterophagosome): These are produced either from phagocytosis or pinocytosis of foreign material by the cell. Actually within the cell the foreign bodies or extra-cellular substances are enclosed within Cell organelles
~ 3 ~ Shanes notes on Cell Biology and Biomolecules the membrane after phagocytosis or pinocytosis and these membrane bound structures are known as phagosomes or pinosomes. These ultimately fuse with primary lysosomes, thus forming secondary lysosomes. This body having engulfed material within membrane has also full complements of acid hydrolases (hydrolytic enzymes). The digested material of these lysosomes passes through the lysosomal membrane and are incorporated into the cell so that they may be reused in metabolic pathways.
3. Residual bodies: These are formed if the digestion is incomplete. In some cells, such as the Amoeba and other protozoa, these residual bodies are eliminated by defecation. Hence lysosomes having undigested material or debris are called residual bodies. These bodies are formed due to lack of certain enzymes in lysosomes. These are rejected from the cell by exocytosis and some time in certain cells these bodies remain in cells for long time causing ageing. These residual bodies also cause diseases in man such as fever, hepatitis, polynephritis, hypertension, congested heart failure, etc. If the debris, mostly lipid in nature, may accumulate and condense into concentric lamellae, it forms myelin fibres. 4. Autophagic vacuole (cytolysosome or autophagosome): It is a special case in which lysosome digests a part of cell (e.g., mitochondria or portions of ER) by the process of autophagy. For example, liver cell shows numerous autophagosomes during starvation among which remnants of mitochondria occur. This is a mechanism by which the cell can achieve degradation of its own constituents without irreparable damage.
Cell organelles
~ 4 ~ Shanes notes on Cell Biology and Biomolecules
Functions of lysosomes: Some of the main functions of Lysosomes are as follows: 1. Intracellular digestion: The word lysosome is derived from (lyso/ lytic or digestive; and soma body) thus helping in digestion. Pinocytic vacuoles formed as a result of absorption of fluid substance into cell or phagocytic vacuoles formed by absorption of solid particles into cell carry protein material to lysosomal region. These foreign proteins may undergo digestion within cell as a result of endocytosis. Endocytosis includes the processes of phagocytosis (Gr., phagein, to eat), pinocytosis (Gr., pinein, to drink) and micropinocytosis. Phagocytosis and pinocytosis are active mechanisms in which cell requires energy for their operation. In endocytosis contraction of microfilaments of actin and myosin present in the peripheral cytoplasm occurs. It causes plasma membrane to invaginate and form the endocytic vacuole. Ingested particles enclosed in membranes derived from the plasma membrane and forming vacuoles are sometimes celled phagosomes. After the entrance of a particle or large body into the cell by endocytosis and the formation of a phagosome, the membranes of the phagosome and a lysosome may fuse to form a single large vacuole. Within this vacuole the lysosomal enzymes begin the process of digestion of the foreign material. Initially the lysosome, known as a primary lysosome containing the complex of enzymes in an inactive state, but after fusion with the phagosome produces a secondary lysosome with a different morphology and active enzymes. After enzymatic digestion the digested material diffuses out into the hyaloplasm of the cell. Some material may remain in the enlarged lysosome vacuole. This remnant vacuole is the residual body, since it contains the residue of digestive process. During starvation also, lysosomes digest stored food materials, i.e., proteins, lipids and glycogen of cytoplasm and provide energy required by the cell. Digestion of proteins usually ends at the level of dipeptide, which can pass through the membrane and be further digested into amino acids. Cell organelles
~ 5 ~ Shanes notes on Cell Biology and Biomolecules
Carbohydrates are usually hydrolysed to monosaccharides which are easily released. However, disaccharides and polysaccharides (cellobiose, inulin or dextrin) are not digested and remain within the lysosome. Digestion of intracellular substances or autophagy: Many cellular components, such as mitochondria, are constantly being removed from the cell by lysosomal system. Cytoplasmic organelles become surrounded by membranes of smooth endoplasmic reticulum, forming vacuoles, then lysosomal enzymes are discharged into autophagic vacuoles and the organelles are digested. Autophagy is a general property of eukaryotic cells. It is related to the normal renovation and turnover of cellular components. The digestion of mitochondria or other cell structures, such as elements of ER, provides a source of energy for these cells. After the digestion of cell structure, the autophagic vacuoles may become residual bodies. 2. Removal of dead cells: Hirsch and Cohn (1964) told that lysosomes help in the removal of dead cells in tissues such as white blood cells with engulfed bacterium in blood, cells in the outer layer of skin and mucous membrane linings of the body. Lysosomal membrane ruptures in these cells, releasing enzymes into body of cell, so that whole cell may be digested. Lysosomes contain a sufficient complement of enzymes to digest most types of biological or organic materials and the digestive process (autolysis) occurs quite rapidly in dead cells. This process of tissue degeneration (necrosis) is due to this lysosomal activity. 3. Role in metamorphosis: Recently lysosomes role has been discovered in the metamorphosis of frog. Disappearance of tail from tadpole larva of frog is due to lysosomal activity (action of cathepsins present in lysosomes) as described by Weber. 4. Help in protein synthesis: Novikoff and Essner (1960) have suggested the possible role of lysosomes in protein synthesis. Recently, the author (Dr. Singh 1972), has correlated lysosomal activity with the protein synthesis. In liver and pancreas of some birds, lysosomes seem to be more active and developed as reported by Singh (1972), showing possible relationship with cell metabolism. 5. Help in fertilization: During fertilization, sperms head secrete some lysosomal enzymes which help in the penetration of sperm into vitelline layer of ovum. Acrosome contains protease and hyaluronidase and abundant acid phosphatase. Hyaluronidase disperse the cells around the oocyte and protease digests the zona pellucida making a channel through which sperm nucleus penetrates. Cell organelles
~ 6 ~ Shanes notes on Cell Biology and Biomolecules 6. Role in osteogenesis: It has been argued that formation of bone cells and also their destruction depends upon lysosomal activity. Likewise, ageing of cells and parthenogenetic development are related with the lysosome activity. The osteoclasts (multinucleated cells) which remove bone, do so by the release of lysosomal enzymes which degrade the organic matrix. This process is activated by the parathyroid hormone. 7. Malfunctioning of lysosomes: Lysosomal malfunction may lead to diseases, for example, when glycogen taken up by lysosomes is not digested (Pompes disease). Ruptures of lysosomes in skin cells exposed to direct sun light leads to pathological changes following sunburn. The enzymes liberated by these lysosomes kill cells in the epidermis, causing blistering and later to peeling of a layer of epidermis. A metabolic disorder, the gout, is caused by the accumulation of sodium urate crystals in the joints. These are picked up by the phagocytes resulting in their lysosomes rupture. This leads to the acute inflammation and increased collagen synthesis. Malfunctioning of lysosomes (either hyper or hypo activity) can cause several disease. Some of these are inborn lysosomal diseases such as Hunters syndrome (mucopolysaccharide disorder), Krabbes disease (abnormal sphingolipidosis), mannosidosis (disorder of glycoproteinmetabolism) and cystinosis (accumulation of cystine) etc. 8. Autolysis in cartilage and bone tissue: The excess of vitamin A causes cell poisoning. It disrupts the lysosomal membrane, causing release of enzymes into the cell and producing autolysis in cartilage and bone tissue. 9. Seed germination in plants: During seed germination, lysosomal enzymes are released by the seedlings which attack stored food material making it available to the developing plant. 10. Chromosomal breaks: Lysosomes contain enzyme deoxyribonuclease (DNAase). This enzyme causes chromosomal breaks and their rearrangement. DNAase has two active sites and breaks down both the strands of DNA. This breaks leads to various genetic disorders.
PEROXISOMES In 1965, de Duve showed that microbodies of rat liver contained a number of oxidases that transfer hydrogen atoms to molecular oxygen, thereby forming hydrogen peroxide (Fig 1). De Duve coined the term peroxisomes for these organelles, although a true peroxidatic activity is generally demonstrable only in vitro. In vivo, conditions favor the removal (or degradation) of hydrogen peroxide by catalase rather than by a peroxidase. However, because hydrogen peroxide is an intermediate in the reaction, the term peroxisome may be appropriate. The chemical and enzymatic relationships between an oxidase, peroxidase, and catalase are shown in Figure 1. Cell organelles
~ 7 ~ Shanes notes on Cell Biology and Biomolecules
Fig 1: chemical interrelationship between oxidase, peroxidase and catalase enzymes A number of enzymes are characteristically present in peroxisomes, including uric acid oxidase (also called uricase), D-amino acid oxidase, acyl-CoA oxidase, polyamine oxidase, - hyroxyacid oxidase, NADH-glyoxylate reductase, NADP-isocitrate dehydrogenase, and catalase. When uric acid oxidase is present in large amounts, it frequently takes the form of a paracrystalline nucleoid at the center of the organelle. The functions of peroxisomes in animal cells are diverse. Peroxisomal catalase is thought to be involved in the degradation of H 2 O 2 , which is extremely toxic, the source of the peroxide being other peroxisomal reactions (e.g., those catalyzed by the flavin oxidases). Uric acid oxidase is important in the catabolic pathway that degrades purines. The early observation that there is an abundance of peroxisomes in cells en- gaged in lipid metabolism suggested that these organelles may be involved in lipid metabolism. Recently; it has been shown that liver peroxisomes contain a major system for the -oxidation of fatty acids however; the enzymes are different from those of mitochondria although they produce the same end product, acetyl-CoA. Electron-microscopic studies of tissue sections often reveal a close proximity between peroxisomes and mitochondria. This is not surprising as the products of peroxisomal activity may serve as substrates for mitochondrial activity. For example, glyoxylate produced in peroxisomes may be converted there to glycine by transamination. After passage to neighboring mito- chondria, the glycine may be further metabolized in a variety of ways including conversion to other amino acids or incorporation into heme. One of the characteristic features of lysosomal enzyme activity is its latency. No latency is exhibited by the peroxisomal enzymes, as relatively large molecules (including the peroxisomal enzyme substrates) readily permeate the peroxisome membrane. Isolation of Peroxisomes: The sedimentation coefficients and densities of peroxisomes in sucrose gradients are close to those of lysosomes and account for the fact that for some time peroxisomal enzyme activities were ascribed to lysosomes. Density gradient centrifugation of the so- called light mitochondrial fraction prepared by preliminary differential centrifugation is the method of choice for isolating peroxiso- mes. The greatest success in peroxisome purification is obtained if the lysosomes are first allowed to accumulate Triton WR-1339. Cell organelles
~ 8 ~ Shanes notes on Cell Biology and Biomolecules Triton-loaded lysosomes are considerably less dense than normal lysosomes and so they are easily floated away from the peroxisomes during sucrose. Formation of Peroxisomes: For many years, it was believed that peroxisomes of both plant and animal tissues arose as outgrowths of the endoplasmic reticulum and that the peroxisomal enzymes were dispatched into the cisternae of the endoplasmic reticulum by attached ribosomes. The enzymes made their way into the organelle prior to its physical separation from the ER. However, the growing tide of recent evidence appears to argue in favor of an alternative model for the biogenesis of peroxisomes. In liver and in other tissues as well, many of the peroxisomal enzymes are synthesized by unattached ribosomes (i.e., ribosomes that are not bound to the ER) and are released into the cytosol. From there, the enzymes are slowly taken up by preexisting peroxisomes. Studies with peroxisomal catalase indicate that the subunits of the enzyme and the heme enter the microbody and are then assembled to form the functional enzyme. The nature of the mechanism that translocates peroxisomal proteins into peroxisomes is uncertain, but it is speculated that the proteins are first bound to a membrane receptor and are then drawn through the membrane and trapped inside as multi-protein complexes. Indeed, dense crystalline aggregates are often ob- served inside peroxisomes. Translocation of proteins through the peroxisome membrane is readily demonstrable in vitro. When newly synthesized peroxisomal proteins are tagged with a radioactive label and incu- bated with isolated peroxisomes, some of the protein may soon be recovered from within the organelles. Peroxisomes often appear in clusters when examined by electron microscopy and occasional dumbbell- shaped images are observed. This suggests that the tails seen on peroxisomes are connections to other peroxisomes. Indeed, either permanent or transient connections between peroxisomes might form a peroxisome reticulum. Such interconnections would explain the remarkable biochemical homogeneity of peroxisomes and the synchronous turnover of peroxisomal proteins. New peroxisomes may be formed either by the fission of preexisting peroxisomes or by budding from the peroxisome reticulum. Functions of peroxisomes: In green plant peroxisomes carry out photorespiration. The oxidase of peroxisomes transfers hydrogen atoms to molecular oxygen, thereby forming hydrogen peroxide. The hydrogen peroxide being extremely toxic is degraded into water and oxygen to peroxisomal catalase or peroxidase.
GLYOXYSOMES Glyoxysomes are generally found in the cells of endosperm only. These are similar to peroxisomes but enzymes of glyoxylate cycle along with peroxisomal enzymes (enzymes of oxidative cycle). These were described by R. W. Brieden-Bach in 1967. These are associated with triglyceride metabolism. These transform stored fat into carbohydrates by gloxylate cycle which is modified Krebs cucle. The enzymes present in Glyoxysomes are isocitrate lyase and maltase synthase and several essential enzymes of Krebs cycle.
Cell organelles
~ 9 ~ Shanes notes on Cell Biology and Biomolecules Differences between peroxisomes and Glyoxysomes Peroxisome: 1. Peroxisomes are single membrane microbodies found in photosynthetic cells of plants and liver and kidney cells of vertebrates. 2. It contains enzymes like peroxides, oxidases and catalases. 3. It is involved in formation and degradation of hydrogen peroxide and detoxification of poisons in the cell. Glyoxysome: 1. Glyoxysomes are also single membrane microbodies but found only in plant cells. 2. It contains many enzymes like isocratic lyase, malate synthetase, glycolate oxidases, etc. for glyoxylate cycle (i.e., metabolism of glycolic acid). 3. It is involved in photorespiration and conversion of fats into carbohydrates.
VACUOLES (storage bin of the cell) Introduction: Structure A vacuole is a unit membrane-bound organelle which is present in all plant (predominantly) and fungal cells and some protists, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain cases they may contain solids which have been engulfed. Vacuoles are formed by the fusion of multiple membrane vesicles and are effectively just larger forms of these. The organelle has no basic shape or size, its structure varies according to the needs of the cell. The membrane covers the vacuole is called tonoplast. The vacuoles are responsible for isolating materials from the cell which may be harmful to it and also, containing waste products within themselves. As an autophagic vesicle, function of this cell organelle is to ingest and destruct any invading bacteria. It is also responsible for maintaining the turgor pressure in the cell and the pH of the cell
Fig: anatomical structure of vacuole Cell organelles
~ 10 ~ Shanes notes on Cell Biology and Biomolecules Functions: Some of the major function preformed by vacuoles in all cell organisms are as follows: Vacuoles used to contain a variety of enzymes. Apart from water and inorganic/organic molecules, different kinds of solid materials (in an engulfed form) are also present in vacuoles. This organelle can be described as kind of large vesicles. This is because; multiple membrane vesicles together form this cell organelle. The vacuoles have an important role in plant and protist cells when compared to that of animal and bacterial cells. The vacuole is considered as a storage bin of cells. It is also one of the important organelles other than cell nucleus. There are many different functions carried out by vacuoles and which are enlisted below: (i) Waste products generated in cells are accumulated in vacuoles. Storage of waste products in vacuoles protects other cell organelles from their harmful effects. (ii) Maintaining the right pH level also is one of the functions of this cell organelle. An acidic internal pH is maintained by vacuoles. (iii) Apart from pH maintenance, turgor and hydrostatic pressure is also maintained in the cells by these organelles. (iv) There are certain substances generated in cells which have the potential to harm/disturb the health of cells. Vacuoles carry out the crucial job of isolating these harmful substances. (v) Unwanted materials present in cells are removed by vacuoles. (vi) Vacuoles play an important role in the process of autophagy. A balance between biogenesis and degradation of substances of cells is maintained during this process by the vacuoles. (vii) Vacuoles are known to protect the cells from certain bacteria. Destroying the invading bacteria is an important function of these cell organelles. (viii) The food vacuoles found in amoeba perform the task of digestion. Food ingested by amoeba is digested with the help of this organ. (ix) The function of vacuole in animal cells is not much important in comparison to that in plant cells.
Cell organelles
~ 11 ~ Shanes notes on Cell Biology and Biomolecules PLASTIDS Introduction: Plastids are found scattered in most plant cells and in some photosynthetic protists. These are absent in most bacteria and fungi and few algae. On the basis of presence or absence of plastids the living world has been separated into plant and animal kingdom. Classification of plastids: Plastids may be divided into two general groups: 1. The chromoplasts or chromatophores having pigment. These further be differentiated into photosynthetically active (chloroplasts, pheoplasts, Rhodoplasts and blue green chromoplasts) and photosynthetically inactive chromoplasts (include crhomoplasts), and 2. Leucoplasts or colourless plastids (Amyloplasts, elaioplasts and aleuronplasts)
CHLOROPLASTS Introduction: The chloroplast which is a widely occurring plastid of green plants may be filamentous, saucer-shaped, spheroid, ovoid, discoid or club shaped. It is vesicular, having a colourless centre. Its size varies from 1 to 3 in thickness and 4 to 10 long (Hall et al., 1974). In higher plants there are usually 20 to 200 chloroplasts per cell. However, in many algae there may be only one or a few chloroplasts, and these may be star-shaped or a spiral ribbon or may have other exotic forms. The leaf of Ricinus communis contain about 4,00,000 chloroplasts per square millimeter of surface area. Distribution: Chloroplasts are frequently packed around the nucleus or close to the cell wall. But in certain cells they are homogeneously distributed within the cytoplasm. Their orientation is specific for different types of cells. Chemical composition: The chloroplasts are mainly composed of proteins, lipids, chlorophyll, cartenoids, RNA and DNA. In it are also present starch granules or osmophilic droplets, vitamin K and E, Phytoferritin etc.
Ultra structure of Chloroplasts: Structurally each plastid consists of two parts: (1) limiting membrane, (2) inner matrix or stroma. 1. Limiting membrane: This is a double membrane and each membrane is trilaminar and lipoproteinous and lacks chlorophyll and cytochromes. All molecular interchanges occur between the cytoplasm and chloroplast through this limiting membrane. The outer membrane is smooth and the inner membrane is much folded into the chloroplast stroma. The inner membrane is highly selectively permeable to molecules. Each membrane is of about 40-60 A 0 in thickness. Intervening space between the two membranes is called the periplastidial space and is 25-75 A 0 . Cell organelles
~ 12 ~ Shanes notes on Cell Biology and Biomolecules In Chlamydomonas, this is continuous with endoplasm. It serves as a semipermeable membrane reticulum and as the boundary of the plastids. Their chemical structure is essentially like that of reticulum. 2. Stroma (matrix): Plastids are filled with a watery, proteinaceous substance called the matrix or stroma. It contains about 50% of the chloroplast proteins and most of these are of the soluble type. It has ribosomes and also DNA. It also contains starch granules and osmiophilic droplets. During inactivity of the plastids the osmiophilic droplets increase, in number. In the stroma are embedded disc-like flattened structures made of double membranes. These discs are called lamellae (thylakoids). The outer surface of the thylakoid is in contact with the stroma and its inner surface encloses an intrathylakoid space. Thylakoids may be stacked like a pile of coins, forming the grana or they may be unstacked called stroma thylakoids. Thus, a granum consists of a series of membrane discs packed back to back, like a stack of coins. However, each disc is interconnected at an angle to all other discs in a granum by tubules called frets. By branching, a fret connects a disc to each of the other discs in turn. Thylakoids provide a large membrane area to hold the photosynthetic pigments and enzymes. Thylakoids containing chlorophyll (photic apparatus) for photosynthesis, permit separation of the light reactions that occur there from the dark reactions in the chloroplast stroma that fix CO 2 to synthesize sugars, starch, fatty acids and some proteins. ATP and reduced coenzymes diffuse from thylakoids where they are formed into the stroma, where they are used as energy source and reductant respectively, for the fixation of carbon dioxide.
Maintenance of grana and frets in a chloroplast requires the presence of inorganic salts. If the salts are replaced by organic buffer, the granum opens up to form an extensive membrane system. Replacement of the ions leads to spontaneous reformation of grana and frets. Cell organelles
~ 13 ~ Shanes notes on Cell Biology and Biomolecules Grana vary in size from 0.3 to 1.7 m in various species. In spinach chloroplasts, they are about 0.6 m in diameter. Membrane lamellae of the grana contain protein, and lipid layers, and in between these two is present the chlorophyll layer (Hodge, 1959). The chloroplasts may contain 40-60 grana in their stroma. Each granum of the chloroplasts of higher plant cells contains 10 to 100 disc-like, superimposed membranous compartments called the thylakoids. Recently Park and Pon (1968) have discovered some smaller paracrystalline spheroid bodies inside the membranes of thylakoids of the grana called quantasomes. These are roughly rectangular (18.0 nm 15.5 nm 10.0 nm). Each quantasome is composed of four subunits (Park and Beggins, 1964). The quantasomes are composed of about equal amounts of lipid and protein, and contain chlorophyll, carotenoids and other components of the photosynthetic apparatus. Quantasomes are plates or elipsoids measuring about 200 A in diameter and 100 A thick, arranged in a regular, lateral array. These quantasomes are believed to be the units of photosynthesis and each quantasome comprises about 300 molecules of chlorophyll, necessary amount for photosynthesis. Functions of Chloroplasts
Cell organelles
~ 14 ~ Shanes notes on Cell Biology and Biomolecules Mitochondria
Some of the main functions of Mitochondria are as follows: 1. Centre of respiration: Mitochondria are the respiratory centres, of cell. This view was postulated by Kingsbury (1912). They store energy in the form of adenosine triphosphate (ATP) which is released during glycolysis, Krebs cycle and electron transport mechanism of cell respiration. Glycolysis occurs in the cytoplasm of the cell, Krebs cycle takes place in the mitochondria. During this process the metabolic end products of various food stuffs, i.e., glucose or glycogen, amino acids and fatty acids, are degraded to CO 2
and water with the use of O 2 , and energy is liberated. This energy is locked in adenosine triphosphate (ATP), which is utilized in various body activities. Glycolysis (Glucose to acetyl coenzyme A): Glucose + 2ADP + 2 phosphate + 4 NAD + 2 Coenzyme A 2 acetyl coenzyme A + 2 2 + 2 ATP + 4 NADH + 4H + (ATP yield-2) Cell organelles
~ 15 ~ Shanes notes on Cell Biology and Biomolecules Krebs cycle: 2 Acetyl coenzyme A + 6H 2 O + 2 ADP + 2 phosphate + 2 FAD + 4 NAD 6CO 2 + 4 ATP + 2 FADH 2 + 10 NADH + 10 H + (ATP yield-2) Respiratory chain: 2 FADH 2 + 4 ADP + 4 phosphate + O 2 2 FAD + 4 ATP + 2H 2 O and (ATP yield-4)
10NADH + 10 H + +30 ADP + 30 phosphate + 5O 2 10 NAD + 30 ATP + 10 H 2 O (ATP yield-30) 2. Proteolytic activity: Horning described that mitochondria have proteolytic enzyme activity. In Protozoa, he has demonstrated both lytic and synthetic activity of mitochondria. In Amoeba, he found that pieces of engulfed food circulate in cytoplasm and later they become associated with mitochondria. He is of opinion that mitochondria are responsible for the production of zymogen granules of pancreas.
3. Histogenesis: Levi and Chevremont believe that myofibrils have mitochondrial organ. According to them, these mitochondria help in histogenesis (especially neurofibrils and muscle fibrils). 4. Fat metabolism: Wotton suggested by studying mitochondria of trypanosomes that these are associated with metabolism of fat. Recently, evidence of fat metabolism comes from Bensley (1947) on mitochondria of liver cells. Finally, he suggested that mitochondria really represent a reserve store of metabolic material for cell. In germinating seeds these help in diastatic activity. 5. Hereditary function: Recently, Luck et al., (1964) have discovered that mitochondria serve as cytoplasmic genes. They carry characters and thus help in genetic continuity of traits. 6. They play an important part in the formation of germ cells during its division (i.e., chondriokinesis). 7. Mitochondria are self-replicating organelles. Replication occurs in response to physiological requirements. They contain DNA, ribosomes and necessary enzymes to carry on protein synthesis as well as synthesis of phospholipids and other low molecular weight constituents. 8. Formation of mitochondrial spiral in the developing sperm: During maturation of a spermatid into a spermatozoon, the mitochondria form a spiral sheath around the axial filament of the middle piece of spermatozoon. Sheath is actually an end-to-end association of elongated mitochondria. They provide energy for the movement of sperm. Cell organelles
~ 16 ~ Shanes notes on Cell Biology and Biomolecules 9. In the oocytes of frog, Rana pipiens and fresh water snail, Planorbis, some of the mitochondria are transformed into yolk bodies. Yolk platelets accumulate inside the mitochondria and displace the cristae toward the periphery, transforming the mitochondria into yolk bodies. 10. Mitochondria is as a reservoir of energy: It is a source of energy for cellular activities. 11. In the flight muscles of insects, the mitochondria (called sarcosomes here) are intimately associated with the endoplasmic reticulum. They presumably develop endoplasmic reticulum (e.g. rat liver cells).