Cell Organelles in Biotechnology PDF
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Helwan National University
Dr. Ahmed Alazzouni
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This document provides an overview of cell organelles, focusing on their structure, function, and role in various biological processes. It covers cell membranes, proteins, receptors, and transport mechanisms, aiming to explain the complex workings of cellular components.
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Molecular structure of cell membranes Cell membranes (biomembranes) are important part of all cells. Their discovery is closely related to the upgrading of microscopic techniques, especially with the design of transmission electron microscopy, by which was observed typical trilaminar structure.Furt...
Molecular structure of cell membranes Cell membranes (biomembranes) are important part of all cells. Their discovery is closely related to the upgrading of microscopic techniques, especially with the design of transmission electron microscopy, by which was observed typical trilaminar structure.Further observations showed that biomembranes in the cell are similar in structure and slight differences in chemical composition are due to cell differentiation and specialization. Every cell is surrounded by the cytoplasmic membrane, which separates intracellular from extracellular space. Its average thickness is 60 – 100 nm. Functions of Cell membrane is selectively permeable boundary, cell which ensures the maintenance of dynamic equilibrium between cell and environment. membrane It contains enzymes, receptors, transport proteins, signalling systems and antigens. It performs different functions, e.g. intake of substances, interactions, recognition of signals, etc. The biological membrane is a part of many important cellular organelles. Dr. Ahmed Alazzouni The main components of cell membranes are phospholipids. The molecule of phospholipids is composed of polar (hydrophilic) head and two non-polar (hydrophobic) fatty acids chains. In the aqueous environment the hydrophilic parts are oriented towards the water around them and fatty acids chains to each other, creating so-called phospholipid bilayer.Given that phospholipids are not chemically bound to each other, their lateral movement is possible. Dr. Ahmed Alazzouni Other important components of biological membranes are proteins (Fig. 14). They may be: integral, which affect the hydrophobic parts of the phospholipid bilayer or transgress it. They are hardly separable from the biomembranes; peripheral, which lie outside the lipid bilayer. They are associated with electrostatic bonds and can be easily separated from biomembranes. Types and number of proteins in biomembrane is variable. It is dependent on cell differentiation and cell cycle phase. Specific protein composition of biomembranes is regulated by cell. Dr. Ahmed Alazzouni Functions of membrane protiens Membrane proteins perform various functions. They are part of biomembrane structure (structural proteins). Some of them are involved in the transport of ions across the membrane (pump and ion channels) or in the transfer of substances along the electrochemical gradient by facilitated diffusion. Many of them are part of the receptors that are able to specifically bind hormones,neurotransmitters and other signal molecules. Some have the role of enzymes. Proteins and glycolipids are part of the antigens. Dr. Ahmed Alazzouni Membrane receptors Membrane receptors Membrane receptors are protein structures located in cell membrane, which are responsible for recognition and binding of signal molecules (e.g. hormones, neurotransmitters etc.). Through these receptors cell interacts with its surroundings. Membrane receptors may be classified into: Dr. Ahmed Alazzouni receptors which are part of the ion channels – these are receptors for transport of cations (e.g. nicotine-acetylcholine receptor and receptor for excited amino acids) and anions (e.g. glycine receptor and receptor for gamma amino-butyric acid); Membrane receptors with enzyme activity are receptors with intracellular protein subunit which catalyzes receptors certain chemical reactions. These include receptors with tyrosine kinase activity, insulin receptor etc.; receptors coupled to G proteins – the largest group of membrane receptors (five families are known). Dr. Ahmed Alazzouni Transport of substances through the membrane Transfer of substances into cells and outside of cells is realized by two basic mechanisms. 1- passive transport. 2- active transport. Dr. Ahmed Alazzouni The passive transport The passive transport ensures transfer of substances in the direction of concentration gradient without consumption of energy (diffusion and osmosis). The speed of transition depends only on the size of the gradient (difference between concentrations in the cell and outside). Given the selective permeability of the cytoplasmic membrane only a few substances with low molecular weight (e.g. water, oxygen, carbon dioxide, urea, methanol and ethanol) can be transported by this way. Dr. Ahmed Alazzouni Diffusion Diffusion is an unordered movement of molecules in solution. It results in the movement of dissolved substances from the higher concentration to places of lower concentration. This movement will stop as soon as the concentration of the substance on both sides membrane equalized. Dr. Ahmed Alazzouni Osmosis Osmosis is a process in which water passes through the cytoplasmic membrane from the environment with a lower concentration in more concentrated environment. The process takes place until equalization of concentrations of both solutions. In case, that both solutions are isotonic to each other and cells that are in it perform no changes. For human cells the isotonic solutions are 0.9% NaCl saline and 5% glucose solution. If the solution in the extracellular environment is more concentrated than inside the cell, it is hypertonic solution. The cells lose water and shrink. In plant cells occurs plasmolysis (separation of the plasma membrane from the cell wall). If the solution outside the cell is of lower concentration as in the cell, it ishypotonic solution. Water penetrates into the cell. Animal cell increases in size and burst (cytolysis); e.g. haemolysis of red blood cells. In plant cells only it increases their turgor – cell wall prevents them against breaking. Phenomena of osmosis in plant and animal cells are presented by figure 17. Dr. Ahmed Alazzouni Facilitated diffusion Facilitated diffusion (Fig. 18) represents the transport of substances mediated by plasma membrane proteins without consumption of energy in the direction of concentration gradient. This process occurs so that the substance is bound to transport protein on the cell surface. It changes its conformation and the substance is released into the cytoplasm (e.g. transport of glucose). Dr. Ahmed Alazzouni Active transport is the transfer of substances against concentration or electrochemical gradient without moving membrane. This process ensures due to consumption of energy obtained by dissociation of ATP (adenosine triphosphate) to ADP (adenosine diphosphate) Active or up to AMP (adenosine monophosphate). transport This process is provided by special transportation systems (channels and pumps), protein complexes, which pass through the membrane. It is a rigorous selective and managed process, often controlled by receptors. There are two basic types of active transport – primary and secondary. Dr. Ahmed Alazzouni Primary active transport is realized against concentration or electrochemical gradient with the energy consumption (obtained by hydrolysis of ATP). This process is performed by cyclic phosphorylation and dephosphorylation of Primary transport proteins. This also changes the affinity to the substrate – alternately on the active outside and inside the membrane. The whole process can be summarized as follows – transport transported substance (substrate) is attached to phosphorylated transport protein; protein is dephosphorylated to open the binding site toward the cytoplasm and the substrate is released. The function of Na+– K+ pump (Na +-K +-ATPase) and H+ pump (H+–ATP–ase) is realized by this way. Dr. Ahmed Alazzouni Primary active transport Dr. Ahmed Alazzouni Secondary active transport secondary active transport, the affinity of membrane transport protein is not changed by phosphorylation, but by the attachment of ions (e.g. Na+). These proteins have two sites, first one for connection with ion and second one for transported substrate. In the case of the substrate and the ions are transported in the same direction, it is cotransport. In the case of transport in the opposite direction, it is antiport. Dr. Ahmed Alazzouni Endocytosis and exocytosis Transport of substances with high molecular weight is carried by Endocytosis endocytosis and exocytosis. Both processes are associated with active and participation of the cytoplasmic exocytosis membrane (changes in its structure or its movement). Dr. Ahmed Alazzouni Endocytosis Endocytosis is the process by which substances are transported into cells. According to transported substances we distinguished pinocytosis (especially the transport of soluble substances) and phagocytosis (transport of solids). Dr. Ahmed Alazzouni pinocytosis pinocytosis the transported substance is attached to receptor in the plasma membrane, which “creates” hollow inside the cytoplasmic membrane. The resulting cavity gradually increasing and surrounded transported substance. Finally, it is closed and creates a pinocytotic vesicle, which is released into cell and travels to the place of further processing. After that plasma membrane integrity is Dr. Ahmed Alazzouni restored Phagocytosis Phagocytosis represents transport of solid particles, for example phagocytosis of bacteria. The cell generates plasma membrane processes (pseudopodia) which surround the transported material. It enters cells in vesicles and is processed. Dr. Ahmed Alazzouni exocytosis exocytosis the substances are transported from the cell into the extracellular environment. Secreted material is located in vesicles, which usually arise from endoplasmic reticulum and Golgi apparatus. Vesicle approaches the plasma membrane, touched her, and merging with it and the substances are released into the environment. Dr. Ahmed Alazzouni Endocytosis and exocytosis Dr. Ahmed Alazzouni Cytoplasm represent basic inner environment of the cell. It is composed mainly of water (70 – 80%). It also includes a considerable number of ions, inorganic and organic compounds (e.g. fatty acids, amino acids, lipids, carbohydrates, nucleic acids, proteins). Cytoplasm is a semi-liquid mass, which should be in form of sol (liquid) or gel. Protein content and their ability to bind water influences its viscosity. The cytoplasm of eukaryotic Cell cells contains a variety of cell organelles. According to their composition cell organelles are organelles distinguished into three basic types: membrane, which are composed from one membrane (endoplasmic reticulum, Golgi apparatus, lysosomes, peroxisomes, vacuoles and other vesicles) or from two membranes (nuclear envelope, mitochondria and chloroplasts); composed of proteins – cytoskeleton (microtubules, microfilaments, intermediate filaments, flagellas, cilia etc.); composed of proteins and nucleic acids – ribosomes, nucleolus. Dr. Ahmed Alazzouni Mitochondria are organelles inevitable for the life of eukaryotic cells. They are involved in generating energy for cells by breaking down of saccharides, lipids and other energy- rich organic compounds. Mitochondria can vary in number and shape according to the type of cell. Their number is in a direct proportion to the intensity of cell’s energy metabolism. They consist of two biological membranes. Mitochondria The outer membrane encloses it while the inner one is folded into the mitochondrial cristae expanding its surface. The enzyme system (H+ATP synthase) is localized in the inner membrane and is responsible for cellular respiration (oxidative phosphorylation). This is where the glucose is broken down with the freed energy bounding into ATP molecules (ADP + E + P = ATP). It also contains other important enzymes, e.g. cytochromes, NADP dehydrogenases etc. Dr. Ahmed Alazzouni The space between the cristae is filled with mitochondrial matrix made up of phospholipoproteins and ions of calcium and magnesium. Moreover it also contains enzymes of the citric acid cycle (the Krebs cycle). Mitochondria have their own circular DNA and ribosomes of the prokaryotic type. This enables their auto-reproduction and synthesis of their own proteins. Most of mitochondrial enzyme substance is however coded by nuclear genes. They are synthesized on endoplasmic reticulum, modified in the endoplasmic reticulum and completed in Golgi apparatus. Consequently, they are transported into mitochondria where they are enhanced with proteins synthesized in mitochondria and become functional. Dr. Ahmed Alazzouni Endoplasmic reticulum Endoplasmic reticulum (ER) is a system of tubules, cisterns and flat vesicles from the biological membrane closely communicating with the nucleus and via transport vesicles with the Golgi apparatus and cytoplasm. ER is responsible for synthetic processes. Dr. Ahmed Alazzouni Smooth ER From the morphological and functional aspect, we distinguish between two types of endoplasmic reticulum. Smooth ER which is the place of synthesis of saccharides and lipids (including parts of the biomembranes), steroid hormones and cholesterol. The detoxication function of the smooth ER is also significant – excessive cell‘s intoxication leads into apoptosis. The muscle cells contain a special form of endoplasmic reticulum – the sarcoplasmic reticulum. It ensures the transport of Ca2+ cations that are inevitable for muscle contraction. Dr. Ahmed Alazzouni Rough endoplasmic reticulum Rough endoplasmic reticulum (Fig. 25b) contains ribosomes on its surface and is the place of protein synthesis. Formed proteins penetrate its structure and are modified for the first time. For further processing they are transported in a vesicle from the membrane into the Golgi apparatus. The ratio of the rough and smooth ER in a cell depends on its function and products it creates. Dr. Ahmed Alazzouni Golgi apparatus It is made up of a system of flat cisterns whose sides are from the functional aspect divided into cis and trans. Proteins synthesized on the endoplasmic reticulum, usually enclosed in a transport vesicle, come to the cis side. Their posttranslational modification continues inside of the Golgi apparatus. They leave Golgi apparatus from the trans side in the form of vesicles to cytoplasm. Some of them fulfill specific tasks in a cell, others join the cytoplasmic membrane and their content comes into the extracellular environment. Some vesicles only take part in “circulation” of cell membranes. Dr. Ahmed Alazzouni Functions of Golgi Apparatus The Golgi apparatus is a complex of stacked membranous sacs that is crucial for the processing, packaging, and further distribution of proteins and lipids coming from the ER. The formation of secretory vesicles also takes place here. Its main function is the packaging and secretion of proteins. It receives proteins from Endoplasmic Reticulum. It packages it into membrane-bound vesicles, which are then transported to various destinations, such as lysosomes, plasma membrane or secretion. The Golgi apparatus is responsible for transporting, modifying, and packaging proteins and lipids into vesicles for delivery to targeted destinations. As the secretory proteins move through the Golgi apparatus, a number of chemical modifications may transpire. Dr. Ahmed Alazzouni Lysosomes and other vesicles Lysosomes are vesicles from single biological membrane created by being detached from the Golgi apparatus as a primary lysosome. They contain digestive enzymes (e.g. hydrolases for protein digestion). Following connection with an unneeded organelle or pinocytotic (fagocytotic) vesicle creates a secondary lysosome and lysosomal enzymes break down the content of these structures. In plant cells and some protists, the function of lysosomes and many other tasks is performed by vacuoles. Dr. Ahmed Alazzouni Peroxisomes belong to the large family of vesicles in cytoplasm, the role of which is to ensure activity of various Peroxisomes enzymes. Peroxisomes contain catalase eliminating the aggressive hydrogen peroxide. Enzymes in the cytosolic vesicles are thus available for the cell; however, they are not directly contained in the cytoplasm, but used when the cell needs them. Dr. Ahmed Alazzouni Ribosomes These might be one of the smallest cell organelles, but they are crucial, because of protein synthesis. They are made up of a large and small sub-unit that is connected only during translation (the protein synthesis). The sub-units are made up of proteins and ribosomal RNA. The main function of ribosome is protein synthesis. Dr. Ahmed Alazzouni ribosomes of eukaryotic cells The ribosomes of eukaryotic cells are larger and their gravitation density is 80S. They consist of four types of rRNA and 82 proteins. They are present freely in cytoplasm, but their prevailing majority participates in the creation of rough ER. This enables the eukaryotic cell to effectively manage the course of protein synthesis. In the eukaryotic cells there are also ribosomes of the prokaryotic type, specifically in mitochondria and chloroplasts. These organelles synthesize their own proteins needed in order to activate their enzymes and auto-reproduction Dr. Ahmed Alazzouni Cytoskeleton Cytoskeleton (Fig. 27) is the internal functional and dynamic skeleton of eukaryotic cells. It participates in forming of the shape of cells, distribution of organelles and performing of some intracellular activities (e.g. movement, contraction etc.). Cytoskeleton consists of protein- based microtubules, microfilaments, intermediate filaments and microtrabecules, while each of these plays a different role Dr. Ahmed Alazzouni Microtubules Microtubules are the same in all eukaryotes. They are long, firm and hollow cylinders about 25 nm in diameter comprising the molecules of tubulin (dimers consist of _ and _ monomers). Microtubules are made longer or shorter through polymerization and depolymerization (adding or removing of tubulin dimers). They participate in the transport of vesicles between endoplasmic reticulum, the Golgi apparatus and cytoplasmic membrane. They play an important role as part of mitotic spindle fibers in the movement of chromosomes during cell division. They also make up the structure of centrioles, tails and cillia. Dr. Ahmed Alazzouni Microfilaments are thin (7 nm in diameter) filaments of helical structure made up of globular protein called actin to which the proteins of myosin are bound together creating (via clutching) a contractile system of muscle cells. Intermediate filaments are think fibers (10 nm in diameter) that are not able of contraction. These include, for example, lamins of the nuclear matrix, epithelial keratins, vimentins of the intercellular substance (EDM) of connective tissues – networks of proteins connecting cells and neurofilaments (the transport system of nerve cells). They provide for cell resistance to pulling and pressing. They also participate in the distribution of organelles and inclusion within a cell. Dr. Ahmed Alazzouni Centrosome The centrosome is a structure of eukaryotic cells located near the nucleus. It is of vital importance for the process of cell division. It consists of two complexes of three couples of orthogonally arranged tubulin fibers (centrioles). The centrosome doubles prior to cell division. Dr. Ahmed Alazzouni Ribosomes These might be one of the smallest cell organelles, but they are crucial, because of protein synthesis. They are made up of a large and small sub-unit that is connected only during translation (the protein synthesis). The sub- units are made up of proteins and ribosomal RNA. Dr. Ahmed Alazzouni The ribosomes of eukaryotic cells are larger and their gravitation density is 80S. They consist of four types of rRNA and 82 proteins. They are present freely in cytoplasm, but their prevailing majority participates in the creation of rough ER. This enables the eukaryotic cell to effectively manage the course of protein synthesis. In the eukaryotic cells there are also ribosomes of the prokaryotic type, specifically in mitochondria and chloroplasts. These organelles synthesize their own proteins needed in order to activate their enzymes and auto- reproduction. Dr. Ahmed Alazzouni Nucleus Nucleus is the coordinating and control center of cells. It contains the genetic information that is embedded in the structure of DNA. The inner structure of nucleus is very complex and is characterized by high dynamics of the processes that occurs there. During the cell cycle is periodically changed. Interphase nucleus is observed between two mitotic divisions. The second form is the mitotic nucleus which morphologically “disappears” during the indirect cell division. Dr. Ahmed Alazzouni Nuclear envelope of eukaryotic cells consists of two membranes. The inner Membrane encloses the contents and the outer is in contact with the cytoplasm and Endoplasmic reticulum. The space between these membranes is wide 20 – 80 nm and is called as perinuclear space. Both membrane in some places are merged and creates nuclear pores. Nuclear pores represent complicated gaps in the nuclear membrane with a diameter of about 70 nm and occupy 5 – 25 % of nuclear envelope surface From upper view the nuclear pore has circular shape Nuclear pores provide active transport of substances from the nucleus to the cytoplasm (especially RNA subunits and ribosomes) and from the cytoplasm to the nucleus (e.g. transport of histones, nutrients and regulatory proteins). Dr. Ahmed Alazzouni nuclear matrix The nuclear matrix consists of the insoluble structural framework of the nucleus, which includes the nuclear lamina and pore complex, an internal ribonucleoprotein network, and residual nucleolus The nuclear matrix is the residual framework scaffolding of the nucleus and consists of the peripheral lamins and pore complexes, an internal ribonucleic protein network, and residual nucleoli The nuclear matrix is a three-dimensional filamentous protein network, found in the nucleoplasm, which provides a structural framework for organising chromatin, while facilitating transcription and replication. Dr. Ahmed Alazzouni Kinetochore Kinetochore, a structure important for cell division, is also made up of proteins. It is the protein complex attaching to the centromere of chromosomes (chromatids) and enabling in the course of cell division free attachment of tubulin fibers of the mitotic spindle. Dr. Ahmed Alazzouni Centrosome The centrosome is a structure of eukaryotic cells located near the nucleus. It is of vital importance for the process of cell division. It consists of two complexes of three couples of orthogonally arranged tubulin fibers (centrioles). The centrosome doubles prior to cell division. Dr. Ahmed Alazzouni centromere vs centrosome Dr. Ahmed Alazzouni chromosome Dr. Ahmed Alazzouni Chromatin Interphase nucleus is filled by seemingly amorphous mass – chromatin which is composed of linear DNA associated with the histon or non-histon proteins. Histon proteins have structural and regulatory function. Non-histon proteins have mainly regulatory functions and manage the internal organization of the nucleus. Dr. Ahmed Alazzouni In eukaryotic cells, we distinguish two types of chromatin – heterochromatin and euchromatin Heterochromatin consists of condensed chromosome segments and produces dense aggregations, which are mostly located near the nuclear envelope. It can be also irregularly distributed throughout the nucleus. Heterochromatin located by the nucleolus is called perinucleolar chromatin. If euchromatin is in the nucleolus, it is intranucleolar chromatin. It is a transcriptional inactive chromatin. Euchromatin has soft, foamy or almost fibrillar structure. It consists of decondensed segments of chromosomes. It is transcriptional active and is present in the cells with high protein synthetic activity. Dr. Ahmed Alazzouni Nucleolus Integral part of eukaryotic cell nucleus is the nucleolus. This functional organelle is most abundant in the G1 phase of the cell cycle. It has spherical shape of about 1-5μm. From a chemical point of view it is formed of RNA molecules and proteins. It is a place of Rrna synthesis, post-transcriptional modification and completisation of ribosome subunits. In electron microscope we distinguished three basic parts of nucleolus: pars granulosa – composed of ribonucleoprotein particles, which are precursors of ribosomes; pars fibrosa – composed of soft filaments, which are stored close to each other. Contains precursors of rRNA; nucleolar organizer region – the place of nucleolus restoration after cell division. Dr. Ahmed Alazzouni cell cycle The cell cycle consists of two main phases, which are interphase and M-phase (mitosis phase). The individual phases of the cell cycle proceed after each other. The process is regulated by a complex of regulatory proteins, which are coded by tumor suppressor genes (they have a control function) and protooncogenes (stimulating division). A failure of their normal function can cause deregulation of the cell cycle and a consecutive malign transformation of the cell, meaning a change to a cancer cell. Dr. Ahmed Alazzouni Interphase is a time between two divisions and is made up of G1, S a G2 phase. The duration of the phases differs and depends on the type of the cell and the life period of the individual. Dr. Ahmed Alazzouni G1 phase G1 phase begins after the end of the previous mitosis. It is characterized by an intensive synthesis of proteins, and usually also growth and of new cell. During this process, in the so called G0 phase, the cell differentiates, to fulfill specialized functions for organism. Time duration of G0 phase is the most variable component of the interphase. At the end of the G1 phase is the so called main checkpoint, in which it is decided weather the cycle will continue or not. Dr. Ahmed Alazzouni S phase (synthetic) S phase (synthetic) is a time during which the duplication (semiconservative replication) of the nuclear DNA occurs. Considering the length of the DNA in the nucleus (in women around 2 m) and the processes of their repeating control and the repair of defects, it is the longest phase of the cell cycle, even tough the replication takes place at numerous places simultaneously. At its end each chromosome is doubled, meaning it consists of two chromatids connected. Dr. Ahmed Alazzouni G2 phase G2 phase is a relatively short period of preparation for mitosis and it contains another checkpoint of the cell cycle. After replication it is important for the cell to check the DNA and repair the potential mistakes. It is also important to prepare the necessary proteins, mainly tubulin, as well as sufficient sources of energy. During this phase the duplication of the centrosome occurs (made up of centrioles). At the same time, on the second centrosome, a so called astral complex and the basis of non- kinetochore microtubules are formed. Dr. Ahmed Alazzouni M phase M phase - mitosis is a part of the cell cycle, during which the division of the nucleus occurs (karyokinesis) and consecutively the division of the whole cell (cytokinesis) happens. It was first described and named by Walther Flemming (1887 – 1880). Mitosis is divided into five phases – prophase, prometaphase, metaphase, anaphase and telophase. Dr. Ahmed Alazzouni 1- prophase Prophase is the first phase of mitosis. This part of mitosis is all about preparing. During this phase, the DNA forms into chromosomes, which we can actually see. Chromosomes are condensed chromatin. In this phase, the chromosomes consist of two identical chromatids called sister chromatids. Steps of prophase are:- 1. 1- Mitotic spindles begin to form. 2. 2- The nucleolus (organelle in the nucleus that makes ribosomes) disappears. 3. 3- The nuclear envelope disintegrates and centrosomes move to opposite ends (poles) of the cell.. Dr. Ahmed Alazzouni prometaphase the movement of the centrosomes towards the poles continues. The condensation of the chromosomes goes ahead and they can be observed as stick-like formations. The process of chromatid separation from the end part of the chromosomes (telomeres) begins. On the outer side of each chromatid centromere functional kinetochores are formed. At the same time, kinetochore microtubules (KMT) “grow out” from each centrosome (elongated by polymerization of tubulin dimers) and enter the area of the former nucleus – “searching” for connection to kinetochores. When kinetochore microtubules connect to both kinetochores of particular doubled chromosome, they begin to elongate and shorten (by depolymerization), to transport the chromosome to the central (equatorial) plain of the cell. This takes a certain amount of time, making prometaphase the longest period of mitosis. Dr. Ahmed Alazzouni Metaphase is a relatively short period during which the duplicated chromosomes are located in the equatorial plain of the cell. The centrosomes are pushed to the opposite sites of the cell – spindle body is finished. All the kinetochores are occupied by kinetochore microtubules. Cohesins, except for the parts between centromeres of sister chromatids, are destroyed. This is why the metaphase chromosomes have the shape of the letter X. By this, all the conditions for the activation of the so called anaphase promotion complex (APC) are fulfilled and mitosis can continue to anaphase – chromatids are separated and transfer of daughter chromosomes can start. Dr. Ahmed Alazzouni anaphase anaphase (Fig. 31) two parallel processes take place. Anaphase A is characterized by the shortening of the KMT, which is responsible for transporting (“pulling”) of the individual daughter chromosome, to the centrosomes. Anaphase B the elongation of the non- kinetochore microtubules continues which elongates the whole cell and creates the space for cytokinesis. Both processes are supported by the activity of the so called motor proteins – dyneins and kinesins. Dr. Ahmed Alazzouni Telophase telophase the nucleus is reformed close to each centrosome. Theformation of two new nuclei in the cell is called karyokinesis. Chromosomes decondense and the functional organization of the nuclei is renewed. Parallelly – the cell divides (cytokinesis) and two new identical daughter cells are formed. Important is, that each of the two daughter cells retains one centrosome near the nucleus with the base of the non-kinetochore microtubules – new cell keeps the essential components necessary for the next division. If the cytokinesis doesn’t take place, a so called syncytium is formed. Dr. Ahmed Alazzouni