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actin filaments and tubulin microtubules

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994 PANEL 16–3: Accessory Proteins that Control the Assembly <strong>and</strong> Position of Cytoskeletal Filaments<br />

ACTIN FILAMENTS<br />

– +<br />

cofilin<br />

binds ADP-<strong>actin</strong> <strong>filaments</strong>,<br />

accelerates disassembly<br />

fimbrin<br />

–<br />

+<br />

formin<br />

nucleates assembly <strong>and</strong> remains<br />

associated with the growing<br />

plus end<br />

thymosin<br />

binds subunits,<br />

prevents assembly<br />

–<br />

–<br />

–<br />

+<br />

+<br />

gelsolin<br />

severs <strong>filaments</strong> <strong>and</strong><br />

binds to plus end<br />

+<br />

<strong>actin</strong> subunits<br />

<strong>actin</strong> filament<br />

a-<strong>actin</strong>in filamin spectrin<br />

ARP complex<br />

nucleates assembly<br />

to form a web <strong>and</strong> remains<br />

associated with the minus end<br />

– +<br />

capping protein<br />

prevents assembly <strong>and</strong><br />

disassembly at plus end<br />

filament bundling, cross-linking, <strong>and</strong> attachment to membranes<br />

–<br />

+<br />

–<br />

–<br />

+<br />

–<br />

–<br />

profilin<br />

binds subunits,<br />

speeds elongation<br />

plasma<br />

membrane<br />

+<br />

+<br />

– +<br />

+<br />

–<br />

tropomyosin<br />

stabilizes filament<br />

Some of the major accessory proteins of the <strong>actin</strong> cytoskeleton. Except for the myosin motor proteins, to be discussed in<br />

a later section, an example of each major type is shown. Each of these is discussed in the text. However, most cells contain<br />

more than a hundred different <strong>actin</strong>-binding proteins, <strong>and</strong> it is likely that there are important types of <strong>actin</strong>-associated<br />

proteins that are not yet recognized.<br />

ERM<br />

+


MICROTUBULES<br />

stathmin<br />

binds subunits,<br />

prevents assembly<br />

– +<br />

kinesin 13<br />

enhances catastrophic<br />

disassembly at plus end<br />

–<br />

+<br />

– +<br />

– +<br />

katanin<br />

severs <strong>microtubules</strong><br />

g-TuRC<br />

nucleates assembly <strong>and</strong><br />

remains associated with<br />

the minus end<br />

ab <strong>tubulin</strong> dimers<br />

microtubule<br />

filament bundling <strong>and</strong> cross-linking<br />

centrosome<br />

MAPs<br />

stabilize tubules by<br />

binding along sides<br />

– +<br />

tau MAP-2 plectin<br />

+TIPs<br />

remain associated with growing plus ends<br />

<strong>and</strong> can link them to other structures,<br />

such as membranes<br />

– +<br />

XMAP215<br />

stabilizes plus ends<br />

<strong>and</strong> accelerates assembly<br />

links to intermediate <strong>filaments</strong><br />

Some of the major accessory proteins of the microtubule cytoskeleton. Except for two classes of motor proteins, to be<br />

discussed in a later section, an example of each major type is shown. Each of these is discussed in the text. However,<br />

most cells contain more than a hundred different microtubule-binding proteins, <strong>and</strong>—as for the <strong>actin</strong>-associated<br />

proteins—it is likely that there are important types of microtubule-associated proteins that are not yet recognized.<br />

995

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