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Essential Cell Biology 5th edition

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Muscle Contraction

601

plus end

minus end

minus end

myosin-ll

plus end

The myosin filament is like a double-headed arrow, with the two sets of

myosin heads pointing outward, away from the middle. One set binds to

actin filaments in one orientation and moves the filaments one way; the

other set binds to other actin filaments in the opposite orientation and

moves the filaments in the opposite direction. As a result, a myosin filament

slides sets of oppositely oriented actin filaments past one another

ECB5 E17.39/17.39

(Figure 17–39). Thus, if actin filaments and myosin filaments are organized

together in a bundle, the bundle can generate a strong contractile

force. This is seen most clearly in muscle contraction, but it also occurs in

the much smaller contractile bundles of actin filaments and myosin-II filaments

(see Figure 17−29B) that assemble transiently in nonmuscle cells,

and in the contractile ring that pinches a dividing cell in two by contracting

and pulling inward on the plasma membrane (see Figure 17–29D).

Actin Filaments Slide Against Myosin Filaments During

Muscle Contraction

In most animals, skeletal muscle fibers are huge, multinucleated individual

cells formed by the fusion of many separate smaller cells. The

nuclei of the contributing cells are retained in the muscle fiber and lie just

beneath the plasma membrane. The bulk of the cytoplasm is made up of

myofibrils, the contractile elements of the muscle cell. These cylindrical

structures are 1–2 μm in diameter and may be as long as the muscle cell

itself (Figure 17–40A).

A myofibril consists of a chain of identical tiny contractile units, or

sarcomeres. Each sarcomere is about 2.5 μm long, and the repeating

pattern of sarcomeres gives the vertebrate myofibril a striped appearance

(Figure 17–40B). Sarcomeres are highly organized assemblies of

two types of filaments—actin filaments and myosin filaments composed

Figure 17–39 A small, bipolar myosin-II

filament can slide two actin filaments

of opposite orientation past each

other. Similar sliding movement mediates

the contraction of interacting actin and

myosin-II filaments in both muscle and

nonmuscle cells. As with myosin-I, a

myosin-II head group walks toward the

plus end of the actin filament with which

it interacts. Note that multiple myosin

molecules are required to generate

movement: when one myosin head releases

the filament to reposition itself, other

myosins must remain attached so the

structure does not fall apart.

QUESTION 17–8

If both the actin and myosin

filaments of muscle are made

up of subunits held together by

weak noncovalent bonds, how is it

possible for a human being to lift

heavy objects?

(A)

(B)

nucleus plasma membrane myofibril

Figure 17–40 A skeletal muscle cell is packed with myofibrils.

(A) In an adult human, these huge, multinucleated cells (also

called muscle fibers) are typically 50 μm in diameter, and they can

be several centimeters long. They contain numerous myofibrils,

in which actin filaments and myosin-II filaments are arranged in

a highly organized structure, giving each myofibril—and skeletal

muscle cell—a striated or striped appearance; for this reason,

skeletal muscle is also called striated muscle. (B) Low-magnification

electron micrograph of a longitudinal section through a skeletal

muscle cell of a rabbit, showing that each myofibril consists of

a repeating chain of sarcomeres, the contractile units of the

myofibrils. (B, courtesy of Roger Craig.)

sarcomere ~2.5 µm

sarcomere

two myofibrils

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