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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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892 Chapter 16: The Cytoskeleton

Figure 16–2 Diagram of changes in

cytoskeletal organization associated

with cell division. The crawling fibroblast

drawn here has a polarized, dynamic

actin cytoskeleton (shown in red) that

assembles lamellipodia and filopodia to

push its leading edge toward the right. The

polarization of the actin cytoskeleton is

assisted by the microtubule cytoskeleton

(green), consisting of long microtubules

that emanate from a single microtubuleorganizing

center located in front of

the nucleus. When the cell divides, the

polarized microtubule array rearranges

to form a bipolar mitotic spindle, which

is responsible for aligning and then

segregating the duplicated chromosomes

(brown). The actin filaments form a

contractile ring at the center of the cell

that pinches the cell in two after the

chromosome segregation. After cell

division is complete, the two daughter

cells reorganize both the microtubule and

actin cytoskeletons into smaller versions of

those that were present in the mother cell,

enabling them to crawl their separate ways.

blood cell, chases and engulfs bacterial and fungal cells that accidentally gain

access to the normally sterile parts of the body, as through a cut in the skin. Like

most crawling cells, neutrophils advance by extending a protrusive structure filled

with newly polymerized actin filaments. When the elusive bacterial prey moves in

a different direction, the neutrophil is poised to reorganize its polarized protrusive

structures within seconds (Figure 16–3).

The Cytoskeleton Determines Cellular Organization and Polarity

MBoC6 m16.02/16.02

In cells that have achieved a stable, differentiated morphology—such as mature

neurons or epithelial cells—the dynamic elements of the cytoskeleton must also

provide stable, large-scale structures for cellular organization. On specialized epithelial

cells that line organs such as the intestine and the lung, cytoskeletal-based

cell-surface protrusions including microvilli and cilia are able to maintain a constant

location, length, and diameter over the entire lifetime of the cell. For the

actin bundles at the cores of microvilli on intestinal epithelial cells, this is only a

few days. But the actin bundles at the cores of stereocilia on the hair cells of the

inner ear must maintain their stable organization for the entire lifetime of the animal,

since these cells do not turn over. Nonetheless, the individual actin filaments

time 0 min 1 min 2 min 3 min

Figure 16–3 A neutrophil in pursuit of

bacteria. In this preparation of human

blood, a clump of bacteria (white arrow) is

about to be captured by a neutrophil. As

the bacteria move, the neutrophil quickly

reassembles the dense actin network at

its leading edge (highlighted in red) to

push toward the location of the bacteria

(Movie 16.1). Rapid disassembly and

reassembly of the actin cytoskeleton in

this cell enables it to change its orientation

and direction of movement within a few

minutes. (From a video recorded by

David Rogers.)

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