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Anatomy and Physiology

10.2 Skeletal Muscle

Anatomy and Physiology10.2 Skeletal Muscle
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  1. Preface
  2. Unit 1: Levels of Organization
    1. 1 An Introduction to the Human Body
      1. Introduction
      2. 1.1 Overview of Anatomy and Physiology
      3. 1.2 Structural Organization of the Human Body
      4. 1.3 Functions of Human Life
      5. 1.4 Requirements for Human Life
      6. 1.5 Homeostasis
      7. 1.6 Anatomical Terminology
      8. 1.7 Medical Imaging
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
    2. 2 The Chemical Level of Organization
      1. Introduction
      2. 2.1 Elements and Atoms: The Building Blocks of Matter
      3. 2.2 Chemical Bonds
      4. 2.3 Chemical Reactions
      5. 2.4 Inorganic Compounds Essential to Human Functioning
      6. 2.5 Organic Compounds Essential to Human Functioning
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    3. 3 The Cellular Level of Organization
      1. Introduction
      2. 3.1 The Cell Membrane
      3. 3.2 The Cytoplasm and Cellular Organelles
      4. 3.3 The Nucleus and DNA Replication
      5. 3.4 Protein Synthesis
      6. 3.5 Cell Growth and Division
      7. 3.6 Cellular Differentiation
      8. Key Terms
      9. Chapter Review
      10. Interactive Link Questions
      11. Review Questions
      12. Critical Thinking Questions
    4. 4 The Tissue Level of Organization
      1. Introduction
      2. 4.1 Types of Tissues
      3. 4.2 Epithelial Tissue
      4. 4.3 Connective Tissue Supports and Protects
      5. 4.4 Muscle Tissue and Motion
      6. 4.5 Nervous Tissue Mediates Perception and Response
      7. 4.6 Tissue Injury and Aging
      8. Key Terms
      9. Chapter Review
      10. Interactive Link Questions
      11. Review Questions
      12. Critical Thinking Questions
  3. Unit 2: Support and Movement
    1. 5 The Integumentary System
      1. Introduction
      2. 5.1 Layers of the Skin
      3. 5.2 Accessory Structures of the Skin
      4. 5.3 Functions of the Integumentary System
      5. 5.4 Diseases, Disorders, and Injuries of the Integumentary System
      6. Key Terms
      7. Chapter Review
      8. Interactive Link Questions
      9. Review Questions
      10. Critical Thinking Questions
    2. 6 Bone Tissue and the Skeletal System
      1. Introduction
      2. 6.1 The Functions of the Skeletal System
      3. 6.2 Bone Classification
      4. 6.3 Bone Structure
      5. 6.4 Bone Formation and Development
      6. 6.5 Fractures: Bone Repair
      7. 6.6 Exercise, Nutrition, Hormones, and Bone Tissue
      8. 6.7 Calcium Homeostasis: Interactions of the Skeletal System and Other Organ Systems
      9. Key Terms
      10. Chapter Review
      11. Review Questions
      12. Critical Thinking Questions
    3. 7 Axial Skeleton
      1. Introduction
      2. 7.1 Divisions of the Skeletal System
      3. 7.2 The Skull
      4. 7.3 The Vertebral Column
      5. 7.4 The Thoracic Cage
      6. 7.5 Embryonic Development of the Axial Skeleton
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    4. 8 The Appendicular Skeleton
      1. Introduction
      2. 8.1 The Pectoral Girdle
      3. 8.2 Bones of the Upper Limb
      4. 8.3 The Pelvic Girdle and Pelvis
      5. 8.4 Bones of the Lower Limb
      6. 8.5 Development of the Appendicular Skeleton
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    5. 9 Joints
      1. Introduction
      2. 9.1 Classification of Joints
      3. 9.2 Fibrous Joints
      4. 9.3 Cartilaginous Joints
      5. 9.4 Synovial Joints
      6. 9.5 Types of Body Movements
      7. 9.6 Anatomy of Selected Synovial Joints
      8. 9.7 Development of Joints
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
    6. 10 Muscle Tissue
      1. Introduction
      2. 10.1 Overview of Muscle Tissues
      3. 10.2 Skeletal Muscle
      4. 10.3 Muscle Fiber Contraction and Relaxation
      5. 10.4 Nervous System Control of Muscle Tension
      6. 10.5 Types of Muscle Fibers
      7. 10.6 Exercise and Muscle Performance
      8. 10.7 Cardiac Muscle Tissue
      9. 10.8 Smooth Muscle
      10. 10.9 Development and Regeneration of Muscle Tissue
      11. Key Terms
      12. Chapter Review
      13. Interactive Link Questions
      14. Review Questions
      15. Critical Thinking Questions
    7. 11 The Muscular System
      1. Introduction
      2. 11.1 Interactions of Skeletal Muscles, Their Fascicle Arrangement, and Their Lever Systems
      3. 11.2 Naming Skeletal Muscles
      4. 11.3 Axial Muscles of the Head, Neck, and Back
      5. 11.4 Axial Muscles of the Abdominal Wall, and Thorax
      6. 11.5 Muscles of the Pectoral Girdle and Upper Limbs
      7. 11.6 Appendicular Muscles of the Pelvic Girdle and Lower Limbs
      8. Key Terms
      9. Chapter Review
      10. Review Questions
      11. Critical Thinking Questions
  4. Unit 3: Regulation, Integration, and Control
    1. 12 The Nervous System and Nervous Tissue
      1. Introduction
      2. 12.1 Basic Structure and Function of the Nervous System
      3. 12.2 Nervous Tissue
      4. 12.3 The Function of Nervous Tissue
      5. 12.4 The Action Potential
      6. 12.5 Communication Between Neurons
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    2. 13 Anatomy of the Nervous System
      1. Introduction
      2. 13.1 The Embryologic Perspective
      3. 13.2 The Central Nervous System
      4. 13.3 Circulation and the Central Nervous System
      5. 13.4 The Peripheral Nervous System
      6. Key Terms
      7. Chapter Review
      8. Interactive Link Questions
      9. Review Questions
      10. Critical Thinking Questions
    3. 14 The Somatic Nervous System
      1. Introduction
      2. 14.1 Sensory Perception
      3. 14.2 Central Processing
      4. 14.3 Motor Responses
      5. Key Terms
      6. Chapter Review
      7. Interactive Link Questions
      8. Review Questions
      9. Critical Thinking Questions
    4. 15 The Autonomic Nervous System
      1. Introduction
      2. 15.1 Divisions of the Autonomic Nervous System
      3. 15.2 Autonomic Reflexes and Homeostasis
      4. 15.3 Central Control
      5. 15.4 Drugs that Affect the Autonomic System
      6. Key Terms
      7. Chapter Review
      8. Interactive Link Questions
      9. Review Questions
      10. Critical Thinking Questions
    5. 16 The Neurological Exam
      1. Introduction
      2. 16.1 Overview of the Neurological Exam
      3. 16.2 The Mental Status Exam
      4. 16.3 The Cranial Nerve Exam
      5. 16.4 The Sensory and Motor Exams
      6. 16.5 The Coordination and Gait Exams
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    6. 17 The Endocrine System
      1. Introduction
      2. 17.1 An Overview of the Endocrine System
      3. 17.2 Hormones
      4. 17.3 The Pituitary Gland and Hypothalamus
      5. 17.4 The Thyroid Gland
      6. 17.5 The Parathyroid Glands
      7. 17.6 The Adrenal Glands
      8. 17.7 The Pineal Gland
      9. 17.8 Gonadal and Placental Hormones
      10. 17.9 The Endocrine Pancreas
      11. 17.10 Organs with Secondary Endocrine Functions
      12. 17.11 Development and Aging of the Endocrine System
      13. Key Terms
      14. Chapter Review
      15. Interactive Link Questions
      16. Review Questions
      17. Critical Thinking Questions
  5. Unit 4: Fluids and Transport
    1. 18 The Cardiovascular System: Blood
      1. Introduction
      2. 18.1 An Overview of Blood
      3. 18.2 Production of the Formed Elements
      4. 18.3 Erythrocytes
      5. 18.4 Leukocytes and Platelets
      6. 18.5 Hemostasis
      7. 18.6 Blood Typing
      8. Key Terms
      9. Chapter Review
      10. Interactive Link Questions
      11. Review Questions
      12. Critical Thinking Questions
    2. 19 The Cardiovascular System: The Heart
      1. Introduction
      2. 19.1 Heart Anatomy
      3. 19.2 Cardiac Muscle and Electrical Activity
      4. 19.3 Cardiac Cycle
      5. 19.4 Cardiac Physiology
      6. 19.5 Development of the Heart
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
    3. 20 The Cardiovascular System: Blood Vessels and Circulation
      1. Introduction
      2. 20.1 Structure and Function of Blood Vessels
      3. 20.2 Blood Flow, Blood Pressure, and Resistance
      4. 20.3 Capillary Exchange
      5. 20.4 Homeostatic Regulation of the Vascular System
      6. 20.5 Circulatory Pathways
      7. 20.6 Development of Blood Vessels and Fetal Circulation
      8. Key Terms
      9. Chapter Review
      10. Interactive Link Questions
      11. Review Questions
      12. Critical Thinking Questions
    4. 21 The Lymphatic and Immune System
      1. Introduction
      2. 21.1 Anatomy of the Lymphatic and Immune Systems
      3. 21.2 Barrier Defenses and the Innate Immune Response
      4. 21.3 The Adaptive Immune Response: T lymphocytes and Their Functional Types
      5. 21.4 The Adaptive Immune Response: B-lymphocytes and Antibodies
      6. 21.5 The Immune Response against Pathogens
      7. 21.6 Diseases Associated with Depressed or Overactive Immune Responses
      8. 21.7 Transplantation and Cancer Immunology
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
  6. Unit 5: Energy, Maintenance, and Environmental Exchange
    1. 22 The Respiratory System
      1. Introduction
      2. 22.1 Organs and Structures of the Respiratory System
      3. 22.2 The Lungs
      4. 22.3 The Process of Breathing
      5. 22.4 Gas Exchange
      6. 22.5 Transport of Gases
      7. 22.6 Modifications in Respiratory Functions
      8. 22.7 Embryonic Development of the Respiratory System
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
    2. 23 The Digestive System
      1. Introduction
      2. 23.1 Overview of the Digestive System
      3. 23.2 Digestive System Processes and Regulation
      4. 23.3 The Mouth, Pharynx, and Esophagus
      5. 23.4 The Stomach
      6. 23.5 The Small and Large Intestines
      7. 23.6 Accessory Organs in Digestion: The Liver, Pancreas, and Gallbladder
      8. 23.7 Chemical Digestion and Absorption: A Closer Look
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
    3. 24 Metabolism and Nutrition
      1. Introduction
      2. 24.1 Overview of Metabolic Reactions
      3. 24.2 Carbohydrate Metabolism
      4. 24.3 Lipid Metabolism
      5. 24.4 Protein Metabolism
      6. 24.5 Metabolic States of the Body
      7. 24.6 Energy and Heat Balance
      8. 24.7 Nutrition and Diet
      9. Key Terms
      10. Chapter Review
      11. Review Questions
      12. Critical Thinking Questions
    4. 25 The Urinary System
      1. Introduction
      2. 25.1 Physical Characteristics of Urine
      3. 25.2 Gross Anatomy of Urine Transport
      4. 25.3 Gross Anatomy of the Kidney
      5. 25.4 Microscopic Anatomy of the Kidney
      6. 25.5 Physiology of Urine Formation
      7. 25.6 Tubular Reabsorption
      8. 25.7 Regulation of Renal Blood Flow
      9. 25.8 Endocrine Regulation of Kidney Function
      10. 25.9 Regulation of Fluid Volume and Composition
      11. 25.10 The Urinary System and Homeostasis
      12. Key Terms
      13. Chapter Review
      14. Review Questions
      15. Critical Thinking Questions
    5. 26 Fluid, Electrolyte, and Acid-Base Balance
      1. Introduction
      2. 26.1 Body Fluids and Fluid Compartments
      3. 26.2 Water Balance
      4. 26.3 Electrolyte Balance
      5. 26.4 Acid-Base Balance
      6. 26.5 Disorders of Acid-Base Balance
      7. Key Terms
      8. Chapter Review
      9. Interactive Link Questions
      10. Review Questions
      11. Critical Thinking Questions
  7. Unit 6: Human Development and the Continuity of Life
    1. 27 The Reproductive System
      1. Introduction
      2. 27.1 Anatomy and Physiology of the Male Reproductive System
      3. 27.2 Anatomy and Physiology of the Female Reproductive System
      4. 27.3 Development of the Male and Female Reproductive Systems
      5. Key Terms
      6. Chapter Review
      7. Interactive Link Questions
      8. Review Questions
      9. Critical Thinking Questions
    2. 28 Development and Inheritance
      1. Introduction
      2. 28.1 Fertilization
      3. 28.2 Embryonic Development
      4. 28.3 Fetal Development
      5. 28.4 Maternal Changes During Pregnancy, Labor, and Birth
      6. 28.5 Adjustments of the Infant at Birth and Postnatal Stages
      7. 28.6 Lactation
      8. 28.7 Patterns of Inheritance
      9. Key Terms
      10. Chapter Review
      11. Interactive Link Questions
      12. Review Questions
      13. Critical Thinking Questions
  8. References
  9. Index
By the end of this section, you will be able to:
  • Describe the layers of connective tissues packaging skeletal muscle
  • Explain how muscles work with tendons to move the body
  • Identify areas of the skeletal muscle fibers
  • Describe excitation-contraction coupling

The best-known feature of skeletal muscle is its ability to contract and cause movement. Skeletal muscles act not only to produce movement but also to stop movement, such as resisting gravity to maintain posture. Small, constant adjustments of the skeletal muscles are needed to hold a body upright or balanced in any position. Muscles also prevent excess movement of the bones and joints, maintaining skeletal stability and preventing skeletal structure damage or deformation. Joints can become misaligned or dislocated entirely by pulling on the associated bones; muscles work to keep joints stable. Skeletal muscles are located throughout the body at the openings of internal tracts to control the movement of various substances. These muscles allow functions, such as swallowing, urination, and defecation, to be under voluntary control. Skeletal muscles also protect internal organs (particularly abdominal and pelvic organs) by acting as an external barrier or shield to external trauma and by supporting the weight of the organs.

Skeletal muscles contribute to the maintenance of homeostasis in the body by generating heat. Muscle contraction requires energy, and when ATP is broken down, heat is produced. This heat is very noticeable during exercise, when sustained muscle movement causes body temperature to rise, and in cases of extreme cold, when shivering produces random skeletal muscle contractions to generate heat.

Each skeletal muscle is an organ that consists of various integrated tissues. These tissues include the skeletal muscle fibers, blood vessels, nerve fibers, and connective tissue. Each skeletal muscle has three layers of connective tissue (called “mysia”) that enclose it and provide structure to the muscle as a whole, and also compartmentalize the muscle fibers within the muscle (Figure 10.3). Each muscle is wrapped in a sheath of dense, irregular connective tissue called the epimysium, which allows a muscle to contract and move powerfully while maintaining its structural integrity. The epimysium also separates muscle from other tissues and organs in the area, allowing the muscle to move independently.

This figure shows the structure of muscle fibers. The top panel shows a skeleton muscle fiber, and a magnified view of the muscle fascicles are shown. The middle panel shows a magnified view of the muscle fascicles with the muscle fibers, perimysium and the endomysium. The bottom panel shows the structure of the muscle fiber with the sarcolemma highlighted.
Figure 10.3 The Three Connective Tissue Layers Bundles of muscle fibers, called fascicles, are covered by the perimysium. Muscle fibers are covered by the endomysium.

Inside each skeletal muscle, muscle fibers are organized into individual bundles, each called a fascicle, by a middle layer of connective tissue called the perimysium. This fascicular organization is common in muscles of the limbs; it allows the nervous system to trigger a specific movement of a muscle by activating a subset of muscle fibers within a bundle, or fascicle of the muscle. Inside each fascicle, each muscle fiber is encased in a thin connective tissue layer of collagen and reticular fibers called the endomysium. The endomysium contains the extracellular fluid and nutrients to support the muscle fiber. These nutrients are supplied via blood to the muscle tissue.

In skeletal muscles that work with tendons to pull on bones, the collagen in the three tissue layers (the mysia) intertwines with the collagen of a tendon. At the other end of the tendon, it fuses with the periosteum coating the bone. The tension created by contraction of the muscle fibers is then transferred though the mysia, to the tendon, and then to the periosteum to pull on the bone for movement of the skeleton. In other places, the mysia may fuse with a broad, tendon-like sheet called an aponeurosis, or to fascia, the connective tissue between skin and bones. The broad sheet of connective tissue in the lower back that the latissimus dorsi muscles (the “lats”) fuse into is an example of an aponeurosis.

Every skeletal muscle is also richly supplied by blood vessels for nourishment, oxygen delivery, and waste removal. In addition, every muscle fiber in a skeletal muscle is supplied by the axon branch of a somatic motor neuron, which signals the fiber to contract. Unlike cardiac and smooth muscle, the only way to functionally contract a skeletal muscle is through signaling from the nervous system.

Skeletal Muscle Fibers

Because skeletal muscle cells are long and cylindrical, they are commonly referred to as muscle fibers. Skeletal muscle fibers can be quite large for human cells, with diameters up to 100 μm and lengths up to 30 cm (11.8 in) in the Sartorius of the upper leg. During early development, embryonic myoblasts, each with its own nucleus, fuse with up to hundreds of other myoblasts to form the multinucleated skeletal muscle fibers. Multiple nuclei mean multiple copies of genes, permitting the production of the large amounts of proteins and enzymes needed for muscle contraction.

Some other terminology associated with muscle fibers is rooted in the Greek sarco, which means “flesh.” The plasma membrane of muscle fibers is called the sarcolemma, the cytoplasm is referred to as sarcoplasm, and the specialized smooth endoplasmic reticulum, which stores, releases, and retrieves calcium ions (Ca++) is called the sarcoplasmic reticulum (SR) (Figure 10.4). As will soon be described, the functional unit of a skeletal muscle fiber is the sarcomere, a highly organized arrangement of the contractile myofilaments actin (thin filament) and myosin (thick filament), along with other support proteins.

This figure shows the structure of the muscle fibers. In the top panel, a sarcolemma is shown with the major parts labeled. In the bottom panel, a magnified view of a single myofibril is shown and the major parts are labeled.
Figure 10.4 Muscle Fiber A skeletal muscle fiber is surrounded by a plasma membrane called the sarcolemma, which contains sarcoplasm, the cytoplasm of muscle cells. A muscle fiber is composed of many fibrils, which give the cell its striated appearance.

The Sarcomere

The striated appearance of skeletal muscle fibers is due to the arrangement of the myofilaments of actin and myosin in sequential order from one end of the muscle fiber to the other. Each packet of these microfilaments and their regulatory proteins, troponin and tropomyosin (along with other proteins) is called a sarcomere.

Interactive Link

Watch this video to learn more about macro- and microstructures of skeletal muscles. (a) What are the names of the “junction points” between sarcomeres? (b) What are the names of the “subunits” within the myofibrils that run the length of skeletal muscle fibers? (c) What is the “double strand of pearls” described in the video? (d) What gives a skeletal muscle fiber its striated appearance?

The sarcomere is the functional unit of the muscle fiber. The sarcomere itself is bundled within the myofibril that runs the entire length of the muscle fiber and attaches to the sarcolemma at its end. As myofibrils contract, the entire muscle cell contracts. Because myofibrils are only approximately 1.2 μm in diameter, hundreds to thousands (each with thousands of sarcomeres) can be found inside one muscle fiber. Each sarcomere is approximately 2 μm in length with a three-dimensional cylinder-like arrangement and is bordered by structures called Z-discs (also called Z-lines, because pictures are two-dimensional), to which the actin myofilaments are anchored (Figure 10.5). Because the actin and its troponin-tropomyosin complex (projecting from the Z-discs toward the center of the sarcomere) form strands that are thinner than the myosin, it is called the thin filament of the sarcomere. Likewise, because the myosin strands and their multiple heads (projecting from the center of the sarcomere, toward but not all to way to, the Z-discs) have more mass and are thicker, they are called the thick filament of the sarcomere.

This figure shows the structure of thick and thin filaments. On the top of the image a sarcomere is shown with the H zone, Z line and M lines labeled. To the right of the bottom panel, the structure of the thick filament is shown in detail. To the left of the bottom panel, the structure of a thin filament is shown in detail.
Figure 10.5 The Sarcomere The sarcomere, the region from one Z-line to the next Z-line, is the functional unit of a skeletal muscle fiber.

The Neuromuscular Junction

Another specialization of the skeletal muscle is the site where a motor neuron’s terminal meets the muscle fiber—called the neuromuscular junction (NMJ). This is where the muscle fiber first responds to signaling by the motor neuron. Every skeletal muscle fiber in every skeletal muscle is innervated by a motor neuron at the NMJ. Excitation signals from the neuron are the only way to functionally activate the fiber to contract.

Interactive Link

Every skeletal muscle fiber is supplied by a motor neuron at the NMJ. Watch this video to learn more about what happens at the NMJ. (a) What is the definition of a motor unit? (b) What is the structural and functional difference between a large motor unit and a small motor unit? (c) Can you give an example of each? (d) Why is the neurotransmitter acetylcholine degraded after binding to its receptor?

Excitation-Contraction Coupling

All living cells have membrane potentials, or electrical gradients across their membranes. The inside of the membrane is usually around -60 to -90 mV, relative to the outside. This is referred to as a cell’s membrane potential. Neurons and muscle cells can use their membrane potentials to generate electrical signals. They do this by controlling the movement of charged particles, called ions, across their membranes to create electrical currents. This is achieved by opening and closing specialized proteins in the membrane called ion channels. Although the currents generated by ions moving through these channel proteins are very small, they form the basis of both neural signaling and muscle contraction.

Both neurons and skeletal muscle cells are electrically excitable, meaning that they are able to generate action potentials. An action potential is a special type of electrical signal that can travel along a cell membrane as a wave. This allows a signal to be transmitted quickly and faithfully over long distances.

Although the term excitation-contraction coupling confuses or scares some students, it comes down to this: for a skeletal muscle fiber to contract, its membrane must first be “excited”—in other words, it must be stimulated to fire an action potential. The muscle fiber action potential, which sweeps along the sarcolemma as a wave, is “coupled” to the actual contraction through the release of calcium ions (Ca++) from the SR. Once released, the Ca++ interacts with the shielding proteins, forcing them to move aside so that the actin-binding sites are available for attachment by myosin heads. The myosin then pulls the actin filaments toward the center, shortening the muscle fiber.

In skeletal muscle, this sequence begins with signals from the somatic motor division of the nervous system. In other words, the “excitation” step in skeletal muscles is always triggered by signaling from the nervous system (Figure 10.6).

 Alt text to come.
Figure 10.6 Motor End-Plate and Innervation At the NMJ, the axon terminal releases ACh. The motor end-plate is the location of the ACh-receptors in the muscle fiber sarcolemma. When ACh molecules are released, they diffuse across a minute space called the synaptic cleft and bind to the receptors.

The motor neurons that tell the skeletal muscle fibers to contract originate in the spinal cord, with a smaller number located in the brainstem for activation of skeletal muscles of the face, head, and neck. These neurons have long processes, called axons, which are specialized to transmit action potentials long distances— in this case, all the way from the spinal cord to the muscle itself (which may be up to three feet away). The axons of multiple neurons bundle together to form nerves, like wires bundled together in a cable.

Signaling begins when a neuronal action potential travels along the axon of a motor neuron, and then along the individual branches to terminate at the NMJ. At the NMJ, the axon terminal releases a chemical messenger, or neurotransmitter, called acetylcholine (ACh). The ACh molecules diffuse across a minute space called the synaptic cleft and bind to ACh receptors located within the motor end-plate of the sarcolemma on the other side of the synapse. Once ACh binds, a channel in the ACh receptor opens and positively charged ions can pass through into the muscle fiber, causing it to depolarize, meaning that the membrane potential of the muscle fiber becomes less negative (closer to zero.)

As the membrane depolarizes, another set of ion channels called voltage-gated sodium channels are triggered to open. Sodium ions enter the muscle fiber, and an action potential rapidly spreads (or “fires”) along the entire membrane to initiate excitation-contraction coupling.

Things happen very quickly in the world of excitable membranes (just think about how quickly you can snap your fingers as soon as you decide to do it). Immediately following depolarization of the membrane, it repolarizes, re-establishing the negative membrane potential. Meanwhile, the ACh in the synaptic cleft is degraded by the enzyme acetylcholinesterase (AChE) so that the ACh cannot rebind to a receptor and reopen its channel, which would cause unwanted extended muscle excitation and contraction.

Propagation of an action potential along the sarcolemma is the excitation portion of excitation-contraction coupling. Recall that this excitation actually triggers the release of calcium ions (Ca++) from its storage in the cell’s SR. For the action potential to reach the membrane of the SR, there are periodic invaginations in the sarcolemma, called T-tubules (“T” stands for “transverse”). You will recall that the diameter of a muscle fiber can be up to 100 μm, so these T-tubules ensure that the membrane can get close to the SR in the sarcoplasm. The arrangement of a T-tubule with the membranes of SR on either side is called a triad (Figure 10.7). The triad surrounds the cylindrical structure called a myofibril, which contains actin and myosin.

 Alt text to come.
Figure 10.7 The T-tubule Narrow T-tubules permit the conduction of electrical impulses. The SR functions to regulate intracellular levels of calcium. Two terminal cisternae (where enlarged SR connects to the T-tubule) and one T-tubule comprise a triad—a “threesome” of membranes, with those of SR on two sides and the T-tubule sandwiched between them.

The T-tubules carry the action potential into the interior of the cell, which triggers the opening of calcium channels in the membrane of the adjacent SR, causing Ca++ to diffuse out of the SR and into the sarcoplasm. It is the arrival of Ca++ in the sarcoplasm that initiates contraction of the muscle fiber by its contractile units, or sarcomeres.

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