Lecture 6 The cytoskeleton

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1 Lecture 6 The cytoskeleton

2 The Cytoskeleton 1.Three kinds of filaments: microtubules, actin filaments, intermediate filaments 2. Functions of the cytoskeleton: structure, attachment, compartmentalization, motility, transport, cell division 3. Methods for studying the cytoskeleton: EM, antibodies, fusion proteins, drugs, cell free assays

3 1) Microtubules - 25nm Cytoskeleton Filamentous network that provides the structural framework of cells 2) Intermediate filaments nm 3) Actin filaments (thin filaments, microfilaments) - 6nm

4 The cytoskeleton is highly dynamic

5 Microtubules Tubulin is a heterodimer consisting of alpha and beta-tubulin. Protofilaments are linear polymers of tubulin. Polymerization is GTP-dependent. Microtubules are cylinders of 25nm in diameter composed of 13 protofilaments.

6 Actin filaments (microfilaments) Made up of actin monomers (G-actin) assembled into filaments (F-actin) Polymerizes in an ATP-dependent manner

7 Structure of actin filaments Direct visualization of secondary structures of F-actin by electron cryomicroscopy. Fujii T, Iwane AH, Yanagida T, Namba K. Nature Oct 7;467(7316):724-8.

8 Actin has diverse and essential functions in every cell

9 Intermediate filaments Includes a large number of ultrastructurally similar (8-10 nm), but chemically distinct filaments Contribute to cell shape and stability

10 Intermediate filaments

11 Intermediate filaments Lamins are intermediate filaments that maintain nuclear shape

12 Figure Molecular Biology of the Cell ( Garland Science 2008)

13 Intermediate filaments may be anchored at junctions Distribution of keratin on an epithelial cell visualized with antikeratin antibodies

14 Intermediate filaments may be anchored at junctions Distribution of keratin on an epithelial cell visualized with antikeratin antibodies

15 Intermediate filaments may be anchored at junctions Weber, G.F., Bjerke, M.A., and DeSimone, D.W. (2012). A mechanoresponsive cadherin-keratin complex directs polarized protrusive behavior and collective cell migration. Developmental Cell 22,

16 Intermediate filaments can withstand deformation

17 Keratin deficiency can cause blistering

18 Functions of the cytoskeleton 1. Maintenance of cell shape, structural support 2. Stabilization of cell attachments 3. Maintenance of local specializations 4. Cell motility 5. Intracellular transport of proteins, organelles, mrnas 6. Mitosis and meiosis

19 1. Maintenance of cell shape, structural support 2. Stabilization of cell attachments 3. Maintenance of local specializations Figure 16-5 Molecular Biology of the Cell ( Garland Science 2008)

20 Figure 16-4 Molecular Biology of the Cell ( Garland Science 2008) 4. Cell motility

21 4. Cell motility David Rogers, Vanderbilt University, late 1950s

22 5. Intracellular transport of proteins, organelles, mrnas Kinesins and dyneins transport cargo along microtubules

23 6. Mitosis and meiosis Microtubules form the spindle: Dividing sand dollar embryo--microtubules and DNA labeled

24 6. Mitosis and meiosis Actin plays a role in cytokinesis: Sea Urchin embryos stained for microtubules (white) and myosin II (magenta), which binds to actin filaments

25 The cytoskeleton 1.Three kinds of filaments: microtubules, actin filaments, intermediate filaments 2. Functions of the cytoskeleton: structure, attachment, compartmentalization, motility, transport, cell division 3. Methods for studying the cytoskeleton: EM, antibodies, fusion proteins, drugs, cell free assays

26 Electron microscopy provides a static, but high resolution view of the cytoskeleton TEM of hemidesmosome with emanating intermediate filaments

27 Electron microscopy provides a static, but high resolution view of the cytoskeleton Platinum Replica Electron Microscopy Tatjana Svitkina Technique: detergent extraction, triple fixation, critical point drying, and shadowing with platinum and carbon

28 Electron microscopy provides a static, but high resolution view of the cytoskeleton Wildtype growth cone Arp2/3 complex is important for filopodia formation, growth cone motility, and neuritogenesis in neuronal cells. Korobova F, Svitkina T. Mol Biol Cell Apr;19(4):

29 Electron microscopy provides a static, but high resolution view of the cytoskeleton Arp2/3 deficient growth cone Arp2/3 complex is important for filopodia formation, growth cone motility, and neuritogenesis in neuronal cells. Korobova F, Svitkina T. Mol Biol Cell Apr;19(4):

30 Antibodies against components of the cytoskeleton reveal their organization in a cell Figure 16-1 Molecular Biology of the Cell ( Garland Science 2008)

31 Fluorescent fusions to cytoskeletal components has allowed the monitoring of cytoskeletal dynamics in live cells GFP-tubulin fusion GFP-actin fusion Memo-RhoA-mDia1 signaling controls microtubules, the actin network, and adhesion site formation in migrating cells. Zaoui K, Honoré S, Isnardon D, Braguer D, Badache A. J Cell Biol Nov 3;183(3):401-8

32 Fluorescent fusions to cytoskeletal components has made it possible to monitor cytoskeletal dynamics in live cells Speckle microscopy: a method to watch individual molecules How microtubules get fluorescent speckles. Waterman-Storer CM, Salmon ED. Biophys J Oct;75(4):

33 Fluorescent fusions to cytoskeletal components has allowed the monitoring of cytoskeletal dynamics in live cells Speckle microscopy: a method to monitor filament dynamics How microtubules get fluorescent speckles. Waterman-Storer CM, Salmon ED. Biophys J Oct;75(4):

34 Fluorescent fusions to cytoskeletal components has allowed the monitoring of cytoskeletal dynamics in live cells Fluorescent speckle microscopy, a method to visualize the dynamics of protein assemblies in living cells. Waterman-Storer CM, Desai A, Bulinski JC, Salmon ED. Curr Biol Nov 5;8(22):

35 Drugs affecting cytoskeletal components in specific ways facilitate the study of cytoskeletal function Table 16-2 Molecular Biology of the Cell ( Garland Science 2008)

36 Drugs affecting cytoskeletal components in specific ways have advanced the study of the cytoskeleton colchicine low colchicine no drug APM

37 Drugs affecting cytoskeletal components in specific ways have advanced the study of the cytoskeleton Colchicine inhibits cancer metastasis Colchicine inhibits pressure-induced tumor cell implantation within surgical wounds and enhances tumor-free survival in mice. David H. Craig, Cheri R. Owen, William C. Conway, Mary F. Walsh, Christina Downey, Marc D. Basson. J Clin Invest. 2008; 118(9):

38 Cell free assays have made it possible to study cytoskeletal components in a controlled fashion In vitro actin polymerization assay

39 Cell free assays have made it possible to study cytoskeletal components in a controlled fashion

40 Cytoskeleton summary 1. Microtubules consist of alpha and beta tubulin dimers assembled into protofilaments, which in turn form large hollow filaments 2. Microfilaments are made up of actin, which polymerize in an ATP-dependent manner. 3. Intermediate filaments (e.g. keratin) provide structural support 4. Functions of the cytoskeleton: structure, attachment, compartmentalization, motility, transport, cell division 5. Methods for studying the cytoskeleton: EM, antibodies, fusion proteins, drugs, cell free assays

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