CHAPTER 9: MUSCULAR SYSTEM OBJECTIVES: 1. Compare and contrast the types of muscle tissues in terms of structure, control, location, and type of contraction, and function. 2. Describe three similarities among the three muscle tissues. 3. Identify the terms used for a muscle fiber's cell membrane and cytoplasm. 4. Describe the functions of muscle tissue. 5. Compare and contrast the functional characteristics of muscle tissue (i.e. excitability, contractility, extensibility, and elasticity). 6. Illustrate how a skeletal muscle is wrapped in four layers of connective tissue. 7. Define the terms tendon, aponeurosis, raphe, and syncytium. 8. Illustrate how the myofibrils that compose skeletal muscle fibers are composed of sarcomeres. Label the thick filaments, thin filaments, A-Band, I-Band and Z-line. 9. Compare and contrast the ultrastructure of thick and thin filaments. 10. Explain the significance of the special membranous organelles found in skeletal muscle tissue. 11. Explain what happens to sarcomere structure when a muscle contracts. 12. Explain the role that calcium plays in contraction. 13. Name the organelle that contains a high concentration of calcium due to the action of a calcium pump. 14. List the sequence of events involved in the power stroke of muscle contraction. 15. Define the terms neuromuscular junction (NMJ), motor unit, motor end plate and neurotransmitter. 16. Identify the neurotransmitter involved in muscle contraction. 17. List the sequence of events involved in skeletal muscle fiber contraction beginning with the necessary motor impulse initiated by the brain. 18. Explain how and why a contracted muscle relaxes. 19. Name the three pathways that regenerate energy/ATP in muscle cells. 20. Outline a general overview of cellular respiration, denoting its two major parts and where each occurs in the cell. Be sure to include starting products, end products, and any additional requirements. Then discuss the significance of this pathway in skeletal muscle contraction (don't forget that the midpoint product can take one of two pathways!!!). 21. Explain how lactic acid is produced and what its accumulation causes. 22. Define the term oxygen debt. 23. Demonstrate the negative feedback mechanisms that maintain thermal homeostasis. 24. Define the term threshold stimulus, and give the numerical value in skeletal muscle cells. 25. A myogram measures a muscle contraction as a twitch. What does this term mean? 26. Describe what is meant by "all or nothing" response in skeletal muscle fibers. 27. Define the term used to describe a myogram that shows a series of twitches with increasing strength. 28. Name the term when a myogram illustrates a sustained contraction that lacks even slight relaxation between twitches. 29. Compare and contrast isometric and isotonic muscle contractions. 30. List the differences between fast and slow muscle fibers, and explain why they are also called white and red fibers, respectively. 31. Distinguish between multi-unit and visceral smooth muscle and give examples of each type. 32. Define peristalsis. 33. List the characteristics of cardiac muscle tissue. 34. Define the terms origin and insertion as they relate to a skeletal muscle. 35. Define the terms prime mover, antagonist, synergists, and fixator as they relate to muscle actions, and use the thigh muscles as an example. 36. For every skeletal muscle listed in this outline, be able to complete the following: A. locate the muscle on a diagram or human muscle model. describe the shape and/or fascicle arrangement of the muscle. B. C. D. identify key origin and insertion sites. describe the action. I. OVERVIEW OF MUSCLE TISSUES: Review from Chapter 5. A. Skeletal B. Similarities: 1. All muscle cells are elongated = muscle fibers; 2. Muscle contraction depends on two kinds of myofilaments (actin and myosin); 3. The cell membrane of a muscle cell is called "sarcolemma", while the cytoplasm of a muscle cell is called "sarcoplasm". C. Skeletal Muscle Characteristics: 1. long, thin and multi-nucleated fibers; 2. striations; 3. voluntary control; 4. arranged into packages called muscles that attach to and cover the bony skeleton; 5. contracts rapidly & vigorously, but tired easily; may exert great force. D. Cardiac Muscle Characteristics: 1. network of branched fibers connected by gap junctions (intercalated disks); 2. only in heart; 3. striations; 4. involuntary control; 5. contracts at rhythmic, steady rate set by "pacemaker". E. Smooth Muscle Characteristics: 1. lacks striations; 2. walls of hollow visceral organs and blood vessels; 3. involuntary control; 4. contractions are slow & sustained. F. Functions: 1. Movement = locomotion & manipulation, vision, facial expression (skeletal), blood pumping (cardiac), food digesting, urination (smooth); 2. Posture Maintenance (skeletal) 3. Joint Stability (skeletal) 4. Heat Generation (skeletal) G. II. Muscle Types: Smooth Cardiac Functional Characteristics of Muscle: 1. Excitability = the ability to receive and respond to stimuli; 2. Contractility = the ability to shorten forcibly when stimulated; 3. Extensibility = the ability to be stretched or extended; 4. Elasticity = the ability to bounce back to original length SKELETAL MUSCLE Introduction: Each skeletal muscle is an organ made up of skeletal muscle fibers, connective tissue coverings, blood vessels, and nerve fibers. A. Structure of a Skeletal Muscle: 1. Connective Tissue Coverings: See Fig 9.2, page 280. a. Each muscle fiber (cell) is wrapped in a thin, delicate layer of CT called endomysium. b. Many muscle fibers are bundled together into groups called fascicles; See Fig 9.3, page 280. o Each fascicle is wrapped in a second layer of CT made of collagen called perimysium. c. Many fascicles are bundled together to form a skeletal muscle. o Each skeletal muscle is covered by a third layer of dense, fibrous CT called epimysium. d. Each skeletal muscle is then covered by a fourth, very tough fibrous layer of CT called deep fascia. o The deep fascia may extend past the length of the muscle (tendon or aponeurosis), and attach that muscle to a bone, cartilage or muscle. o See Fig 9.1, page 279 to compare these two extensions. B. Skeletal Muscle Fibers Recall that skeletal muscle tissue possesses striations: See Fig 9.7, page 282. 1. A muscle fiber is a long, thin cell; a. Each muscle fiber is composed of myofibrils; o Each myofibril is composed of two types of protein filaments (cytoskeletal elements): See Fig 9.4, page 281 1. Thick filaments primarily composed of the protein myosin; 2. Thin filaments primarily composed of the protein actin. o Striations are caused by the arrangement of thick and thin filaments within the myofibrils: See Fig 9.5, page 281. 1. A-Band = dark area = overlapping of thick and thin filaments; 2. I-Band = light area = thin filaments alone. o The length of each myofibril is divided into sarcomeres: 1. Sarcomeres meet one another at an area called the Z-line. o See Fig 9.4, Fig 9.5, Fig 9.6 page 281 and Fig 9.7, page 282 II. SKELETAL MUSCLE B. Skeletal Muscle Fibers 2. 3. Thick filaments = protein myosin. a. rod-like tail (axis) that terminates in two globular heads or cross bridges; b. Cross bridges interact with active sites on thin filaments; Thin filaments = protein actin. a. coiled helical structure (resembles twisted strands of pearls): b. Tropomyosin = rod-shaped protein spiraling around actin backbone to stabilize it; c. Troponin = complex of polypeptides: o o o d. Both tropomyosin and troponin help control actin's interaction with myosin during contraction. 4. Within the sarcoplasm of a muscle fiber, there are two specialized membranous organelles: See Fig 9.7, page 282. a. Sarcoplasmic reticulum (SR) o Network of membranous channels that surrounds each myofibril and runs parallel to it. o Same as endoplasmic reticulum in other cells. o SR has high concentrations of calcium ions compared to the sarcoplasm (maintained by active transport calcium pump). o When stimulated by muscle impulse, membranes become more permeable to calcium ions and calcium diffuses out of SR and into sarcoplasm. b. Transverse tubules (TT) o set of membranous channels that extends into the sarcoplasm as invaginations continuous with muscle cell membrane (sarcolemma). o TTs are filled with extracellular fluid and extend deep into the cell. o Each TT runs between two enlarged portions of SR called cisternae. These structures form a triad near the region where actin and myosin overlap. c. SR and TT are involved in activating the muscle contraction mechanism (discussed in greater detail later). d. Because one TT is associated with two SR they are termed the Triad II. SKELETAL MUSCLE C. Skeletal Muscle Contraction: 1. Neuromuscular Junction - Stimulation of Skeletal Muscle Cell: See Figure 9.9, page 283. a. b. 2. Stimulus for Contraction See Table 9.1, page 287. a. b. c. d. e. II. Neuromuscular Junction (NMJ) = the site where a motor nerve fiber and a skeletal muscle fiber meet; (also called a synapse or synaptic cleft) In order for a skeletal muscle to contract, its fibers must first be stimulated by a motor neuron. c. Motor Unit = one motor neuron and many skeletal muscle fibers; See Fig 9.9, page 283. d. Motor End Plate = the specific part of a skeletal muscle fiber's sarcolemma directly beneath the NMJ. e. Neurotransmitter = chemical substance released from a motor end fiber, causing stimulation of the sarcolemma of muscle fiber; acetylcholine (ACh). f. Synaptic cleft – small space between neuron and muscle Introduction: The function of skeletal muscle is to move bones of the skeleton under voluntary control. Contraction of a skeletal muscle fiber is a complex interaction of several cellular and chemical constituents. The final result is a movement whereby actin and myosin filaments slide past one another. Accordingly, the muscle fiber shortens and pulls on its attachments. The process begins when a motor impulse is initiated by the brain, travels down the spinal cord, into a motor neuron, which branches into many motor nerve fibers/endings; o Each motor nerve fiber extends to the motor end-plate of a skeletal muscle fiber forming a neuromuscular junction (NMJ); When the motor impulse reaches the end of the motor nerve fiber/ending, the membrane is depolarized (-70mV to -55mV); o calcium ions rush into motor nerve fiber, and o neurotransmitter (Acetylcholine) is released into the NMJ (via exocytosis). Acetylcholine diffuses across the NMJ & stimulates/depolarizes the motor end-plate (sarcolemma) of a skeletal muscle fiber from 100mV to –70mV; The muscle impulse travels over the surface of the skeletal muscle fiber and deep into the muscle fiber by means of the TT SKELETAL MUSCLE C. Skeletal Muscle Contraction: II. 3. Excitation Contraction Coupling a. The muscle impulse reaches the sarcoplasmic reticulum, which releases calcium ions into the sarcoplasm of the muscle fiber; b. Calcium binds to troponin, moving tropomyosin and exposing myosin binding sites on actin filament; c. Cross-bridges (linkages) form between actin and myosin; d. Actin filaments are pulled inward by myosin cross-bridges; e. The muscle fiber shortens as contraction occurs. 4. Sliding Filament Theory: a. most popular theory concerning muscle contraction; b. first proposed by Hugh Huxley in 1954; c. states that muscle contraction involves the sliding movement of the thin filaments (actin) past the thick filaments (myosin); d. Sliding continues until the overlapping between the thin & thick filaments is complete. *Remember that in a relaxed muscle cell, overlapping of thick and thin filaments is only slight. e. Changes in muscle during contraction: See Fig 9.8, page 283. o The distance between the Z-lines of the sarcomeres decreases; o The I-Bands (light bands) shorten; o The A-Bands move closer together, but do not diminish in length. f. The Role of Calcium in Contraction Mechanism: o In a resting muscle cell (i.e. in the absence of calcium ions): Tropomyosin blocks or inhibits the myosin binding sites on actin. See Fig 9.10a, page 285. o When calcium ions (Ca++) are present: Ca++ binds to troponin causing a conformational change in the troponin-complex, which causes: 1. Tropomyosin to move 2. which "opens" or exposes the myosin binding sites on actin; 3. This results in interaction between the active sites on actin and the heads (or cross bridges) of myosin. See Fig 9.10b, page 285 SKELETAL MUSCLE C. Skeletal Muscle Contraction: 5. Cross-Bridge Cycling See Fig 9.11, page 286. When calcium ions are present, the myosin binding sites on actin are exposed: a. Cross-bridge attaches. o ATP breakdown provides energy to “cock” myosin head. o “Cocked” myosin attaches to exposed actin binding site b. Cross-bridge (myosin head) springs from cocked position and pulls on actin filament. c. Cross bridges break. o ATP binds to cross-bridge (but is not yet broken down) o Myosin heads are released from actin. * As long as calcium ions and ATP are present, this walking continues until the muscle fiber is fully contracted. 6. Relaxation: a. Acetylcholinesterase is an enzyme present in the NMJ; b. It immediately destroys acetylcholine, so the motor end-plate is no longer stimulated (i.e. it cannot cause continuous muscle contraction). c. Calcium ions are transported from sarcoplasm back into sarcoplasmic reticulum. d. Linkages between actin and myosin are broken. e. The muscle fiber relaxes. 7. Energy Sources for Contraction: a. Introduction: The energy used to power the interaction between actin and myosin comes from ATP. b. ATP stored in skeletal muscle lasts only about six seconds. o ATP must be regenerated continuously if contraction is to continue. o There are three pathways in which ATP is regenerated: 1. Coupled Reaction with Creatine Phosphate (CP) 2. Anaerobic Cellular Respiration 3. Aerobic Cellular Respiration c. Coupled Reaction with Creatine Phosphate (CP) See Fig 9.12, page 287. o o o II. CP + ADP <------> creatine + ATP Muscle stores a lot of CP, This coupling reaction allows for about 10 seconds worth of ATP. SKELETAL MUSCLE C. Skeletal Muscle Contraction: 8. Oxygen Supply and Cellular Respiration: See Fig 9.13, page 288. Review from Chapter 4. a. Anaerobic Respiration o Steps are called glycolysis. o Steps occur in the cytoplasm of the cell. o Results in production of pyruvic acid and 2 ATP. b. Aerobic Respiration o Steps are called citric acid cycle and electron transport chain. o Oxygen is required. o Steps occur in the mitochondrion of the cell. o Results in CO2, water and 36ATP. * Read box at the bottom of page 309, which illustrates how a runner utilizes the sources discussed above. 9. Muscle Fatigue a. Muscle fatigue is a state of physiological inability to contract; b. If no oxygen is available in muscle cells to complete aerobic respiration, pyruvic acid is converted to lactic acid, which causes muscle fatigue and soreness; See Fig 9.13 and 9.14, pages 288 + 289. c. Results from a relative deficit of ATP and/or accumulation of lactic acid (which decreases pH). 10. a. b. 11. II. Oxygen Debt: See Fig 9.14, page 289. The oxygen debt is the amount of oxygen necessary to support the conversion of lactic acid to glycogen. needed to replenish spent glycogen stores. Heat Production a. Almost half of the energy released during muscle contraction is lost to heat, which helps maintain our body temperature at 37o C. b. Excessive heat is lost through many negative feedback mechanisms (discussed in chapter 1) including sweating, dilation of superficial blood vessels, increased breathing rate, and increased heart rate. SKELETAL MUSCLE D. MUSCULAR RESPONSES 1. 2. 3. II. Threshold Stimulus a. The minimal strength of stimulation required to cause contraction. b. A skeletal muscle fiber’s resting membrane potential must be depolarized from –100mV to –70mv before an impulse begins; Therefore the threshold stimulus is +30mV. Recording a Muscle Contraction: See Fig 9.15, page 290. a. A myogram is a recording of a muscle contraction. b. A twitch is a single contraction that lasts a fraction of a second, followed by relaxation. c. The delay between stimulation and contraction is called the latent period. d. A muscle fiber must return to its resting state (-100mV) before it can be stimulated again. This is called the refractory period. e. All-or-Nothing Response o If a muscle fiber is brought to threshold or above, it responds with a complete twitch. o If the stimulus is sub-threshold, the muscle fiber will not respond. f. Staircase Effect (treppe): See Fig 9.16a, page 291. o Most muscle fiber contraction is “all-or-nothing”. o However a muscle fiber that has been inactive can be subjected to a series of stimuli and: the fiber undergoes a series of twitches with relaxation between, and the strength of each successive contraction increases slightly. o This phenomenon is small and brief and involves excess calcium in sarcoplasm. Summation: See Fig 9.16b, page 291. a. When several stimuli are delivered in succession to a muscle fiber, it cannot completely relax between contractions. b. The individual twitches begin to combine and the muscle contraction becomes sustained. In a sustained contraction, the force of individual twitches combines in a process called summation. When the resulting sustained contraction lacks even slight relaxation, it is called titanic contraction; c. Fig 9.16c, page 291. SKELETAL MUSCLE D. MUSCULAR RESPONSES 4. Motor Units: See Fig 9.9, page 283. a. Definition: A motor unit is a motor neuron and the many skeletal 5. 6. 7. muscle fibers it stimulates. b. Because the motor neuron branches into several motor nerve endings, it can stimulate many skeletal muscles fibers simultaneously, which then contract simultaneously. c. The number of muscle fibers in a motor unit varies from 10hundreds. Recruitment of Motor Units a. Because a whole muscle is composed of many motor units, controlled by many different motor neurons, simultaneous contraction of all units does not necessarily occur. b. As the intensity of stimulation increases, recruitment of motor units increases, until all contract simultaneously. Sustained Contractions a. Even when a muscle is at rest, a certain amount of sustained contraction is occurring in its fibers. This is called muscle tone. o Muscle tone is very important in maintaining posture. Types of Contractions: See Fig 9.18, page 295. a. Isotonic contractions o The muscle shortens and its attachment(s) move(s). b. Isometric contraction, o The muscle becomes taut, but the attachment(s) do not move; tensing a muscle; c. Most muscular movements involve both isotonic and isometric contractions. d. Read Clinical Application 9.2, page 293 re: use and disuse of skeletal muscles. 8. III. Fast and Slow Muscle Fibers a. Muscle fibers vary in contraction speed (i.e. slow or fast twitch). b. Slow-Twitch Fibers are also called red fibers. o contain oxygen carrying pigment, myoglobin, receive a rich blood supply, and contain many mitochondria o can generate ATP fast enough to keep up with breakdown. o These fibers contract for long periods without fatiguing. c. Fast-twitch fibers are also called white fibers. o contain less myoglobin, blood, and fewer mitochondria. o contain extensive sarcoplasmic reticulum to store and reabsorb calcium. o These fibers contract rapidly, but fatigue easily due to lactic acid accumulation. d. Read box at bottom of page 293 re: dark and white meat. SMOOTH MUSCLE: A. Introduction: The contraction mechanism of smooth muscle is similar to that of skeletal muscle in that interaction occurs between actin and myosin, however the transverse tubules and sarcoplasmic reticula are greatly reduced, and troponin is absent. B. Smooth Muscle Fibers - Two types: 1. Multi-unit smooth muscle a. location: o irises of eyes o walls of blood vessels b. Contraction is rapid and vigorous (similar to skeletal muscle tissue). 2. C. Visceral smooth muscle a. Location = the walls of hollow organs b. Contraction is slow and sustained. o Rhythmicity = pattern of repeated contractions; o Peristalsis = wave-like motion that helps push substances through passageways. c. Structure: o random arrangement of actin and myosin filaments. o Two layers of muscle surround the passageway. 1. inner circular layer 2. outer longitudinal layer Smooth Muscle Contraction: 1. A protein, calmodulin binds to calcium ions (no troponin) and activates the contraction mechanism. 2. Most calcium diffuses in to smooth muscle cells from the extracellular fluid (reduced SR). 3. Norepinephrine and acetylcholine are smooth muscle neurotransmitters. 4. Contraction is slow and sustained. IV. CARDIAC MUSCLE: 296. See Fig 9.19, page Will be studied in greater detail in Chapter 15. A. Location: 1. Only in heart. B. Anatomy: 1. Striated uninuclear cells joined end-to-end forming a network. a. Cell junctions are called intercalated discs. o gap junctions 2. Arrangement of actin and myosin not as organized as skeletal muscle. 3. Contains sarcoplasmic reticula, transverse tubules, and numerous mitochondria: a. Sarcoplasmic reticulum is less developed than SR in skeletal muscle and stores much less calcium. C. Physiology 1. Self-exciting tissue (i.e. “Pacemaker”); 2. Rhythmic contractions (60-100 beats/minute); 3. Involuntary, all-or-nothing contractions a. Function as a “syncytium” (all-or-nothing) 4. Pumps blood to: a. Lungs for oxygenation; b. Body for distribution of oxygen and nutrients. V. SKELETAL MUSCLE ACTIONS A. Introduction: Skeletal muscles generate a great variety of body movements. The action of a muscle primarily depends upon the joint associated with it and the manner in which the muscle is attached on either side of that joint. B. Origin and Insertion: Recall that skeletal muscles are usually attached to a fixed body part and a movable body part: See Fig 9.20, page 297. 1. The origin of a muscle is its immovable (anchored) end. 2. The insertion of a muscle is the movable end of a muscle. *When a muscle contracts and shortens, its insertion is pulled toward its origin. C. Review of Skeletal Muscle Actions: 1. Flexion = decreasing the angle between 2 bones; a. Dorsiflexion = decreasing the angle between the foot and shin; b. Plantar flexion = pointing toes; 2. Extension = increasing the angle between 2 bones; 3. Abduction = moving a body part away from the midline; 4. Adduction = moving a body part toward the midline; 5. Circumduction = movement in a circular (cone-shaped) motion; 6. Rotation = turning movement of a bone about its long axis; (i.e. atlas/axis); 7. Supination = thumbs up; 8. Pronation = thumbs down; 9. Inversion = sole of foot in; 10. Eversion = sole of foot out; 11. Elevation = lifting a body part; (i.e. shoulder shrug); 12. Depression = returning a body part to pre-elevated position. D. Interactions of Skeletal Muscles 1. Prime Mover (agonist) = the primary muscle responsible for a movement. The biceps brachii in flexing the arm at the elbow, 2. Antagonist(s) = the muscle(s) in opposition to the action of the prime mover. The antagonist relaxes (or stretches) during the prime movement. The triceps brachii is the antagonist of the biceps brachii when we flex the arm at the elbow. 3. Synergist(s) = muscles that assist the prime mover. The brachialis helps the biceps brachii during elbow flexion. 4. Fixators = muscle groups that stabilize the origin of the prime mover (i.e. hold it in place) so that the prime mover can act more efficiently. The scapula is the origin for many arm muscles, but it must be held in place by fixator muscles in order to function in this way. a. serratus anterior b. pectoralis minor VI. MAJOR SKELETAL MUSCLES - Naming CHARACTERISTIC EXAMPLES EXAMPLES IN HUMANS Direction of fascicles relative to midline rectus = parallel transverse = perpendicular oblique = at 45o angle Rectus abdominis Transversus abdominis External Oblique Location (i.e. the bone or body part that a muscle covers) frontal bone tibia Frontalis Tibialis Anterior Relative Size maximus = largest longus = longest brevis = shortest Gluteus maximus Palmaris longus Peroneus longus Number of Origins (Heads) biceps = 2 origins triceps = 3 origins Biceps brachii Triceps brachii Shape deltoid = triangle trapezius = trapezoid serratus = saw-toothed orbicularis = circular Deltoid Trapezius Serratus anterior Orbicularis oris Location of Origin and/or Insertion origin = sternum insertion = mastoid process Sternocleidomastoid Action of Muscle flexion extension adduction Flexor carpi radialis Extensor digitorum Adductor longus VII. MAJOR SKELETAL MUSCLES (Keyed at the end of this outline) Use the figures and tables on pages 298-325 to complete the following information. If a muscle is starred (*), please include its origin and insertion. Also see References Plates on pages 332-336. A. Muscles of Facial Expression: See Fig 9.22 on page 298, and Table 9.3 on page 299. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Epicranius Frontalis Occipitalis Orbicularis oris Zygomaticus (*) Buccinator Platysma Orbicularis oculi B. Muscles of Mastication: See Fig 9.22 on page 298, and Table 9.4 on page 300. NAME OF MUSCLE Masseter(*) Temporalis LOCATION/ DESCRIPTION ACTION VII. MAJOR SKELETAL MUSCLES (continued) C. Muscles that move the Head and Vertebral Column: See Fig 9.23 on page 301, and Table 9.5 on page 302. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Sternocleidomastoid (*) Erector Spinae D. Muscles that move the Pectoral Girdle: See Fig 9.25 on page 304, Fig 9.26 on page 305, and Table 9.6 on page 304. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Trapezius (*) Pectoralis minor Serratus anterior E. Muscles that move the Arm (Humerus): See Fig 9.25, page 304, Fig 9.26, page 305, and Table 9.7, page 306. NAME OF MUSCLE LOCATION/ DESCRIPTION Pectoralis major (*) Latissimus dorsi Deltoid VII. MAJOR SKELETAL MUSCLES (continued) ACTION F. Muscles that move the Forearm (radius & ulna): See Fig 9.28, page 307, Fig 9.29, page 309, and Table 9.8, page 308. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Biceps Brachii (*) Brachialis Brachioradialis Triceps brachii G. Muscles that move the Hand See Fig 9.30, page 310, Fig 9.31, page 311, and Table 9.9, page 311. NAME OF MUSCLE Flexor carpi radialis Flexor carpi ulnaris Palmaris longus Extensor digitorum LOCATION/ DESCRIPTION ACTION VII. MAJOR SKELETAL MUSCLES (continued) H. Muscles of the Abdominal Wall: See Fig 9.32, page 313, and Table 9.10, page 312. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Rectus abdominis (*) External Oblique Internal Oblique Transversus abdominis I. Muscles of the Pelvic Outlet See Fig 9.33 page 314 and Table 9.11 page 315. NAME OF MUSCLE LOCATION/ DESCRIPTION Levator ani Coccygeus Superficial transversus perinei Bulbospongiosus Ischiocavernosus Sphincter urethrae VII. MAJOR SKELETAL MUSCLES (continued) ACTION J. Muscles that move the Thigh (Femur): See Fig 9.18, page 341, Fig 9.37, page 319, and Table 9.12, page 319. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Gluteus Maximus (*) Gluteus Medius Adductor Longus K. Muscles that move the Leg (Tibia & Fibula): See Fig 9.35, page 317, Fig 9.37, page 319, and Table 9.13, page 320. NAME OF MUSCLE Rectus femoris Vastus lateralis Vastus Medialis Vastus intermedius Sartorius (*) Biceps femoris Semitendinosus Semimembranosus LOCATION/ DESCRIPTION ACTION VII. MAJOR SKELETAL MUSCLES (continued) L. Muscles that move the Foot & Toes See Fig 9.39, page 323, Fig 9.40, page 324, Fig 9.41, page 325, and Table 9.14, page 348. NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Tibialis anterior Fibularis longus Gastrocnemius (*) Soleus Note the location of the Calcaneal Tendon in Fig 9.40, page 324. VIII. LIFE SPAN CHANGES A. Supplies of ATP, myoglobin, and creatine phosphate in muscle fibers begin to decline in one’s forties. B. Half of one’s muscle mass has been replaced by connective and adipose tissue by age 80, and reflexes are reduced. C. Exercise is the best way to maintain muscle function. IX. Homeostatic Imbalances/Disorders A. Tendinitis (page 278) B. Compartment Syndrome (page 278) C. Muscle Strain (page 282) D. Poliomyelitis (page 283) E. Myasthenia Gravis (CA 9.1, page 284) F. Duchenne Muscular Dystrophy (page 282) G. Rigor Mortis (page 287) H. Botulism (page 284) I. Use and Disuse of Skeletal Muscles (CA 9.2, page 293) J. TMJ (CA 9.3, page 300) K. Parkinson's Disease (page 312) X. Innerconnections of the Muscular System: See page 326. XI. Clinical Terms Related to the Muscular System. See page 327. SKELETAL MUSCLE SUMMARY TABLES A. Muscles of Facial Expression NAME OF MUSCLE LOCATION/ DESCRIPTION Covers cranium Epicranius Frontalis Occipitalis ACTION elevates eyebrow over forehead over occipital Orbicularis oris circular muscle around the mouth closes lips (“kissing muscle”) Zygomaticus (*) Origin: zygomatic arch Insertion: corners of orbicularis oris muscle that connects zygomatic arch to corner of mouth elevates corners of mouth (“smiling muscle”) Buccinator hollow of cheek compresses cheeks “trumpeter’s muscle” Platysma over lower jaw to neck depresses mandible Orbicularis oculi circular muscle around eye closes eye B. Muscles of Mastication NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Masseter(*) Origin: Zygomatic Arch Insertion: Lateral Mandible over lateral mandible elevates mandible Temporalis convergent muscle over temporal bone elevates mandible SKELETAL MUSCLE SUMMARY TABLES C. Muscles that move the Head and Vertebral Column: NAME OF MUSCLE LOCATION/ DESCRIPTION Sternocleidomastoid(*) Origin: sternoclavicular region Insertion: mastoid process of temporal Major neck muscle flexion of head toward chest (both contracted) rotation/abduction of head (as antagonists) Erector Spinae Group of midline dorsum muscles Maintain posture D. ACTION Muscles that move the Pectoral Girdle NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Trapezius (*) Origin: occipital bone & spines of C7-T12 Insertion: clavicle and acromion process of scapulae Trapezoid shaped muscle in posterior neck and upper back elevates pectoral girdle (“shoulder shrug”) Pectoralis minor Muscle deep to Pectoralis major scapula fixator Serratus anterior Saw-toothed lateral thoracic muscle scapula fixator E. Muscles that move the Arm (Humerus) NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Pectoralis major (*) Origin: clavicle, sternum, & costal cartilages of ribs 1-6 Insertion: Greater tubercle of humerus Large, convergent chest muscle flexes arm medially (pull arms forward and together) Latissimus dorsi Large, back muscle adduction of humerus Deltoid Triangular shaped shoulder muscle abduction of humerus SKELETAL MUSCLE SUMMARY TABLES F. Muscles that move the Forearm (radius & ulna) NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Biceps Brachii (*) Origin: Coracoid process Insertion: Radial tuberosity fusiform, parallel, anterior upper arm muscle (two origins) flexion of arm at elbow (prime mover) Brachialis muscle beneath biceps brachii flexion of arm at elbow (synergist) Brachioradialis lateral muscle between upper and forearm flexion of arm at elbow (synergist) Triceps brachii posterior upper arm muscle (three heads) extension of arm at elbow G. Muscles that move the Hand NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Flexor carpi Radialis anterior, lateral forearm muscle flexion of wrist Flexor carpi Ulnaris anterior, medial forearm muscle flexion of wrist Palmaris longus anterior forearm muscle located between two above flexion of wrist Extensor digitorum posterior forearm muscle extension of wrist/fingers SKELETAL MUSCLE SUMMARY TABLES H. Muscles of the Abdominal Wall NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Rectus abdominis (*) Origin: pubic crest & symphysis Insertion: xiphoid process & costal cartilages of 5-7th ribs strap like muscle from costal cartilages to ilium tenses abdominal wall External Oblique superficial/lateral oblique abdominal muscle tenses abdominal wall Internal Oblique deep oblique abdominal muscle tenses abdominal wall Transversus abdominis deep abdominal muscle that runs perpendicular to rectus abdominis tenses abdominal wall I. Muscles of the Pelvic Outlet NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Levator ani Thin sheet surrounding anus, urethra and vagina Supports viscera and aids sphincters Coccygeus Fan shaped, posterior to levator ani Same as above Superficial transversus perinei Band of muscle anterior to levator ani Supports viscera Bulbospongiosus Surrounds base of penis or vagina Male – assists ejaculation Female – constricts vaginal orifice Ischiocavernosus Band of muscle lateral to bulbospongiosus Same as above Sphincter urethrae Surround urethra Opens and closes urethral orifice SKELETAL MUSCLE SUMMARY TABLES J. Muscles that move the Thigh (Femur) NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Gluteus Maximus (*) Origin: dorsal ilium, sacrum, coccyx Insertion: posterior femur buttocks, largest muscle in body extension of hip at thigh (as in walking or climbing stairs) Gluteus Medius lateral hip muscle abduction of femur Adductor Longus medial thigh muscle adduction of femur K. Muscles that move the Tibia & Fibula NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Rectus femoris anterior thigh; quadriceps extension of leg at knee Vastus lateralis lateral anterior thigh; quadriceps extension of leg at knee Vastus Medialis medial anterior thigh; quadriceps extension of leg at knee Vastus intermedius deep anterior thigh; quadriceps extension of leg at knee Sartorius (*) Origin: iliac spine Insertion: medial tibia parallel strap-like muscle that crosses thigh flexion of knee forward Biceps femoris posterior thigh; hamstring flexion of leg at knee Semitendinosus posterior thigh; hamstring flexion of leg at knee Semimembranosus posterior thigh; hamstring flexion of leg at knee SKELETAL MUSCLE SUMMARY TABLES L. Muscles that move the Foot & Toes NAME OF MUSCLE LOCATION/ DESCRIPTION ACTION Tibialis anterior anterior to tibia dorsiflexion Peroneus longus lateral to fibula eversion Gastrocnemius (*) Origin: condyles of femur Insertion: calcaneus posterior lower leg (i.e. calf muscle); two origins plantar flexion (prime mover) Soleus deep to gastrocnemius plantar flexion (synergist) Shier, Butler, and Lewis: Hole’s Human Anatomy and Physiology, 10 th ed. Chapter 9: Muscular System I. Structure of a Skeletal Muscle A. Introduction 1. A__________________________________ is an organ of the skeletal system. 2. A skeletal muscle is composed of ____________________________________ B. Connective Tissue Coverings 1. Fascia is ________________________________________________________ 2. A tendon is ______________________________________________________ 3. Tendons connect a muscle to ________________________________________ 4. An aponeurosis is _________________________________________________ 5. Epimysium is ____________________________________________________ 6. Perimysium is____________________________________________________ 7. A fascicle is _____________________________________________________ 8. Endomysium is___________________________________________________ 9. Deep fascia is ____________________________________________________ 10. Subcutaneous fascia is ____________________________________________ 11. Subserous fascia is _______________________________________________ C. Skeletal Muscle Fibers 1. A skeletal muscle fiber is a single ____________________________________ 2. The sarcolemma is ________________________________________________ 3. The sarcoplasm is _________________________________________________ 4. The sarcoplasm contains ___________________________________________ 5. Myofibrils are _________________ and are located in the ________________ 6. Myofibrils play a fundamental role in _________________________________ 7. Thick myofilaments are composed of _________________________________ 8. Thin myofilaments are composed of __________________________________ 9. The organization of myofilaments produce _____________________________ __________________________________________________________________ 10. A sarcomere is __________________________________________________ 11. Myofibrils may be thought of as _____________________ joined end to end. 12. I bands are composed of __________________________________________ 13. Z lines are ______________________________________________________ 14. A bands are composed of __________________________________________ 15. The H zone is ___________________________________________________ 16. The M line is ___________________________________________________ 17. Titin connects ___________________________________________________ 18. A sarcomere extends from one _____________________________________ 19. Each myosin molecule consists of ___________________________________ __________________________________________________________________ 20. Thin filaments consist of double ____________________________________ 21. Actin has a binding site to which ___________________________ can attach. 22. Troponin and tropomyosin associate with _____________________________ 23. Sarcoplasmic reticulum is _________________________________________ 24. Transverse tubules are ____________________________________________ 25. Cisternae are____________________________________________________ 26. A triad is formed by ______________________________________________ II. Skeletal Muscle Contraction A. Neuromuscular Junction 1. Each skeletal muscle is connected to ____________________________ 2. A motor neuron passes out from _____________________________________ 3. Normally, a skeletal muscle fiber contracts only upon ____________________ __________________________________________________________________ 4. A neuromuscular junction is ________________________________________ 5. A motor end plate is _______________________________________________ 6. A motor unit is ___________________________________________________ 7. A synaptic cleft separates___________________________________________ 8. Synaptic vesicles store _____________________________________________ B. Stimulus for Contraction 1. __________________________ is the neurotrasnmitter that motor neurons use to control skeletal muscle. 2. ACh is synthesized in the cytoplasm of ________________________________ and is stored in _____________________________________________________ 3. When a nerve impulse ________________________ , acetylcholine is released into the synaptic cleft. 4. ACh combines with ____________________________ on the motor end plate, and stimulates ______________________________________________________ 5. A muscle impulse is _______________________________________________ 6. A muscle impulse changes __________________________________________ 7. Ultimately the muscle impulse reaches ________________________________ C. Excitation Contraction Coupling 1. The sarcoplasmic reticulum has a high concentration of ___________________ 2. In response to a muscle impulse, the membranes ________________________ ___________________________, and the calcium ________________________ __________________________________________________________________ 3. When a muscle fiber is at rest, ________________________ block the binding sites on the actin molecules. 4. Calcium ions bind to ____________________, changing its _______________ and altering ________________________________________________________ 5. The movement of the tropomyosin molecule exposes _____________________ __________________________, allowing________________________________ D. The Sliding Filament Theory 1. The functional unit of skeletal muscles is ______________________________ 2. According to the sliding filament theory, when sarcomeres shorten, _________ __________________________________________________________________ 3. As contraction occurs, ________________ and ______________ get narrower and _____________________________________________ move closer together. E. Cross-bridge Cycling 1. The force that shortens the sarcomeres comes from ______________________ __________________________________________________________________ 2. A myosin cross-bridge attaches to actin in order to _______________________ __________________________________________________________________ 3. The cross-bridge can release, straighten, and combine with ________________ __________________________________________________________________ 4. Myosin cross-bridges contain the enzyme ______________________________ 5. ATPase catalyzes _________________________________________________ 6. The force for muscle contraction is provided by _________________________ __________________________________________________________________ 7. Breaking down of ATP puts the myosin cross-bridge in a __________ position. 8. When a muscle is stimulated to contract, a ____________________ attaches to ___________________________ and pulls ______________________________ , shortening the ______________________________________________________ 9. When another ATP binds, the ______________________________ is released, then breaks down the ____________________to return to ___________________ 10. The cross-bridge cycle may repeat over and over as long as _______________ __________________________________________________________________ F. Relaxation 1. In order for a muscle fiber to relax, acetylcholine must be decomposed by an enzyme called ________________________________________________ 2. The action of acetycholinesterase prevents _____________________________ __________________________________________________________________ 3. When acetylcholine is broken down, __________________________________ ____________________________________________________________ ceases. 4. The calcium pump moves calcium ___________________________________ __________________________________________________________________ 5. When calcium is removed from the cytoplasm, the _______________________ break and _____________________________________ rolls back into its groove, preventing _________________________________________________________ 6. ATP is necessary for both muscle ____________________________________ and _______________________________________________________________ 7. The trigger for contraction is ________________________________________ G. Energy Sources for Contraction 1. Creatine phosphate is ________________________________________ 2. Creatine phosphate contains __________________________________ 3. As ATP is decomposed to ADP, the energy from __________________ is transferred back to __________________________________________ 4. After creatine phosphate is used, a muscle cell must depend on _______ as a source of energy for synthesizing _______________________________________________ 5. Typically a muscle stores glucose in the form of __________________ H. Oxygen Supply and Cellular Respiration 1. Glycolysis occurs _______________________ and is ____________________ 2. Glycolysis releases a few ___________________________________________ 3. The complete break down of glucose occurs in __________________________ and requires ________________________________________________________ 4. The citric acid cycle and electron transport chain produce _________________ __________________________________________________________________ 5. Oxygen is carried in the blood stream bound to _________________________ 6. Myoglobin is _____________________________________________ in color. 7. Myoglobin stores ___________________________________ in muscle tissue. I. Oxygen Debt 1. Lactic acid threshold is ____________________________________________ 2. Under anaerobic conditions, glycolysis breaks down glucose into ___________ and converts it to ____________________________________________________ 3. Lactic acid is carried by the blood to the _______________________________ 4. Liver cells can convert lactic acid to __________________________________ 5. Oxygen debt reflects ______________________________________________ __________________________________________________________________ J. Muscle Fatigue 1. Fatigue is _______________________________________________________ 2. Fatigue may result from ____________________________________________ __________________________________________________________________ 3. A cramp is ______________________________________________________ 4. Physically fit people make less lactic acid because _______________________ __________________________________________________________________ K. Heat Production 1. Heat is a by-product of_____________________________________________ 2. Blood transports heat _____________________________________________ , which helps ________________________________________________________ III. Muscle Responses A. Introduction 1. One way to observe muscle contraction is to____________________________ __________________________________________________________________ 2. An________________________ is usually used to produce muscle contraction. B. Threshold Stimulus 1. Threshold stimulus is ________________________________________ 2. An impulse in _______________________________ normally releases enough ______________________ to bring the muscle fibers in its __________________ to ________________________________________________________________ C. Recording a Muscle Contraction 1. A twitch is ______________________________________________________ 2. A myogram is ____________________________________________________ 3. Three periods of a muscle fiber contraction are__________________________ __________________________________________________________________ 4. During the period of contraction, a muscle fiber is _______________________ __________________________________________________________________ 5. The latent period is________________________________________________ 6. The period of relaxation is __________________________________________ 7. The refractory period is ____________________________________________ 8. An all-or-none response is __________________________________________ __________________________________________________________________ 9. The length to which a muscle is stretched before stimulation affects _________ __________________________________________________________________ 10. If a muscle is stretched well beyond its normal resting length, the force will __________________________________________________________________ 11. At very short muscle lengths, the sarcomere becomes ___________________ and ___________________________________________________ is not possible. 12. In the whole muscle, the degree of tension reflects ______________________ __________________________________________________________________ D. Summation 1. Twitches in a muscle can combine to become ___________________________ 2. Summation is ____________________________________________________ 3. Tetanic contractions are ____________________________________________ E. Recruitment of Motor Units 1. The _______________ in the motor units, the more ________________ the movements can be produced in a particular muscle. 2. All muscle fibers in a motor unit are stimulated at __________________ time. 3. Multiple motor unit summation is ____________________________________ 4. Recruitment is ___________________________________________________ 5. As the intensity of stimulation increases, ______________________________ continues until all possible motor units are activated in a muscle. F. Sustained Contractions 1. During sustained contractions, __________ tend to be recruited earlier. 2. The larger motor units respond ______________________________________ and more __________________________________________________________ 3. Muscle movements are smooth because _______________________________ __________________________________________________________________ 4. Muscle tone is ___________________________________________________ 5. Muscle tone is important for maintaining ______________________________ G. Types of Contractions 1. An isotonic contraction is __________________________________________ 2. A concentric contraction is _________________________________________ 3. An eccentric contraction is__________________________________________ 4. An isometric contraction is _________________________________________ 5. An example of an isometric contraction is _____________________________ 6. An example of an isotonic contraction is _______________________________ H. Fast and Slow Twitch Muscle Fibers 1. Type I fibers are __________________________________________________ 2. Examples of type I fibers are ________________________________________ 3. Type I fibers are _________________ in color because ___________________ __________________________________________________________________ 4. Type I fibers are resistant to_________________________________________ 5. Type IIa fibers are ________________________________________________ 6. Type IIa fibers are ________________ in color because __________________ __________________________________________________________________ 7. Type IIb fibers are ________________________________________________ 8. A muscles contain ____________________________________ of fiber types. IV. Smooth Muscles A. Smooth Muscle Fibers 1. Compared to skeletal muscle fibers, smooth muscle fibers _________________ __________________________________________________________________ 2. Two major types of smooth muscle are ________________________________ __________________________________________________________________ 3. Multiunit smooth muscle is located ___________________________________ __________________________________________________________________ 4. Visceral smooth muscle is located ____________________________________ __________________________________________________________________ 5. Fibers of visceral smooth muscle are connected by_______________________ 6. Rhythmicity is ___________________________________________________ 7. Peristalsis is _____________________________________________________ 8. Peristalsis helps force ______________________________________________ B. Smooth Muscle Contraction 1. Compared to skeletal muscle fibers, smooth muscle fibers lack _____________ and use _______________________________________ to bind calcium instead. 2. Two neurotransmitters that affect smooth muscle are _____________________ __________________________________________________________________ 3. Hormones affect smooth muscle by___________________________________ __________________________________________________________________ 4. Stretching of smooth muscle can trigger _______________________________ 5. Smooth muscle is _________________ to contract and ___________________ to relax than skeletal muscle. 6. Unlike skeletal muscle, smooth muscle fibers can change length without changing __________________________________________________________ V. Cardiac Muscle A. Cardiac muscle appears only in ___________________________________________ B. Cardiac muscle is composed of _______________________________________ cells, forming fibers that are ______________________________________________________ C. Cardiac muscle fibers can contract ___________________ then skeletal muscle fibers. D. Intercalated discs are ____________________________________________________ E. Intercalated discs allow muscle impulses to __________________________________ F. A syncytium is _________________________________________________________ VI. Skeletal Muscle Actions A. Introduction 1. Skeletal muscles generate __________________________________________ 2. The action of each muscle mostly depends upon_________________________ __________________________________________________________________ B. Origin and Insertion 1. The origin of a muscle is ___________________________________________ 2. The insertion of a muscle is _________________________________________ 3. When a muscle contracts, its ________________________________________ is pulled toward its __________________________________________________ 4. The head of a muscle is ____________________________________________ 5. The origins of the biceps brachii are __________________________________ 6. The insertion of the bicep brachii is ___________________________________ 7. When the biceps brachii contracts, the __________________________ bends. C. Interaction of Skeletal Muscles 1. A prime mover is ___________________________________________ 2. A synergist is ____________________________________________________ 3. An antagonist is __________________________________________________ VII. Major Skeletal Muscles A. Introduction 1. The name of a muscle may reflect ____________________________________ __________________________________________________________________ 2. An example of a muscle named for it size is ____________________________ 3. An example of a muscle named for its shape is __________________________ 4. An example of a muscle named for its function is________________________ 5. An example of a muscle named for its number of origins is ________________ 6. An example of a muscle named for its attachments site is _________________ __________________________________________________________________ 7. An example of a muscle named of the direction of its muscle fibers is _______ __________________________________________________________________ B. Muscles of Facial Expression 1. As a group, muscles of facial expression connect ________________________ __________________________________________________________________ 2. For the following muscles, list their origins, insertions, and actions. Epicranius Orbicularis oculi Orbicularis oris Buccinator Zygomaticus Platysma C. Muscles of Mastication 1. Muscles of mastication produce______________________ movements. 2. For the following muscles, list their origins, insertions, and actions Masseter Temporalis Medial pterygoid Lateral pterygoid D. Muscles That Move the Head and Vertebral Column 1. For the following muscles, list their origins, insertions, and actions Sternocleidomastoid Splenius capitis Semispinalis capitis Erector spinae E. Muscles That Move the Pectoral Girdle 1. For the following muscles, list their origins, insertions, and actions Trapezius Rhomboideus major Levator spinae Serratus anterior Pectoralis major F. Muscles That Move the Arm 1. For the following muscles, list their origins, insertions, and actions Coracobrachialis Pectoralis major Teres major Latissimus dorsi Supraspinatus Deltoid Subscapularis Infraspinatus Teres minor G. Muscles That Move the Forearm 1. For the following muscles, list their origins, insertions, and actions Biceps brachii Brachialis Brachioradialis Triceps brachii Supinator Pronator teres Pronator quadratus H. Muscles That Move the Hand 1. For the following muscles, list their origins, insertions, and actions Flexor carpi radialis longus Flexor carpi ulnaris Palmaris longus Flexor digitorum profundus Flexor digitorum superficialis Extensor carpi radialis Extensor carpi radialis brevic Extensor carpi ulnaris Extensor digitorum I. Muscles of the Abdominal Wall 1. The linea aspera is ________________________________________________ 2. For the following muscles, list their origins, insertions, and actions External oblique Internal oblique Transversus abdominis Rectus abdominis J. Muscles of the Pelvic Outlet 1. The pelvic diaphragm forms ________________________________________ 2. The urogenital diaphragm fills _______________________________________ 3. For the following muscles, list their origins, insertions, and actions Levator ani Coccygeus Superficial transversus perinea Bulbospongiosus Ischiocavernosus Sphincter urethra K. Muscles That Move the Thigh 1. For the following muscles, list their origins, insertions, and actions Psoas major Iliacus Gluteus maximus Gluteus medius Gluteus minimus Pectineus Adductor longus Adductor magnus Gracilis L. Muscles That Move the Leg 1. For the following muscles, list their origins, insertions, and actions Biceps femoris Semitendinosus Semimembranosus Sartorius Quadriceps femoris group M. Muscle That Move the Foot 1. For the following muscles, list their origins, insertions, and actions Tibialis anterior Fibularis tertius Extensor digitorum longus Gastrocnemius Soleus Flexor digitorum longus Tibialis posterior Fibularis longus VIII. Life-Span Changes A. Signs of aging of the muscular system begin to appear _________________________ B. At a microscopic level, ___________________________________________ decline. C. _______________________________________ being to replace some muscle tissue. D. _________________________________ can help maintain a healthy muscular system E. According to the National Institute on Aging, exercise should include _____________ ________________________________________________________________________