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Cytoskeleton Providing structural support to the cell, the cytoskeleton also functions in cell motility and regulation Structural Support • Mechanical support – Maintains shape • Fibers act like a geodesic dome to stabilize and balance opposing forces • Provides anchorage for organelles • Dynamic – Dismantles in one spot and reassembles in another to change cell shape • Introduction • The cytoskeleton is a network of fibers extending throughout the cytoplasm. • The cytoskeleton organizes the structures and activities of the cell. • The cytoskeleton also plays a major role in cell motility. – This involves both changes in cell location and limited movements of parts of the cell. • The cytoskeleton interacts with motor proteins. – In cilia and flagella motor proteins pull components of the cytoskeleton past each other. – This is also true in muscle cells. Fig. 7.21a • Motor molecules also carry vesicles or organelles to various destinations along “monorails’ provided by the cytoskeleton. • Interactions of motor proteins and the cytoskeleton circulates materials within a cell via streaming. • Recently, evidence is accumulating that the cytoskeleton may transmit mechanical signals that rearrange the nucleoli and other structures. Fig. 7.21b • There are three main types of fibers in the cytoskeleton: microtubules, microfilaments, and intermediate filaments. • Microtubules, the thickest fibers, are hollow rods about 25 microns in diameter. – Microtubule fibers are constructed of the globular protein, tubulin, and they grow or shrink as more tubulin molecules are added or removed. • They move chromosomes during cell division. • Another function is as tracks that guide motor proteins carrying organelles to their destination. Fig. 7.21b • In many cells, microtubules grow out from a centrosome near the nucleus. – These microtubules resist compression to the cell. • In animal cells, the centrosome has a pair of centrioles, each with nine triplets of microtubules arranged in a ring. • During cell division the centrioles replicate. Fig. 7.22 • Microtubules are the central structural supports in cilia and flagella. – Both can move unicellular and small multicellular organisms by propelling water past the organism. – If these structures are anchored in a large structure, they move fluid over a surface. • For example, cilia sweep mucus carrying trapped debris from the lungs. Fig. 7.2 • Cilia usually occur in large numbers on the cell surface. – They are about 0.25 microns in diameter and 2-20 microns long. • There are usually just one or a few flagella per cell. – Flagella are the same width as cilia, but 10-200 microns long. • A flagellum has an undulatory movement. – Force is generated parallel to the flagellum’s axis. Fig. 7.23a • Cilia move more like oars with alternating power and recovery strokes. – They generate force perpendicular to the cilia’s axis. Fig. 7.23b • In spite of their differences, both cilia and flagella have the same ultrastructure. – Both have a core of microtubules sheathed by the plasma membrane. – Nine doublets of microtubules arranged around a pair at the center, the “9 + 2” pattern. – Flexible “wheels” of proteins connect outer doublets to each other and to the core. – The outer doublets are also connected by motor proteins. – The cilium or flagellum is anchored in the cell by a basal body, whose structure is identical to a centriole. Fig. 7.24 • The bending of cilia and flagella is driven by the arms of a motor protein, dynein. – Addition to dynein of a phosphate group from ATP and its removal causes conformation changes in the protein. – Dynein arms alternately grab, move, and release the outer microtubules. – Protein cross-links limit sliding and the force is expressed as bending. Fig. 7.25 • Microfilaments, the thinnest class of the cytoskeletal fibers, are solid rods of the globular protein actin. – An actin microfilament consists of a twisted double chain of actin subunits. • Microfilaments are designed to resist tension. • With other proteins, they form a threedimensional network just inside the plasma membrane. Fig. 7.26 The shape of the microvilli in this intestinal cell are supported by microfilaments, anchored to a network of intermediate filaments. • In muscle cells, thousands of actin filaments are arranged parallel to one another. • Thicker filaments, composed of a motor protein, myosin, interdigitate with the thinner actin fibers. – Myosin molecules walk along the actin filament, pulling stacks of actin fibers together and shortening the cell. Fig. 7.21a • In other cells, these actin-myosin aggregates are less organized but still cause localized contraction. – A contracting belt of microfilaments divides the cytoplasm of animals cells during cell division. – Localized contraction also drives amoeboid movement. • Pseudopodia, cellular extensions, extend and contract through the reversible assembly and contraction of actin subunits into microfilaments. Fig. 7.21b • In plant cells (and others), actin-myosin interactions and sol-gel transformations drive cytoplasmic streaming. – This creates a circular flow of cytoplasm in the cell. – This speeds the distribution of materials within the cell. Fig. 7.21c • Intermediate filaments, intermediate in size at 8 12 nanometers, are specialized for bearing tension. – Intermediate filaments are built from a diverse class of subunits from a family of proteins called keratins. • Intermediate filaments are more permanent fixtures of the cytoskeleton than are the other two classes. • They reinforce cell shape and fix organelle location. Fig. 7.26