Microscopic Anatomy and Contraction of Skeletal Muscle
We have already examined the structure of skeletal muscle as seen with the light microscope. As you know, skeletal muscle tissue has alternating light and dark bands, giving it a striated appearance. The electron microscope shows that these bands are due to the arrangement of myofilaments in a muscle fiber.
A muscle fiber contains the usual cellular components, but special names have been assigned to some of these components (Table 7.1 and Figure 7.3). The plasma membrane is called the sarcolemma; the cytoplasm is the sarcoplasm; and the endoplasmic reticulum is the sarcoplasmic reticulum. A muscle fiber also has some unique anatomical characteristics. One feature is its T (for transverse) system; the sarcolemma forms T (transverse) tubules that penetrate, or dip down, into the cell so that they come into contact-but do not fuse-with expanded portions of the sarcoplasmic reticulum. The expanded portions of the sarcoplasmic reticulum are calcium storage sites. Calcium ions (Ca2+), as we shall see, are essential for muscle contraction. The sarcoplasmic reticulum encases hundreds and sometimes even thousands of myofibrils, each about 1 µm in diameter, which are the contractile portions of the muscle fibers. Any other organelles, such as mitochondria, are located in the sarcoplasm between the myofibrils. The sarcoplasm also contains glycogen, which provides stored energy for muscle contraction, and the red pigment myoglobin, which binds oxygen until it is needed for muscle contraction.
Figure 7.3 Anatomy of a muscle fiber. A muscle fiber containsmany myofibrils with the components shown. A myofibril hasmany sarcomeres that contain myosin and actin filaments whose arrangement gives rise to the striations so characteristic of skeletal muscle. Muscle contraction occurs when sarcomeres contract and actin filaments slide past myosin filaments.
Myofibrils and Sarcomeres
Myofibrils are cylindrical in shape and run the length of the muscle fiber. The striations of skeletal muscle fibers are formed by the placement of myofilaments within units of myofibrils called sarcomeres. A sarcomere extends between two dark lines called the Z lines. A sarcomere contains two types of protein myofilaments. The thick filaments are made up of a protein called myosin, and the thin filaments are made up of a protein called actin. Other proteins are also present. The I band is light colored because it contains only actin filaments attached to a Z line. The dark regions of the A band contain overlapping actin and myosin filaments, and its H zone has only myosin filaments.
The thick and thin filaments differ in the following ways: Thick Filaments A thick filament is composed of several hundred molecules of the protein myosin. Each myosin molecule is shaped like a golf club, with the straight portion of the molecule ending in a double globular head, or crossbridge. Cross-bridges are slanted away from the middle of a sarcomere.
Thin Filaments Primarily, a thin filament consists of two intertwining strands of the protein actin. Two other proteins, called tropomyosin and troponin, are also present, as we will discuss later in this section.
Sliding Filaments We will also see that when muscles are innervated, impulses travel down a T tubule, and calcium is released from the sarcoplasmic reticulum. Now the muscle fiber contracts as the sarcomeres within the myofibrils shorten. When a sarcomere shortens, the actin (thin) filaments slide past the myosin (thick) filaments and approach one another. This causes the I band to shorten and the H zone to almost or completely disappear. The movement of actin filaments in relation to myosin filaments is called the sliding filament theory of muscle contraction. During the sliding process, the sarcomere shortens even though the filaments themselves remain the same length. ATP supplies the energy for muscle contraction. Although the actin filaments slide past the myosin filaments, it is the myosin filaments that do the work. Myosin filaments break down ATP and have crossbridges that pull the actin filaments toward the center of the sarcomere.
Skeletal Muscle Contraction
Muscle fibers are innervated-that is, they are stimulated to contract by motor neurons whose axons are found in nerves. The axon of one motor neuron has several branches and can stimulate from a few to several muscle fibers of a particular muscle. Each branch of the axon ends in an axon terminal that lies in close proximity to the sarcolemma of a muscle fiber. A small gap, called a synaptic cleft, separates the axon bulb from the sarcolemma. This entire region is called a neuromuscular junction (Fig. 7.4). Axon terminals contain synaptic vesicles that are filled with the neurotransmitter acetylcholine (ACh). When nerve impulses traveling down a motor neuron arrive at an axon terminal, the synaptic vesicles release a neurotransmitter into the synaptic cleft. It quickly diffuses across the cleft and binds to receptors in the sarcolemma. Now the sarcolemma generates impulses that spread over the sarcolemma and down T tubules to the sarcoplasmic reticulum. The release of calcium from the sarcoplasmic reticulum causes the filaments within the sarcomeres to slide past one another. Sarcomere contraction results in myofibril contraction, which in turn results in muscle fiber, and finally muscle, contraction.
Figure 7.4 Neuromuscular junction. The branch of an axon ends in an axon terminal that meets but does not touch a muscle fiber. A synaptic cleft separates the axon terminal from the sarcolemma of the muscle fiber. Nerve impulses traveling down an axon cause synaptic vesicles to discharge acetylcholine, which diffuses across the synaptic cleft. When the neurotransmitter is received by the sarcolemma of a muscle fiber, impulses begin and lead to muscle fiber contractions.