Chapter 17

Rank the following cytoskeletal filaments from smallest to largest in diameter (1 = smallest in diameter, 4 = largest) ______ intermediate filaments ______ microtubules ______ actin filament ______ myofibril

2, 3, 1, 4

In the budding yeast, activation of the GTP-binding protein Cdc42 occurs on binding of an external signal (pheromone) to a G-protein-coupled receptor. Activation of Cdc42 promotes actin polymerization. Predict what would happen to actin polymerization, in comparison with pheromone-treated cells, in the following cases. A. You add pheromone to an inhibitor of G-protein-coupled receptors. B. You add pheromone to a nonhydrolyzable analog of GTP.

A. Less actin polymerization B. More actin polymerization

Microtubules are formed from the tubulin heterodimer, which is composed of the nucleotide-binding __________________ protein and the __________________ protein. Tubulin dimers are stacked together into protofilaments; __________________ parallel protofilaments form the tubelike structure of a microtubule. __________________ rings are important for microtubule nucleation and are found in the __________________ , which is usually found near the cell's nucleus in cells that are not undergoing mitosis. A microtubule that is quickly growing will have a __________________ cap that helps prevent the loss of subunits from its growing end. Stable microtubules are used in cilia and flagella; these microtubules are nucleated from a __________________ and involve a "__________________ plus two" array of microtubules. The motor protein __________________ generates the bending motion in cilia; the lack of this protein can cause Kartagener's syndrome in humans. α-tubulin dynein nine ATP four thirteen basal body γ-tubulin twenty-one β-tubulin GTP UTP centrosome kinesintwo vimentin δ-tubulin myosin

B-tubulin, a-tubulin, thirteen, y-Tubulin, centrosome, GTP, basal body, nine, dynein

Actin can adopt a variety of shapes. Match the name of the actin form with the type of actin structure depicted as black lines within the cells in Q17-43. Figure Q17-43 _____ lamellipodia _____ contractile bundles _____ contractile ring _____ microvilli

C, B, D, A

Place the following in order of size, from the smallest to the largest. A. protofilament B. microtubule C. α-tubulin D. tubulin dimer E. mitotic spindle

C, D, A, B, E

Match the type of intermediate filament with its appropriate location. lamins______________ neurofilaments____________ vimentins______________ keratins___________ A. nerve cells B. epithelia C. nucleus D. connective tissue

CADB

The following proteins are important for cell movement. Match the following proteins with their function. myosin _______ ARP proteins ________ profiling _______ integrins _______ formins ________ A. nucleation of new actin filaments at the side of an existing filament B. regulation of the availability of actin monomers C. important for the growth of straight, unbranched actin filaments D. contracting the rear of the cell E. involvement in focal contacts

D, A, B, E, C

Do you agree or disagree with the following statement? Explain your answer. When skeletal muscle receives a signal from the nervous system to contract, the signal from the motor neuron triggers the opening of a voltage-sensitive Ca2+ channel in the muscle cells' plasma membrane, allowing Ca2+ to flow into the cell.

Disagree (The increase in intracellular Ca2+ during muscle contraction comes from an intracellular source. The Ca2+ is released from the lumen of the sarcoplasmic reticulum, which is a specialized region of endoplasmic reticulum inside a muscle cell. The signal from the nerve terminal triggers an action potential in the muscle cell plasma membrane, which causes a voltage-sensitive transmembrane protein in the membranous transverse tubules to open a Ca2+-release channel in the membrane of the sarcoplasmic reticulum.)

Do you agree or disagree with this statement? Explain your answer. Minus-end directed microtubule motors (like dyneins) deliver their cargo to the periphery of the cell, whereas plus-end directed microtubule motors (like kinesins) deliver their cargo to the interior of the cell.

Disagree (The plus ends of microtubules usually point toward the cell periphery, whereas the minus ends point toward the cell center. This is because the γ-tubulin in the centrosome serves to nucleate microtubule growth. Because the centrosomes are near the center of the cell, the minus ends of microtubules are located there. Therefore, a minusend directed microtubule motor would direct its cargo toward the center of the cell, and a plus-end directed microtubule motor would direct its cargo toward the cell periphery.)

All intermediate filaments are of similar diameter because ____________. (a) the central rod domains are similar in size and amino acid sequence. (b) the globular domains are similar in size and amino acid sequence. (c) covalent bonds among tetramers allow them to pack together in a similar fashion. (d) there is only a single type of intermediate filament in every organism.

a

Compared to the normal situation, in which actin monomers carry ATP, what do you predict would happen if actin monomers that bind a nonhydrolyzable form of ATP were incorporated into actin filaments? (a) Actin filaments would grow longer. (b) Actin filaments would grow shorter because depolymerization would be enhanced. (c) Actin filaments would grow shorter because new monomers could not be added to the filaments. (d) No change, as addition of monomers binding nonhydrolyzable ATP would not affect actin filament length.

a

The graph in Figure Q17-31 shows the time course of the polymerization of pure tubulin in vitro. Assume that the starting concentration of free tubulin is higher than it is in cells. Figure Q17-31 Three parts of the curve are labeled above it as A, B, and C. You conduct a similar in vitro tubulin-polymerization experiment, only you include purified centrosomes in your preparation. When you plot your data, which part of your graph should be most dissimilar to the curve shown in Figure Q17-31? (a) A (b) B (c) C (d) None. The shape of my graph should be identical to the graph produced when tubulin is polymerized in the absence of purified centrosomes

a

Which of the following items is not important for flagellar movement? (a) sarcoplasmic reticulum (b) ATP (c) dynein (d) microtubules

a

Which of the following statements about microtubules is true? (a) Motor proteins move in a directional fashion along microtubules by using the inherent structural polarity of a protofilament. (b) The centromere nucleates the microtubules of the mitotic spindle. (c) Because microtubules are subject to dynamic instability, they are used only for transient structures in a cell. (d) ATP hydrolysis by a tubulin heterodimer is important for controlling the growth of a microtubule.

a

Which of the following statements about the function of the centrosome is false? (a) Microtubules emanating from the centrosome have alternating polarity such that some have their plus end attached to the centrosome while others have their minus end attached to the centrosome. (b) Centrosomes contain hundreds of copies of the γ-tubulin ring complex important for microtubule nucleation. (c) Centrosomes typically contain a pair of centrioles, which is made up of a cylindrical array of short microtubules. (d) Centrosomes are the major microtubule-organizing center in animal cells.

a

Which of the situations below will enhance microtubule shrinkage? (a) addition of a drug that inhibits GTP exchange on free tubulin dimers (b) addition of a drug that inhibits hydrolysis of the GTP carried by tubulin dimers (c) addition of a drug that increases the affinity of tubulin molecules carrying GDP for other tubulin molecules (d) addition of a drug that blocks the ability of a tubulin dimer to bind to γ- tubulin

a

Which of the following statements is false? (a) Cytochalasins prevent actin polymerization. (b) Actin filaments are usually excluded from the cell cortex. (c) Integrins are transmembrane proteins that can bind to the extracellular matrix. (d) ARPs can promote the formation of branched actin filaments

b

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. A. monomer that binds ATP

actin

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. C. important for formation of the contractile ring during cytokinesis

actin

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. I. directly involved in muscle contraction

actin

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. J. abundant in filopodia

actin

You discover a protein, MtA, and find that it binds to the plus ends of microtubules in cells. The hypothesis that best explains this localization is ________________. (a) MtA is involved in stabilizing microtubules. (b) MtA binds to GTP-bound tubulin on microtubules. (c) MtA is important for the interaction of microtubules with the centrosome. (d) MtA will not bind to purified microtubules in a test tube.

b

Consider the mechanism by which actin and tubulin polymerize. Which of the items below does not describe something similar about the polymerization mechanisms of actin and microtubules? (a) Although both filaments can grow from both ends, the growth rate is faster at the plus ends. (b) Depolymerization initiates at the plus ends of filaments. (c) Nucleotide hydrolysis promotes depolymerization of filaments. (d) Free subunits (actin and tubulin) carry nucleoside triphosphates.

b

Figure Q17-57shows an electron micrograph of a skeletal muscle fiber, where various points along a fiber and various regions have been labeled. Figure Q17-57 Which of the following statements is true about muscle contraction? (a) Point A will move closer to point B. (b) Point B will move closer to point C. (c) Region D will become smaller. (d) Region E will shrink in size.

b

Intermediate filaments are made from elongated fibrous proteins that are assembled into a ropelike structure. Figure Q17-10 shows the structure of an intermediate filament subunit. You are interested in how intermediate filaments are formed, and you create an intermediate filament subunit whose α-helical region is twice as long as that of a normal intermediate filament by duplicating the normal α-helical region while keeping a globular head at the N-terminus and a globular tail at the C-terminus; you call this subunit IFαd. If you were to assemble intermediate filaments using IFαd as the subunit, which of the following predictions describes the most likely outcome? Figure Q17-10 (a) Filaments assembled using IFαd will interact with different cytoskeletal components. (b) Filaments assembled using IFαd will form dimers that are twice as long as dimers assembled from normal intermediate filaments. (c) Sixteen tetramers assembled from IFαd will be needed for a ropelike structure to form. (d) Dimers of IFαd will form by interactions with the N-terminal globular head and the C-terminal globular tail.

b

Intermediate filaments help protect animal cells from mechanical stress because ____________. (a) filaments directly extend from the interior of the cell to the extracellular space and into the next cell, linking one cell to the next, helping to distribute locally applied forces. (b) filaments in each cell are indirectly connected to the filaments of a neighboring cell through the desmosome, creating a continuous mechanical link between cells. (c) filaments remain independent of other cytoskeletal elements and keep the mechanical stress away from other cellular components. (d) filaments make up the desmosome junctions that connect cells; these junctions are more important than the internal network of filaments for protecting cells against mechanical stress.

b

Which of the following conditions is likely to decrease the likelihood of skeletal muscle contraction? (a) partial depolarization of the T-tubule membrane, such that the resting potential is closer to zero (b) addition of a drug that blocks Ca2+ binding to troponin (c) an increase in the amount of ATP in the cell (d) a mutation in tropomyosin that decreases its affinity for the actin filament

b

Which of the following statements about organellar movement in the cell is false? (a) Organelles undergo saltatory movement in the cell. (b) Only the microtubule cytoskeleton is involved in organellar movement. (c) Motor proteins involved in organellar movement use ATP hydrolysis for energy. (d) Organelles are attached to the tail domain of motor proteins.

b

Which of the following statements about skeletal muscle contraction is false? (a) When a muscle cell receives a signal from the nervous system, voltagegated channels open in the T-tubule membrane. (b) The changes in voltage across the plasma membrane that occur when a muscle cell receives a signal from the nervous system cause an influx of Ca2+ into the sarcoplasmic reticulum, triggering a muscle contraction. (c) A change in the conformation of troponin leads to changes in tropomyosin such that it no longer blocks the binding of myosin heads to the actin filament. (d) During muscle contraction, the Z discs move closer together as the myosin heads walk toward the plus ends of the actin filaments.

b

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. E. It uses dynein motors

both

Figure Q17-40A shows how the movement of dynein causes the flagellum to bend. If instead of the normal situation, the polarity of the adjacent doublet of microtubules were to be reversed (see Figure Q17-40B), what do you predict would happen? Figure Q17-40 (a) No bending would occur. (b) Bending would occur exactly as diagrammed in Figure Q17-40A. (c) Bending would occur, except that the right microtubule doublet would move down relative to the left one. (d) The two microtubule doublets would slide away from each other

c

Figure Q17-52 shows the leading edge of a lamellipodium. Which of the following statements is false? Figure Q17-52 (a) Nucleation of new filaments near the leading edge pushes the plasma membrane forward. (b) ARP proteins nucleate the branched actin filaments in the lamellipodium. (c) Capping proteins bind to the minus end of actin filaments. (d) There is more ATP-bound actin at the leading edge than in the actin filaments away from the leading edge.

c

The microtubules in a cell form a structural framework that can have all the following functions except which one? (a) holding internal organelles such as the Golgi apparatus in particular positions in the cell (b) creating long, thin cytoplasmic extensions that protrude from one side of the cell (c) strengthening the plasma membrane (d) moving materials from one place to another inside a cell

c

Which of the following statements about the cytoskeleton is false? (a) The cytoskeleton is made up of three types of protein filaments. (b) The cytoskeleton controls the location of organelles in eukaryotic cells. (c) Covalent bonds between protein monomers hold together cytoskeletal filaments. (d) The cytoskeleton of a cell can change in response to the environment.

c

Which of the following statements is correct? Kinesins and dyneins ____________________. (a) have tails that bind to the filaments. (b) move along both microtubules and actin filaments. (c) often move in opposite directions to each other. (d) derive their energy from GTP hydrolysis.

c

Which of the following structures shorten during muscle contraction? (a) myosin filaments (b) flagella (c) sarcomeres (d) actin filaments

c

You are examining a cell line in which activation of the Rho family member Rac promotes lamellipodia formation. Which of the following statements is most likely to be true? (a) Cells carrying a Rac mutation that makes Rac act as if it is always bound to GTP will polymerize more unbranched actin filaments than normal cells. (b) Cells carrying a Rac mutation that makes Rac unable to exchange GDP for GTP will polymerize more unbranched actin filaments than normal cells. (c) Cells carrying a Rac mutation that makes Rac act as if it is always bound to GTP will polymerize more branched actin filaments than normal cells. (d) Cells carrying a Rac mutation that makes Rac unable to exchange GDP for GTP will polymerize more branched actin filaments than normal cells.

c

You are studying nuclear lamins in yeast. Using recombinant DNA technology, you alter the coding sequence of a nuclear lamin gene such that the gene now codes for a nuclear lamin protein that can no longer be phosphorylated when the nuclear envelope is broken down during mitosis. What do you predict would happen if the yeast cell only had the altered nuclear lamin gene (and not the unaltered version)? (a) Mitosis should proceed as usual because the dephosphorylation of the lamin is what is important for nuclear lamina assembly during mitosis, so phosphorylation will not be necessary. (b) Disassembly of the nuclear lamins will occur prematurely because the lamins cannot be phosphorylated. (c) Nuclear lamins will no longer disassemble properly during mitosis. (d) Nuclear lamins will be unable to produce dimers, as coiled-coil formation will be disrupted

c

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. A. The basal body is the organizing center.

ciliary microtubules

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. C. It is arranged in a "9 + 2" array

ciliary microtubules

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. G. It is involved in moving fluid over the survace of cells

ciliary microtubules

Cell movement involves the coordination of many events in the cell. Which of the following phenomena is not required for cell motility? (a) Myosin-mediated contraction at the rear of the moving cell. (b) Integrin association with the extracellular environment. (c) Nucleation of new actin filaments. (d) Release of Ca2+ from the sarcoplasmic reticulum.

d

Consider the in vitro motility assay using purified kinesin and purified polymerized microtubules shown in Figure Q17-63. The three panels are images taken at 1-second intervals. In this figure, three microtubules have been numbered to make it easy to identify them. Which of the following statements about this assay is false? Figure Q17-63 (a) Kinesin molecules are attached by their tails to a glass slide. (b) The microtubules used in this assay must be polymerized using conditions that stabilize tubule formation or else they would undergo dynamic instability. (c) ATP must be added for this assay to work. (d) Addition of the nonhydrolyzable ATP analog (AMP-PNP) would cause the microtubules to move faster.

d

For both actin and microtubule polymerization, nucleotide hydrolysis is important for ______. (a) stabilizing the filaments once they are formed. (b) increasing the rate at which subunits are added to the filaments. (c) promoting nucleation of filaments. (d) decreasing the binding strength between subunits on filaments.

d

Keratins, neurofilaments, and vimentins are all categories of intermediate filaments. Which of the following properties is not true of these types of intermediate filaments? (a) They strengthen cells against mechanical stress. (b) Dimers associate by noncovalent bonding to form a tetramer. (c) They are found in the cytoplasm. (d) Phosphorylation causes disassembly during every mitotic cycle.

d

Microtubules are important for transporting cargo in nerve cell axons, as diagrammed in Figure Q17-33. Notice that the two types of cargo are traveling in opposite directions. Which of the following statements is likely to be false? Figure Q17-33 (a) The gray cargo is attached to dynein. (b) The black cargo and the gray cargo require ATP hydrolysis for their motion. (c) The black cargo moving toward the axon terminal contains a domain that specifically interacts with the tail domain of a particular kind of motor. (d) The black cargo and the gray cargo are moving along microtubules of opposite polarity.

d

The hydrolysis of GTP to GDP carried out by tubulin molecules ________________. (a) provides the energy needed for tubulin to polymerize. (b) occurs because the pool of free GDP has run out. (c) tips the balance in favor of microtubule assembly. (d) allows the behavior of microtubules called dynamic instability

d

Which of the following statements about actin is false? (a) ATP hydrolysis decreases actin filament stability. (b) Actin at the cell cortex helps govern the shape of the plasma membrane. (c) Actin filaments are nucleated at the side of existing actin filaments in lamellipodia. (d) The dynamic instability of actin filaments is important for cell movement.

d

Which of the following statements about the cytoskeleton is true? (a) All eukaryotic cells have actin, microtubules, and intermediate filaments in their cytoplasm. (b) The cytoskeleton provides a rigid and unchangeable structure important for the shape of the cell. (c) The three cytoskeletal filaments perform distinct tasks in the cell and act completely independently of one another. (d) Actin filaments and microtubules have an inherent polarity, with a plus end that grows more quickly than the minus end.

d

Which of the following statements about the structure of microtubules is false? (a) Microtubules are built from protofilaments that come together to make a hollow structure. (b) The two ends of a protofilament are chemically distinct, with α-tubulin exposed at one end and β-tubulin exposed at the other end. (c) Within a microtubule, all protofilaments are arranged in the same orientation, giving the microtubule structural polarity. (d) α-Tubulin and β-tubulin are covalently bound to make the tubulin dimer that then assembles into protofilaments

d

Which of the following statements regarding dynamic instability is false? (a) Each microtubule filament grows and shrinks independently of its neighbors. (b) The GTP cap helps protect a growing microtubule from depolymerization. (c) GTP hydrolysis by the tubulin dimer promotes microtubule shrinking. (d) The newly freed tubulin dimers from a shrinking microtubule can be immediately captured by growing microtubules and added to their plus end.

d

Which of the statements below about intermediate filaments is false? (a) They can stay intact in cells treated with concentrated salt solutions. (b) They can be found in the cytoplasm and the nucleus. (c) They can be anchored to the plasma membrane at a cell-cell junction. (d) Each filament is about 10 µm in diameter

d

You are interested in understanding the regulation of nuclear lamina assembly. To create an in vitro system for studying this process you start with partly purified nuclear lamina subunits to which you will add back purified cellular components to drive nuclear lamina assembly. Before you start doing experiments, your instructor suggests that you consider what type of conditions would be most amenable to the assembly of the nuclear lamina from its individual subunits in vitro. Which of the following conditions do you predict would be most likely to enhance the assembly of the nuclear lamina? (a) addition of phosphatase inhibitors (b) addition of ATP (c) addition of a concentrated salt solution that is 10 times the concentration normally found in the nucleoplasm (d) addition of protein kinase inhibitors

d

Your friend works in a biotech company that has just discovered a drug that seems to promote lamellipodia formation in cells. Which of the following molecules is unlikely to be involved in the pathway that this drug affects? (a) Rac (b) ARP (c) actin (d) myosin

d

Intermediate filaments are elongated fibrous proteins with an N-terminal globular _________________ region and a C-terminal globular _________________ region; these regions flank the elongated rod domain. The α-helical region of the rod interacts with the α-helical region of another monomer in a _____________________ configuration to form a dimer. ______________ dimers will line up to form a staggered tetramer. ______________ strands of tetramers come together and twist together to form the _________________ nm filament. The ___________________ domains are exposed on the surface of the intermediate filament, allowing for interaction with cytoplasmic components. antiparallel four tail β barrel globular ten coiled-coil head trimeric covalent rod twenty-five eight seven two

head, tail, coiled-coil, two, eight, ten, globular

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. B. includes keratin and neurofilaments

intermediate filaments

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. D. supports and strengthens the nuclear envelope

intermediate filaments

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. H. can be connected through desmosomes

intermediate filaments

Intermediate filaments are found mainly in cells that are subject to mechanical stress. Gene mutations that disrupt intermediate filaments cause some rare human genetic diseases. For example, the skin of people with epidermolysis bullosa simplex is very susceptible to mechanical injury; people with this disorder have mutations in their __________________ genes, which code for the intermediate filament found in epithelial cells. These filaments are usually connected from cell to cell through junctions called __________________s. The main filaments found in muscle cells belong to the __________________ family; people with disruptions in these intermediate filaments can have muscular dystrophy. In the nervous system, __________________s help strengthen the extremely long extensions often present in nerve cell axons; disruptions in these intermediate filaments can lead to neurodegeneration. People who carry mutations in the gene for __________________, an important protein for cross-linking intermediate filaments, have a disease that combines symptoms of epidermolysis bullosa simplex, muscular dystrophy, and neurodegeneration. Humans with progeria, a disease that causes premature aging, carry mutations in a nuclear ____________. desmosome lamin synapse keratin neurofilament vimentin kinase plectin

keratin, desmosomes, vimentin, neurofilaments, plectin, lamin

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. E. their stability involves a GTP cap

microtubules

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. F. used in the eukaryotic flagellum

microtubules

Indicate which of the three major classes of cytoskeletal elements each statement below refers to. G. a component of the mitotic spindle

microtubules

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. D. It is nucleated at the centrosome

microtubules of the mitotic spindle

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. B. The monomer is sequestered by profilin

neither

Indicate whether each of the following statements refers to a ciliary microtubule, a microtubule of the mitotic spindle, both types of microtubule, or neither type of microtubule. F. It is involved in sperm motility

neither