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In This Chapter: Types of Human Muscle 62 Skeletal Muscle 62 Properties 62 Muscle Teamwork 64 Structure 64 Muscle Fibre Types 69 Nerve–Muscle Interaction 70 Motor Unit 70 Intra-muscle Coordination 72 Inter-muscle Coordination 72 Sport-specific Training 72 Muscle’s Adaptation to Strength Training 74 Summary 75 CHAPTER 3 CHAPTER Let’s explore muscle structure and function.... 3 Muscle Structure and Function After completing this chapter you should be able to: n describe the macro and micro structures of skeletal muscle; n describe muscle contraction and explain the sliding filament theory; n demonstrate an understanding of nerve–muscle interaction; n differentiate among types of muscle fibres; n describe group action of muscles; n discuss muscle’s adaptation to strength training. 61 62 Structure determines function. This is a statement that defines the essence of human anatomy and physiology. Muscle tissue – the contraction specialist – provides a prime example of how the structure of a tissue is well-adapted to perform a specific function. With approximately 660 muscles in the adult human body, comprising nearly half of our body weight, the importance of muscular activity is obvious. The various structures and types of muscle tissue support numerous life functions, such as ventilation, physical activity and exercise, digestion, and of course, pumping life-sustaining blood throughout the body via specialized cardiac muscle. The focus in this chapter will be on skeletal muscle, which permits voluntary movement, and is unique among other types of muscle in other important ways. We often look at muscle as a single entity, but in so doing, fail to recognize the molecular complexity and hierarchical structure of this tissue. This specialized structure enables muscle to shorten and develop tension, allowing a myriad of human movements to occur. From movements as simple as waving good-bye or picking up a book, to more complex actions such as those required in athletics, the muscular system is vital to our daily functioning. But how does muscle activity integrate with the nervous system to produce movement? And what are the fundamental contractile properties of muscle? Types of Human Muscle On the basis of their structures, contractile properties, and control mechanisms there are three types of muscle in the human body: (1) skeletal muscle; (2) smooth muscle; and (3) cardiac muscle. Most skeletal muscle is attached to bone and its contraction is responsible for supporting and moving the skeleton. The contraction of skeletal muscle is initiated by impulses in the motor neurons to the muscle and is usually under voluntary control. Smooth muscle is under the control of the Foundations of Exercise Science autonomic nervous system and is called involuntary. Smooth muscle forms the walls of blood vessels and body organs, such as the respiratory tract, the iris of the eye, and the gastrointestinal tract. The contractions of smooth muscle are slow and uniform and are very fatigue resistant. Smooth muscle functions to alter the activity of various body parts to meet the needs of the body at the time. Cardiac muscle, the muscle of the heart, has characteristics of both skeletal and smooth muscle. Cardiac muscle functions to provide the contractile activity of the heart and has its own intrinsic beat. Like skeletal muscle, the contractile activity of cardiac muscle can be graduated; however, cardiac muscle is very fatigue resistant. Like smooth muscle, the activation of cardiac muscle is involuntary. Although fitness training can benefit all three types of muscle systems, this chapter will deal primarily with the skeletal muscle. Skeletal Muscle Properties Skeletal muscle refers to a number of muscle fibres bound together by connective tissue and is usually linked to bone by bundles of collagen fibres, known as tendons. Tendons are located at each end of the muscle (Figure 3.1 A). During muscle contraction, skeletal muscle shortens, and as a result of the tendinous attachments to bone, functions to move the various parts of the skeleton with respect to one another (joints) to allow changes in position of one skeletal segment in relation to another. Positioning several muscles on each “side” of a joint allows movement in several planes, and through graded activation the speed and smoothness of the movement can be graduated. Skeletal muscles are capable of rapid contraction and relaxation. Intensive activity causes them to show early signs of fatigue. The assessment of the movement and the sequential pattern of muscle activation acting through joints to move Studying Human Movement and Health 63 Origin–Insertion In order for muscles to contract they must be attached to bones to create movement. This is accomplished by tendons, strong fibrous tissues at the ends of each muscle. The end of the muscle attached to the bone that does not move is called the origin, while the point of attachment of the muscle on the bone that moves is the insertion (Figure 3.1 A). The origin tends to be the more proximal attachment (closer to the body), while the insertion is the more distal attachment (further from the body). A Tendon (origin) Tendon (origin) Biceps Triceps Tendon (insertion) Tendon (insertion) B Biceps contracts Triceps contracts Extension Flexion Figure 3.1 Bending or straightening the elbow requires the coordinated interplay of the biceps and triceps muscles. 64 Foundations of Exercise Science body segments is termed biomechanics of human movement (see Chapter 7). Striated or Voluntary Muscle Muscle attached to the skeleton to make it move is known as skeletal muscle. It is also known as voluntary or striated muscle. Skeletal muscle is considered striated because of the alternating light and dark bands (created by the organization of the muscle fibres, or cells) that appear when viewed under a light microscope. Its description as voluntary comes from the fact that we can contract skeletal muscle when we want to, voluntarily, i.e., flex the biceps. Muscle Teamwork Muscles work in perfect synchrony. When one muscle contracts (draws together) to move a bone, the other relaxes, allowing the bone to move. The muscle or group of muscles producing a desired effect is known as the agonist, the prime mover. A muscle or group of muscles opposing the action is called an antagonist. An agonist–antagonist relationship occurs between the biceps and triceps of the upper arm. When the biceps (agonist) contracts to bend the elbow, the triceps (antagonist) relaxes and allows the bend. When the triceps (agonist) contracts to straighten the arm, the biceps (antagonist) in turn relaxes (Figure 3.1 B). The cooperation of biceps and triceps is typical of what takes place throughout the body. When entire groups of muscles get involved the interaction between agonist and antagonist muscles becomes more complex. The muscles surrounding the joint being moved and supporting it in the action are called synergists (complementing the action of a prime mover). Other muscle groups called fixators will steady joints closer to the body axis so that the desired action can occur. For example, if you want to climb a rope hand over hand, the muscles holding your shoulder girdle tightly to your rib cage are fixators, enabling you to use the muscles acting over the shoulder, elbow, wrist, and finger joints to perform their job and pull you up the rope. Structure Skeletal muscle is comprised of numerous cylinder-shaped cells called muscle fibres, and each fibre is made up of a number of myofilaments (Figure 3.2). The diameter of each fibre varies between 0.05 and 0.10 mm, with the length being dependent mainly on the distance between skeletal attachments (in the case of the biceps, the length of a fibre is approximately 15 cm). Each cell (fibre) is surrounded by a connective tissue sheath called the sarcolemma, and a variable number of fibres are enclosed together by a thicker connective tissue sheath to form a bundle of fibres (Figure 3.2 B). Each fibre contains not only the contractile machinery needed to develop force (Figure 3.3), but also the cell organelles necessary for cellular respiration (see Chapter 5, Energy for Muscular Activity). Also located outside each fibre is a supply of capillaries from which the cell obtains nutrients and eliminates waste. A large number of individual thread-like fibres known as myofibrils run lengthwise and parallel to one another within a muscle fibre. The myofibrils contain contractile units that are responsible for muscle contraction (Figure 3.2 D). Muscle’s Tug of War In some muscles, the individual fibres extend the entire length of the muscle, but in most the fibres are shorter. The shorter fibres, anchored to the connective-tissue network surrounding the muscle fibres, are placed at an angle to the longitudinal axis of the muscle. When muscle pulls on the bone during the transmission of force, it is like a number of people pulling on a rope, each person corresponding to a single fibre and the rope corresponding to the connective tissue and tendons. Studying Human Movement and Health 65 Muscle: The Contractile Machinery here, but you should still be able to appreciate all the intricate anatomical structures involved with every move we make. Within each myofibril, a number of contractile units, called sarcomeres (Figure 3.3 A), are organized in series, i.e., attached end to end. Each sarcomere is comprised of two types of protein myofilaments: myosin, the so-called thick filament, and actin, termed the thin filament. Looking at the filaments in a cross-section, i.e., looking at the myofilaments end-on, we see that each myosin filament is surrounded by actin filaments (Figure 3.3 A). Examining the sarcomere longitudinally, i.e., length-wise, we see the distinctive banding pattern (striations) characteristic of skeletal, or striated, muscle (Figure 3.4). Projecting out from each of the myosin filaments at an angle of approximately 45 degrees are tiny contractile elements called myosin bridges; from this view, these elements look similar to the projections of oars from a rowing shell (Figure 3.3). The Sliding Filament Theory D C B Muscle Belly Muscle Fibre Bundle Muscle Fibre Myofibril During the contraction of a muscle, it is the sliding of the thin actin filaments over the thick myosin filaments that causes shortening of the muscle to create movement. This phenomenon is called the sliding filament theory. It is far more complex than described A Figure 3.2 Components of skeletal muscle. A. Muscle belly (50 mm in diameter). B. Muscle fibre bundle (0.5 mm). C. Muscle fibre (0.05-0.1 mm). D. Myofibril (0.001-0.002 mm). 66 Foundations of Exercise Science Myofibril Sarcomere Sarcomere Actin filaments Myosin filament Myosin bridge A B C D Figure 3.3 Longitudinal section of a myofibril and simplified representation of muscular contraction: A. At rest. B. Contraction. C. Powerful stretching. D. Powerful contraction. Studying Human Movement and Health Rowing Simulation When a signal comes from the motor nerve activating the fibre, the heads of the myosin filaments temporarily attach themselves to the actin filaments (Figure 3.3 D), a process termed cross bridge formation. In a manner similar to the stroking of the oars, and the subsequent movement of a rowing shell, the movement of the cross bridges causes a movement of the actin filaments in relation to the myosin filaments, leading to shortening of the sarcomere. A single “stroke” shortens the sarcomere by approximately 1 percent of its length, and the nervous system is capable of activating cross bridge formation at a rate of 7-50 per second. Since the sarcomeres are attached to one another in series, the shortening of each sarcomere is additive. The total amount of fibre shortening amounts to some 25-40 percent of myofibril length. To produce an efficient rowing stroke, the oars must be optimally placed, i.e., reaching far enough, but not too far; similarly, for optimal cross bridge formation, the sarcomeres should be an optimal distance apart. For muscle contraction, this optimal distance is 0.0019-0.0022 mm. When the sarcomeres are separated by this distance, an optimal number of cross bridges can be formed per unit time. If the sarcomeres are farther apart, or closer together, than this optimal distance, then fewer cross bridges can be formed, resulting in less force development. If the sarcomeres are stretched 67 further apart, as occurs when the muscle is in a lengthened (i.e., extended or stretched) position (Figure 3.3 C), fewer cross bridges can form as the myosin projections have difficulty in reaching the actin filaments; this results in a decreased ability to produce force. When the sarcomeres are too close together, as would occur when the muscle is shortened (flexed), the cross bridges in fact interfere with one another as they try to form, resulting in a fewer number of effective cross bridges being formed, and again a decreased ability to develop force (Figure 3.3 D). The distance between sarcomeres depends on the state of muscle stretch, which in turn is a product of the position of the joint. What this means to the development of muscle force is that maximal force is developed when an optimal number of cross bridges are formed, which occurs at an optimal joint angle. Thus, as muscle force depends on muscle length, maximal muscle force occurs at optimal muscle length. As a joint moves through its range of motion, the muscle(s) connecting the two segments of the joint will move from a stretched position to a compressed position, and therefore, at some point in the movement, will pass through a position, termed the optimal joint angle, at which the muscle is at optimal length for maximal force development (Figure 3.5). This means that there would be an optimal joint angle for maximal force development Figure 3.4 High magnification view of a single sarcomere within a single myofibril. The characteristic “striped” appearance of skeletal muscle is a result of the arrangement of the actin and myosin strands. 68 Foundations of Exercise Science Maximal Force (kp) 80 60 40 20 0 30 60 90 120 150 Angle (degrees) Figure 3.5 Maximal muscle force changes continuously throughout elbow flexion according to the joint angle. Table 3.1 Relative involvement of muscle fibre types in sport events. Event Slow Twitch – Type I Fast Twitch – Type II 100-m sprint Low High 800-m run High High Marathon High Low Olympic weightlifting Low High Barbell squat Low High Soccer High High Field hockey High High Football wide receiver Low High Football lineman High High Basketball Low High Distance cycling High Low Studying Human Movement and Health 69 for each movement of a joint. Knowledge of the joint angle at which maximal force can be developed is important in the development of optimal biomechanics of the movement. Muscle Fibre Types There are also different types of skeletal muscle fibres. Some fibres can reach maximum tension more quickly than others. Based on this distinction, muscle fibres can be divided into the categories of fast twitch (FT or Type II) (also called white fibres based on their microscopic appearance) and slow twitch (ST or Type I) (also called red fibres based on their microscopic appearance). FT fibres are more anaerobic, larger, fatigue faster, and have a faster contraction speed than ST fibres. This makes these fibres ideal for actions that are short and require quick bursts of power and energy, such as sprinting or jumping. On the other hand, events that require endurance, such as long-distance running, swimming, or cycling, depend on the smaller, slower contracting, fatigue-resistant ST fibres that rely on oxygen (Table 3.1). There are also fibre types that fall in between these extremes with characteristics of both fibre types. Thus, Type II is further divided into Type IIa and Type IIb fibres. The distinction between the two is mainly in their contractile strength and their capacity for aerobicoxidative energy supply. The Type IIa fibres have greater capacity for aerobic metabolism and more capillaries surrounding them than Type IIb and therefore show greater resistance to fatigue. The muscle fibre composition of an individual is dictated through heredity. The fibre composition of an individual cannot be altered by training, i.e., the transformation of a fibre from one type to another as a result of the training stimulus does not occur. Most skeletal muscles, however, contain both FT Fast Twitch (Type II) Fibres (large diameter) Capillary Blood Vessels Slow Twitch (Type I) Fibres (small diameter) Figure 3.6 Muscle biopsy. 70 Foundations of Exercise Science and ST fibres, with the amount of each varying from one muscle to another, as well as among different individuals. Therefore, individual performance differences occur as a result of varying percentages of the muscle fibre types, making some individuals suited to some activities more than others (that is not to say that training will not improve what fibres you do have). information to the brain where it is processed and acted upon. This sensory information is also stored away so that future responses to similar input can be acted upon more quickly. The motor section (Figure 3.7 D) is involved directly in conducting the signals from the CNS to activate muscle contraction (see Chapter 17, Information Processing in Motor Learning). Muscle Biopsy Motor Unit Muscle fibre type is determined from a muscle biopsy. In this procedure, a small incision (about 5-7 mm) is made in the skin and fascia of the muscle following injection of a local anaesthetic into the muscle. A tiny piece of tissue is cut and removed from the muscle and then analyzed under a microscope (Figure 3.6). In addition to determining muscle fibre type, it is possible to study the metabolic characteristics of the muscle, and to assess changes in metabolic capability following various types of training programs. Biopsy = bio (life) + opsis (sight) Nerve–Muscle Interaction As with bone, muscle is a living tissue, and as such, is richly supplied with blood vessels and nerves. Skeletal muscle activation is initiated through neural activation (Figure 3.7), and therefore, is under conscious control. The nervous system is organized at two levels – the central (CNS) and peripheral (PNS) nervous systems, with the central system being composed of the brain and spinal cord (Figure 3.7 A), and the peripheral system being made up of numerous nerves of various sizes (Figure 3.7 C). The nervous system can also be divided in terms of function, namely motor and sensory activity. The sensory section (Figure 3.7 E) collects information from the various sensors located throughout the body and transmits the Motor nerves extend from the spinal cord to the muscle fibres. Each fibre is activated through impulses delivered via its motor end plate. A group of fibres activated via the same nerve is termed a motor unit, the basic functional entity of muscular activity. A muscle can be composed of a different number of motor units and each motor unit can in turn consist of a different number of muscle fibres. All muscle fibres of one particular motor unit, however, are always of the same fibre type (FT or ST fibres). Muscles that need to perform delicate and precise movements (the eye and finger muscles) generally consist of a large number of motor units (1,500-3,000), each containing only a few muscle fibres (8-50). Relatively unrefined movement, however, is carried out by muscles composed of fewer motor units with many fibres (approximately 600-2,000), each of which innervates up to 1,500 muscle fibres. In the tibialis anterior muscle, approximately 650 muscle fibres are innervated by each motor unit; in the gastrocnemius muscle, the number is approximately 1,600, and in the extensors of the back, it is about 2,000. The specific number of fibres in a motor unit of any given muscle can vary. The biceps may be composed of motor units that innervate 1,000, 1,200, 1,400, or 1,600 fibres. Furthermore, each muscle fibre can be innervated by only one motor unit. This cannot be altered through exercise. All-or-none Principle Muscle movement is controlled by the motor nerve Studying Human Movement and Health 71 impulses transmitted from the CNS and spinal cord out to the motor unit, which when activated causes the muscle fibres to contract. Whether or not a motor unit activates upon the arrival of an impulse depends upon the so-called all-or-none principle. This principle, discussed in more detail in Chapter 17, requires an impulse of a certain magnitude (or strength) to cause the innervated fibres to contract. The principle is analogous to firing a gun. Once a sufficient amount of pressure is placed on the trigger, the gun fires; pulling on the trigger harder will not cause the bullet to go faster or further. Activation Threshold Every motor unit has a specific threshold that must be reached for such activation to occur. For the biceps muscle, for example, all of the 1,500 fibres that may comprise B a single motor unit will contract maximally providing the nerve impulse has reached a certain magnitude. However, if the nerve impulse does not reach the required magnitude, then none of the fibres will contract. A weak nerve impulse activates only those motor units that have a low threshold of activation. A stronger nerve impulse will additionally activate motor units with higher thresholds. As the resistance increases, more motor units must be activated by stronger, more intensive impulses. An athlete needs increasingly more will power to exceed the excitatory thresholds of the motor units. This process is extremely fatiguing as a result of lactic acid accumulation in the muscle tissue and blood, the depletion of high-energy compounds, and the fatigue of the nervous system processes. C D E F A Figure 3.7 Sensory neurons transfer messages to the central nervous system, where they are analyzed and responded to by motor neurons. Activation of a motor unit and its innervation systems: A. Spinal cord. B. Cytosomes. C. Spinal nerve. D. Motor nerve. E. Sensory nerve. F. Muscle with muscle fibres. 72 Intra-muscle Coordination The capacity to activate motor units simultaneously is known as intra-muscle coordination. Although it is impossible to use all the motor units of a muscle at the same time, many highly trained power athletes, such as weightlifters, wrestlers, and shot-putters, are able to activate up to 85 percent of their available muscle fibres simultaneously, thus generating great strength. Untrained individuals, on the other hand, can normally activate only up to 60 percent of their fibres. Research has shown that under hypnosis a trained athlete can elevate the maximal force application for a given muscle by approximately 10 percent. The difference between assisted and voluntarily generated maximal force is regarded as the muscle force deficit of the muscle contraction. For untrained individuals, this deficit is much larger (approximately 20-35 percent). Trained athletes have not only a larger muscle mass than untrained individuals, but can also exploit a larger number of muscle fibres to produce force. However, for this reason, such athletes are more restricted than untrained individuals in further developing strength by improving intramuscle coordination. For this same reason, trained individuals can further increase strength only by increasing muscle diameter. Inter-muscle Coordination Any physical movement requires considerable effort by the muscles or muscle groups to master a given movement. This requires an optimal level of inter-muscle coordination. The interplay between muscles that generate movement through contraction, the agonists or prime movers, and muscles responsible for opposing movement, the antagonists, is of particular importance to the quality of intermuscle coordination. The cooperation between agonist and antagonist muscles during the bench Foundations of Exercise Science press, for example, provides a useful illustration. From a supine position, an athlete explosively stretches his or her arms against a high resistance. During the movement, a considerable number of motor units in the triceps and in cooperating muscles are synchronously activated, while the motor units of the antagonist muscles relax. The greater the participation of muscles and muscle groups, the higher the importance of inter-muscle coordination for strength capacity. To benefit from strength training, technically demanding sport-specific movements are often broken down into partial movements, so that the individual muscle groups responsible for these movements can be trained in relative isolation. The exercises used closely resemble the movement structure of the sport-specific movement, such that the training allows for the key muscle groups to be loaded relatively heavily. Sport-specific Training Consider the following exercises, which are beneficial to shot-putters: the bench press (Figure 3.8 A), lateral trunk curl (Figure 3.8 B), knee bend or squat (Figure 3.8 C), and heel or calf raise (Figure 3.8 D). An athlete whose muscles have been trained and developed in isolation using such exercises must subsequently engage in training that coordinates these muscles within the complete, sport-specific movement. Difficulties may occur if the athlete fails to develop all the relevant muscles in a balanced manner. For instance, a shot-putter who uses exercises that increase strength in only the arm and leg extensors, but not the trunk muscles, may experience major disturbances of inter-muscle coordination. As a result, performance may not improve or reach the level desired by the athlete. High-level inter-muscle coordination greatly improves strength performance and also enhances the flow, rhythm, and precision of movement. Unlike an ordinary individual, a highly trained athlete is able to translate strength potential more effectively into strength performance through enhanced inter-muscle coordination. Studying Human Movement and Health A B C D Figure 3.8 A shot-putter’s training includes exercises that work several prime movers in isolation. A. Bench press. B. Lateral trunk curl. C. Knee bend or squat. D. Heel or calf raise. 73 74 Foundations of Exercise Science Trainable vs. Non-trainable Factors The performance capacity of muscle is determined by several trainable and non-trainable factors. Trainable factors: Non-trainable factors: • • • • • • • • number of fibres • fibre structure (ST or FT fibres) fibre diameter intra-muscle coordination nerve impulse frequency inter-muscle coordination elasticity of muscle and its tendons energy stores of muscle and liver capillary density of muscle Muscle’s Adaptation to Strength Training In strength training, an individual’s performance improvements occur through a process of biological adaptation, which is reflected in the body’s increased strength. Similar types of adaptation may occur in any form of training. Indeed, they are the building blocks for improved performance in any athletic activity. In strength training, the adaptation process proceeds at different time rates for different functional systems and physiological processes. The adaptation depends on a variety of factors, in particular on intensity levels used in training and on an athlete’s unique biological make-up. Specific substances of the metabolism, such as enzymes, adapt within hours; the energy supply in the liver and muscle increases at a more moderate pace, within 10-14 days, by which time the first adaptations in the cardiovascular circulation also occur (see Chapter 6). The muscle mass increases slowly, within four to six weeks, its growth caused by an increase in the structural proteins in the skeletal muscle fibres. Recruitment of Muscle Fibres Proportion of Muscle Fibre % 100 Fibre Reserve 80 FT fibres, Glycolytic 60 FT fibres, Oxidative 40 ST fibres 20 20 40 60 Resistance 80 100 % Recruitment of muscle fibres during resistance work depends on the level of muscle tension. As the tension rises, more and more of the various fibre types are recruited into the movement as shown by the curves. Muscle tension below 25 percent of one’s maximal resistance recruits mostly ST fibres. At higher resistance, FT fibres also become active. Furthermore, which fibre is involved depends upon the muscle force that needs to be mobilized, and also the rate of acceleration of the mass to be moved. High accelerations of small loads and low accelerations of high loads require the intensive involvement of the FT fibres. Also, it is primarily the FT fibres that generate the explosive-type movements requiring a lot of strength. Studying Human Movement and Health Summary Muscles attached to skeletal bones work together and with tendons to enable body movement. Thin fibres called myofibrils constitute muscle, and end-to-end units called sarcomeres within each myofibril enable muscles to contract, causing movement in response to motor nerve stimulation. Motor nerves extend from the spinal cord to muscles throughout the body, and each motor unit is specific to either fast twitch or slow twitch muscle types. FT fibres, which are anaerobic in nature and fatigue faster than ST fibres, are best suited for activities requiring short bursts of 75 power and energy. Endurance events such as longdistance running, swimming, or cycling make use of the fatigue-resistant ST fibres that rely on oxygen. Motor units require threshold levels of nerve impulses before they can react – and some motor units have higher resistance thresholds than companion units in the same muscle. Movement requires precise coordination of muscles and the muscle fibres themselves. Intramuscle coordination is the ability to activate motor units simultaneously, while inter-muscle coordination refers to the synchronization of different muscles and muscle groups. Cooperation of the agonists and antagonists is necessary for smooth, controlled motion. Key Words Actin Agonist All-or-none principle Antagonist Biological adaptation Cardiac muscle Cross bridge formation Fast twitch (FT) fibre Fixator Insertion Inter-muscle coordination Intra-muscle coordination Involuntary muscle Motor end plate Motor unit Muscle biopsy Muscle fibre Muscle force deficit Myofibril Myofilament Myosin Origin Prime mover Sarcolemma Sarcomere Skeletal muscle Sliding filament theory Slow twitch (ST) fibre Smooth muscle Striated muscle Synergist Tendon Voluntary muscle Discussion Questions 1. What are the three types of muscle found in the human body? 4. What are the three types of muscle fibres? Give two characteristics of each type of fibre. 2. Describe the structure of a muscle from the largest structural unit to the smallest. 5. Explain nerve–muscle interaction. 3. Explain how the sarcomere contracts, resulting in muscle shortening. 6. Discuss the differences between inter- and intra-muscle coordination.