Essential Cell Biology 5th edition

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608 CHAPTER 17 CytoskeletonQUESTION 17–16When cells enter mitosis, their existing array of cytoplasmicmicrotubules has to be rapidly broken down and replacedwith the mitotic spindle that forms to pull the chromosomesinto the daughter cells. The enzyme katanin, named afterJapanese samurai swords, is activated during the onset ofmitosis, and chops microtubules into short pieces. Whatdo you suppose is the fate of the microtubule fragmentscreated by katanin? Explain your answer.QUESTION 17–17The drug Taxol, extracted from the bark of yew trees, hasan opposite effect to the drug colchicine, an alkaloid fromautumn crocus. Taxol binds tightly to microtubules andstabilizes them; when added to cells, it causes much of thefree tubulin to assemble into microtubules. In contrast,colchicine prevents microtubule formation. Taxol is justas pernicious to dividing cells as colchicine, and both areused as anticancer drugs. Based on your knowledge ofmicrotubule dynamics, suggest why both drugs are toxic todividing cells despite their opposite actions.QUESTION 17–18A useful technique for studying microtubule motors is toattach them by their tails to a glass cover slip (which canbe accomplished quite easily because the tails stick avidlyto a clean glass surface) and then allow them to settle.Microtubules may then be viewed in a light microscope asthey are propelled over the surface of the cover slip by theheads of the motor proteins. Because the motor proteinsattach at random orientations to the cover slip, however,how can they generate coordinated movement of individualmicrotubules rather than engaging in a tug-of-war? In whichdirection will microtubules crawl on a “bed” of kinesinmolecules (i.e., will they move plus-end first or minus-endfirst)?QUESTION 17–19A typical time course of polymerization of purified tubulin toform microtubules is shown in Figure Q17–19.A. Explain the different parts of the curve (labeled A, B,and C). Draw a diagram that shows the behavior of tubulinsubunits in each of the three phases.B. How would the curve in the figure change if centrosomeswere added at the outset?percentage of tubulin moleculesin microtubulesAFigure Q17–19BCtime at 37°C(A)Figure Q17–20QUESTION 17–20The electron micrographs shown in Figure Q17–20Awere obtained from a population of microtubules thatwere growing rapidly. Figure Q17–20B was obtainedfrom microtubules undergoing “catastrophic” shrinking.Comment on any differences between A and B, and suggestlikely explanations for the differences that you observe.QUESTION 17–21The locomotion of fibroblasts in culture is immediatelyECB5 eQ17.20/Q17.20halted by the drug cytochalasin, whereas colchicine causesfibroblasts to cease to move directionally and to beginextending lamellipodia in seemingly random directions.Injection of fibroblasts with antibodies to the intermediatefilament protein vimentin has no discernible effect on theirmigration. What do these observations suggest to youabout the involvement of the three different cytoskeletalfilaments in fibroblast locomotion?QUESTION 17–22(B)Complete the following sentence accurately, explaining yourreason for accepting or rejecting each of the four phrases(more than one can be correct). The role of calcium inmuscle contraction is:A. to detach myosin heads from actin.B. to spread the action potential from the plasmamembrane to the contractile machinery.C. to bind to troponin, cause it to move tropomyosin, andthereby expose actin filaments to myosin heads.D. to maintain the structure of the myosin filament.QUESTION 17–23Which of the following changes takes place when a skeletalmuscle contracts?A. Z discs move farther apart.B. Actin filaments contract.C. Myosin filaments contract.D. Sarcomeres become shorter.(Micrographs courtesy of Eva Mandelkow.)ECB5 eQ17.19/Q17.19

CHAPTER EIGHTEEN18The Cell-Division Cycle“Where a cell arises, there must be a previous cell, just as animals canonly arise from animals and plants from plants.” This statement, whichappears in a book written by German pathologist Rudolf Virchow in 1858,carries with it a profound message for the continuity of life. If every cellcomes from a previous cell, then all living organisms—from a unicellularbacterium to a multicellular mammal—are products of repeated roundsof cell growth and division, stretching back to the beginnings of life morethan 3 billion years ago.A cell reproduces by carrying out an orderly sequence of events in whichit duplicates its contents and then divides in two. This cycle of duplicationand division, known as the cell cycle, is the essential mechanismby which all living things reproduce. The details of the cell cycle varyfrom organism to organism and at different times in an individual organism’slife. In unicellular organisms, such as bacteria and yeasts, each celldivision produces a complete new organism, whereas many rounds ofcell division are required to make a new multicellular organism from afertilized egg. Certain features of the cell cycle, however, are universal,as they allow every cell to perform the fundamental task of copying andpassing on its genetic information to the next generation of cells.To explain how cells reproduce, we have to consider three majorquestions: (1) How do cells duplicate their contents—including the chromosomes,which carry the genetic information? (2) How do they partitionthe duplicated contents and split in two? (3) How do they coordinate allthe steps and machinery required for these two processes? The first questionis considered elsewhere in this book: in Chapter 6, we discuss howDNA is replicated, and in Chapters 7, 11, 15, and 17, we describe howOVERVIEW OF THE CELL CYCLETHE CELL-CYCLE CONTROLSYSTEMG 1 PHASES PHASEM PHASEMITOSISCYTOKINESISCONTROL OF CELL NUMBERSAND CELL SIZE

608 CHAPTER 17 Cytoskeleton

QUESTION 17–16

When cells enter mitosis, their existing array of cytoplasmic

microtubules has to be rapidly broken down and replaced

with the mitotic spindle that forms to pull the chromosomes

into the daughter cells. The enzyme katanin, named after

Japanese samurai swords, is activated during the onset of

mitosis, and chops microtubules into short pieces. What

do you suppose is the fate of the microtubule fragments

created by katanin? Explain your answer.

QUESTION 17–17

The drug Taxol, extracted from the bark of yew trees, has

an opposite effect to the drug colchicine, an alkaloid from

autumn crocus. Taxol binds tightly to microtubules and

stabilizes them; when added to cells, it causes much of the

free tubulin to assemble into microtubules. In contrast,

colchicine prevents microtubule formation. Taxol is just

as pernicious to dividing cells as colchicine, and both are

used as anticancer drugs. Based on your knowledge of

microtubule dynamics, suggest why both drugs are toxic to

dividing cells despite their opposite actions.

QUESTION 17–18

A useful technique for studying microtubule motors is to

attach them by their tails to a glass cover slip (which can

be accomplished quite easily because the tails stick avidly

to a clean glass surface) and then allow them to settle.

Microtubules may then be viewed in a light microscope as

they are propelled over the surface of the cover slip by the

heads of the motor proteins. Because the motor proteins

attach at random orientations to the cover slip, however,

how can they generate coordinated movement of individual

microtubules rather than engaging in a tug-of-war? In which

direction will microtubules crawl on a “bed” of kinesin

molecules (i.e., will they move plus-end first or minus-end

first)?

QUESTION 17–19

A typical time course of polymerization of purified tubulin to

form microtubules is shown in Figure Q17–19.

A. Explain the different parts of the curve (labeled A, B,

and C). Draw a diagram that shows the behavior of tubulin

subunits in each of the three phases.

B. How would the curve in the figure change if centrosomes

were added at the outset?

percentage of tubulin molecules

in microtubules

A

Figure Q17–19

B

C

time at 37°C

(A)

Figure Q17–20

QUESTION 17–20

The electron micrographs shown in Figure Q17–20A

were obtained from a population of microtubules that

were growing rapidly. Figure Q17–20B was obtained

from microtubules undergoing “catastrophic” shrinking.

Comment on any differences between A and B, and suggest

likely explanations for the differences that you observe.

QUESTION 17–21

The locomotion of fibroblasts in culture is immediately

ECB5 eQ17.20/Q17.20

halted by the drug cytochalasin, whereas colchicine causes

fibroblasts to cease to move directionally and to begin

extending lamellipodia in seemingly random directions.

Injection of fibroblasts with antibodies to the intermediate

filament protein vimentin has no discernible effect on their

migration. What do these observations suggest to you

about the involvement of the three different cytoskeletal

filaments in fibroblast locomotion?

QUESTION 17–22

(B)

Complete the following sentence accurately, explaining your

reason for accepting or rejecting each of the four phrases

(more than one can be correct). The role of calcium in

muscle contraction is:

A. to detach myosin heads from actin.

B. to spread the action potential from the plasma

membrane to the contractile machinery.

C. to bind to troponin, cause it to move tropomyosin, and

thereby expose actin filaments to myosin heads.

D. to maintain the structure of the myosin filament.

QUESTION 17–23

Which of the following changes takes place when a skeletal

muscle contracts?

A. Z discs move farther apart.

B. Actin filaments contract.

C. Myosin filaments contract.

D. Sarcomeres become shorter.

(Micrographs courtesy of Eva Mandelkow.)

ECB5 eQ17.19/Q17.19

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