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A top-down view of skeletal muscle

Muscle (from Latin musculus, diminutive of mus "mouse"^[1]) is the contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to produce force and cause motion. Muscles can cause either locomotion of the organism itself or movement of internal organs. Cardiac and smooth muscle contraction occurs without conscious thought and is necessary for survival. Examples are the contraction of the heart and peristalsis which pushes food through the digestive system. Voluntary contraction of the skeletal muscles is used to move the body and can be finely controlled. Examples are movements of the eye, or gross movements like the quadriceps muscle of the thigh. There are two broad types of voluntary muscle fibers: slow twitch and fast twitch. Slow twitch fibers contract for long periods of time but with little force while fast twitch fibers contract quickly and powerfully but fatigue very rapidly.

Muscles are predominately powered by the oxidation of fats and carbohydrates, but anaerobic chemical reactions are also used, particularly by fast twitch fibers. These chemical reactions produce adenosine triphosphate (ATP) molecules which are used to power the movement of the myosin heads.

Embryology

All muscles derive from paraxial mesoderm.^[2] The paraxial mesoderm is divided along the embryo's length into somites, corresponding to the segmentation of the body (most obviously seen in the vertebral column.^[2] Each somite has 3 divisions, sclerotome (which forms vertebrae), dermatome (which forms skin), and myotome (which forms muscle).^[2] The myotome is divided into two sections, the epimere and hypomere, which form epaxial and hypaxial muscles, respectively.^[2] Epaxial muscles in humans are only the erector spinae and small intervertebral muscles, and are innervated by the dorsal rami of the spinal nerves.^[2] All other muscles, including limb muscles, are hypaxial muscles, formed from the hypomere, and inervated by the ventral rami of the spinal nerves.^[2]

During development, myoblasts (muscle progenitor cells) either remain in the somite to form muscles associated with the vertebral column or migrate out into the body to form all other muscles.^[2] Myoblast migration is preceded by the formation of connective tissue frameworks, usually formed from the somatic lateral plate mesoderm.^[2] Myoblasts follow chemical signals to the appropriate locations, where they fuse into elongate skeletal muscle cells.^[2]

Types

Types of muscle (shown at different magnifications)

There are three types of muscle:

• Skeletal muscle or "voluntary muscle" is anchored by tendons (or by aponeuroses at a few places) to bone and is used to effect skeletal movement such as locomotion and in maintaining posture. Though this postural control is generally maintained as a subconscious reflex, the muscles responsible react to conscious control like non-postural muscles. An average adult male is made up of 42% of skeletal muscle and an average adult female is made up of 36% (as a percentage of body mass).^[3]
• Smooth muscle or "involuntary muscle" is found within the walls of organs and structures such as the esophagus, stomach, intestines, bronchi, uterus, urethra, bladder, blood vessels, and the arrector pili in the skin (in which it controls erection of body hair). Unlike skeletal muscle, smooth muscle is not under conscious control.
• Cardiac muscle is also an "involuntary muscle" but is more akin in structure to skeletal muscle, and is found only in the heart.

Cardiac and skeletal muscles are "striated" in that they contain sarcomeres and are packed into highly regular arrangements of bundles; smooth muscle has neither. While skeletal muscles are arranged in regular, parallel bundles, cardiac muscle connects at branching, irregular angles (called intercalated discs). Striated muscle contracts and relaxes in short, intense bursts, whereas smooth muscle sustains longer or even near-permanent contractions.

Skeletal muscle is further divided into several subtypes:

• Type I, slow oxidative, slow twitch, or "red" muscle is dense with capillaries and is rich in mitochondria and myoglobin, giving the muscle tissue its characteristic red color. It can carry more oxygen and sustain aerobic activity.
• Type II, fast twitch muscle, has three major kinds that are, in order of increasing contractile speed:^[4]
  • Type IIa, which, like slow muscle, is aerobic, rich in mitochondria and capillaries and appears red.
  • Type IIx (also known as type IId), which is less dense in mitochondria and myoglobin. This is the fastest muscle type in humans. It can contract more quickly and with a greater amount of force than oxidative muscle, but can sustain only short, anaerobic bursts of activity before muscle contraction becomes painful (often incorrectly attributed to a build-up of lactic acid). N.B. in some books and articles this muscle in humans was, confusingly, called type IIB.^[5]
  • Type IIb, which is anaerobic, glycolytic, "white" muscle that is even less dense in mitochondria and myoglobin. In small animals like rodents this is the major fast muscle type, explaining the pale color of their flesh.

Anatomy

The anatomy of muscles includes both gross anatomy, comprising all the muscles of an organism, and, on the other hand, microanatomy, which comprises the structures of a single muscle.

Gross anatomy

Muscles, anterior view (See Gray's muscle pictures for detailed pictures)

Muscles, posterior view (See Gray's muscle pictures for detailed pictures)

The gross anatomy of a muscle is the most important indicator of its role in the body. The action a muscle generates is determined by the origin and insertion locations. The cross-sectional area of a muscle (rather than volume or length) determines the amount of force it can generate by defining the number of sarcomeres which can operate in parallel. The amount of force applied to the external environment is determined by lever mechanics, specifically the ratio of in-lever to out-lever. For example, moving the insertion point of the biceps more distally on the radius (farther from the joint of rotation) would increase the force generated during flexion (and, as a result, the maximum weight lifted in this movement), but decrease the maximum speed of flexion. Moving the insertion point proximally (closer to the joint of rotation) would result in decreased force but increased velocity. This can be most easily seen by comparing the limb of a mole to a horse - in the former, the insertion point is positioned to maximize force (for digging), while in the latter, the insertion point is positioned to maximize speed (for running).

One particularly important aspect of gross anatomy of muscles is pennation or lack thereof. In most muscles, all the fibers are oriented in the same direction, running in a line from the origin to the insertion. In pennate muscles, the individual fibers are oriented at an angle relative to the line of action, attaching to the origin and insertion tendons at each end. Because the contracting fibers are pulling at an angle to the overall action of the muscle, the change in length is smaller, but this same orientation allows for more fibers (thus more force) in a muscle of a given size. Pennate muscles are usually found where their length change is less important than maximum force, such as the rectus femoris.

There are approximately 639 skeletal muscles in the human body. However, the exact number is difficult to define because different sources group muscles differently.

Microanatomy

Muscle is mainly composed of muscle cells. Within the cells are myofibrils; myofibrils contain sarcomeres, which are composed of actin and myosin. Individual muscle fibres are surrounded by endomysium. Muscle fibers are bound together by perimysium into bundles called fascicles; the bundles are then grouped together to form muscle, which is enclosed in a sheath of epimysium. Muscle spindles are distributed throughout the muscles and provide sensory feedback information to the central nervous system.

Skeletal muscle is arranged in discrete muscles, an example of which is the biceps brachii. It is connected by tendons to processes of the skeleton. Cardiac muscle is similar to skeletal muscle in both composition and action, being made up of myofibrils of sarcomeres, but anatomically different in that the muscle fibers are typically branched like a tree and connect to other cardiac muscle fibers through intercalcated discs, and form the appearance of a syncytium.

Physiology

The three types of muscle (skeletal, cardiac and smooth) have significant differences. However, all three use the movement of actin against myosin to create contraction. In skeletal muscle, contraction is stimulated by electrical impulses transmitted by the nerves, the motor nerves and motoneurons in particular. Cardiac and smooth muscle contractions are stimulated by internal pacemaker cells which regularly contract, and propagate contractions to other muscle cells they are in contact with. All skeletal muscle and many smooth muscle contractions are facilitated by the neurotransmitter acetylcholine.

Muscular activity accounts for much of the body's energy consumption. All muscle cells produce adenosine triphosphate (ATP) molecules which are used to power the movement of the myosin heads. Muscles conserve energy in the form of creatine phosphate which is generated from ATP and can regenerate ATP when needed with creatine kinase. Muscles also keep a storage form of glucose in the form of glycogen. Glycogen can be rapidly converted to glucose when energy is required for sustained, powerful contractions. Within the voluntary skeletal muscles, the glucose molecule can be metabolized anaerobically in a process called glycolysis which produces two ATP and two lactic acid molecules in the process (note that in aerobic conditions, lactate is not formed; instead pyruvate is formed and transmitted through the citric acid cycle). Muscle cells also contain globules of fat, which are used for energy during aerobic exercise. The aerobic energy systems take longer to produce the ATP and reach peak efficiency, and requires many more biochemical steps, but produces significantly more ATP than anaerobic glycolysis. Cardiac muscle on the other hand, can readily consume any of the three macronutrients (protein, glucose and fat) aerobically without a 'warm up' period and always extracts the maximum ATP yield from any molecule involved. The heart, liver and red blood cells will also consume lactic acid produced and excreted by skeletal muscles during exercise.

Nervous control

Efferent leg

The efferent leg of the peripheral nervous system is responsible for conveying commands to the muscles and glands, and is ultimately responsible for voluntary movement. Nerves move muscles in response to voluntary and autonomic (involuntary) signals from the brain. Deep muscles, superficial muscles, muscles of the face and internal muscles all correspond with dedicated regions in the primary motor cortex of the brain, directly anterior to the central sulcus that divides the frontal and parietal lobes.

In addition, muscles react to reflexive nerve stimuli that do not always send signals all the way to the brain. In this case, the signal from the afferent fiber does not reach the brain, but produces the reflexive movement by direct connections with the efferent nerves in the spine. However, the majority of muscle activity is volitional, and the result of complex interactions between various areas of the brain.

Nerves that control skeletal muscles in mammals correspond with neuron groups along the primary motor cortex of the brain's cerebral cortex. Commands are routed though the basal ganglia and are modified by input from the cerebellum before being relayed through the pyramidal tract to the spinal cord and from there to the motor end plate at the muscles. Along the way, feedback, such as that of the extrapyramidal system contribute signals to influence muscle tone and response.

Deeper muscles such as those involved in posture often are controlled from nuclei in the brain stem and basal ganglia.

Afferent leg

The afferent leg of the peripheral nervous system is responsible for conveying sensory information to the brain, primarily from the sense organs like the skin. In the muscles, the muscle spindles convey information about the degree of muscle length and stretch to the central nervous system to assist in maintaining posture and joint position. The sense of where our bodies are in space is called proprioception, the perception of body awareness. More easily demonstrated than explained, proprioception is the "unconscious" awareness of where the various regions of the body are located at any one time. This can be demonstrated by anyone closing their eyes and waving their hand around. Assuming proper proprioceptive function, at no time will the person lose awareness of where the hand actually is, even though it is not being detected by any of the other senses.

Several areas in the brain coordinate movement and position with the feedback information gained from proprioception. The cerebellum and red nucleus in particular continuously sample position against movement and make minor corrections to assure smooth motion.

Exercise

Exercise is often recommended as a means of improving motor skills, fitness, muscle and bone strength, and joint function. Exercise has several effects upon muscles, connective tissue, bone, and the nerves that stimulate the muscles. One such effect is muscle hypertrophy, an increase in size. This is used in bodybuilding.

Various exercises require a predominance of certain muscle fiber utilization over another. Aerobic exercise involves long, low levels of exertion in which the muscles are used at well below their maximal contraction strength for long periods of time (the most classic example being the marathon). Aerobic events, which rely primarily on the aerobic (with oxygen) system, use a higher percentage of Type I (or slow-twitch) muscle fibers, consume a mixture of fat, protein and carbohydrates for energy, consume large amounts of oxygen and produce little lactic acid. Anaerobic exercise involves short bursts of higher intensity contractions at a much greater percentage of their maximum contraction strength. Examples of anaerobic exercise include sprinting and weight lifting. The anaerobic energy delivery system uses predominantly Type II or fast-twitch muscle fibers, relies mainly on ATP or glucose for fuel, consumes relatively little oxygen, protein and fat, produces large amounts of lactic acid and can not be sustained for as long a period as aerobic exercise. The presence of lactic acid has an inhibitory effect on ATP generation within the muscle; though not producing fatigue, it can inhibit or even stop performance if the intracellular concentration becomes too high. However, long-term training causes neovascularization within the muscle, increasing the ability to move waste products out of the muscles and maintain contraction. Once moved out of muscles with high concentrations within the sarcomere, lactic acid can be used by other muscles or body tissues as a source of energy, or transported to the liver where it is converted back to pyruvate. The ability of the body to export lactic acid and use it as a source of energy depends on training level.^{[citation needed]}

Humans are genetically predisposed with a larger percentage of one type of muscle group over another. An individual born with a greater percentage of Type I muscle fibers would theoretically be more suited to endurance events, such as triathlons, distance running, and long cycling events, whereas a human born with a greater percentage of Type II muscle fibers would be more likely to excel at anaerobic events such as a 200 meter dash, or weightlifting.^{[citation needed]}

Delayed onset muscle soreness is pain or discomfort that may be felt one to three days after exercising and subsides generally within two to three days later. Once thought to be caused by lactic acid buildup, a more recent theory is that it is caused by tiny tears in the muscle fibers caused by eccentric contraction, or unaccustomed training levels. Since lactic acid disperses fairly rapidly, it could not explain pain experienced days after exercise.^[6]

Muscular, spinal and neural factors all affect muscle building. Sometimes a person may notice an increase in strength in a given muscle even though only its opposite has been subject to exercise, such as when a bodybuilder finds her left biceps stronger after completing a regimen focusing only on the right biceps. This phenomenon is called cross education.^{[citation needed]}

Disease

Symptoms of muscle diseases may include weakness, spasticity, myoclonus and myalgia. Diagnostic procedures that may reveal muscular disorders include testing creatine kinase levels in the blood and electromyography (measuring electrical activity in muscles). In some cases, muscle biopsy may be done to identify a myopathy, as well as genetic testing to identify DNA abnormalities associated with specific myopathies and dystrophies.

Neuromuscular diseases are those that affect the muscles and/or their nervous control. In general, problems with nervous control can cause spasticity or paralysis, depending on the location and nature of the problem. A large proportion of neurological disorders leads to problems with movement, ranging from cerebrovascular accident (stroke) and Parkinson's disease to Creutzfeldt-Jakob disease.

A non-invasive elastography technique that measures muscle noise is undergoing experimentation to provide a way of monitoring neuromuscular disease. The sound produced by a muscle comes from the shortening of actomyosin filaments along the axis of the muscle. During contraction, the muscle shortens along its longitudinal axis and expands across the transverse axis, producing vibrations at the surface.^[7]

Atrophy

There are many diseases and conditions which cause a decrease in muscle mass, known as muscle atrophy. Examples include cancer and AIDS, which induce a body wasting syndrome called cachexia. Other syndromes or conditions which can induce skeletal muscle atrophy are congestive heart disease and some diseases of the liver.

During aging, there is a gradual decrease in the ability to maintain skeletal muscle function and mass, known as sarcopenia. The exact cause of sarcopenia is unknown, but it may be due to a combination of the gradual failure in the "satellite cells" which help to regenerate skeletal muscle fibers, and a decrease in sensitivity to or the availability of critical secreted growth factors which are necessary to maintain muscle mass and satellite cell survival. Sarcopenia is a normal aspect of aging, and is not actually a disease state yet can be linked to many injuries in the elderly population as well as decreasing quality of life^[8].

Atrophy is of particular interest to the manned spaceflight community, since the weightlessness experienced in spaceflight results is a loss of as much as 30% of mass in some muscles^[9][10].

Physical inactivity and atrophy

Inactivity and starvation in mammals lead to atrophy of skeletal muscle, accompanied by a smaller number and size of the muscle cells as well as lower protein content.^[11] In humans, prolonged periods of immobilization, as in the cases of bed rest or astronauts flying in space, are known to result in muscle weakening and atrophy. Such consequences are also noted in small hibernating mammals like the golden-mantled ground squirrels and brown bats.^[12]

Bears are an exception to this rule; species in the family Ursidae are famous for their ability to survive unfavorable environmental conditions of low temperatures and limited nutrition availability during winter by means of hibernation. During that time, bears go through a series of physiological, morphological and behavioral changes.^[13] Their ability to maintain skeletal muscle number and size at time of disuse is of a significant importance.

During hibernation, bears spend four to seven months of inactivity and anorexia without undergoing muscle atrophy and protein loss.^[12] There are a few known factors that contribute to the sustaining of muscle tissue. During the summer period, bears take advantage of the nutrition availability and accumulate muscle protein. The protein balance at time of dormancy is also maintained by lower levels of protein breakdown during the winter time.^[12] At times of immobility, muscle wasting in bears is also suppressed by a proteolytic inhibitor that is released in circulation.^[11] Another factor that contributes to the sustaining of muscle strength in hibernating bears is the occurrence of periodic voluntary contractions and involuntary contractions from shivering during torpor.^[14] The three to four daily episodes of muscle activity are responsible for the maintenance of muscle strength and responsiveness in bears during hibernation.^[14]

Strength

A display of "strength" (e.g. lifting a weight) is a result of three factors that overlap: physiological strength (muscle size, cross sectional area, available crossbridging, responses to training), neurological strength (how strong or weak is the signal that tells the muscle to contract), and mechanical strength (muscle's force angle on the lever, moment arm length, joint capabilities). Contrary to popular belief, the number of muscle fibres cannot be increased through exercise; instead the muscle cells simply get bigger. Muscle fibres have a limited capacity for growth through hypertrophy and some believe they split through hyperplasia if subject to increased demand.^{[citation needed]}

The "strongest" human muscle

Since three factors affect muscular strength simultaneously and muscles never work individually, it is misleading to compare strength in individual muscles, and state that one is the "strongest". But below are several muscles whose strength is noteworthy for different reasons.

• In ordinary parlance, muscular "strength" usually refers to the ability to exert a force on an external object'for example, lifting a weight. By this definition, the masseter or jaw muscle is the strongest. The 1992 Guinness Book of Records records the achievement of a bite strength of 4,337 N (975 lb_f) for 2 seconds. What distinguishes the masseter is not anything special about the muscle itself, but its advantage in working against a much shorter lever arm than other muscles.
• If "strength" refers to the force exerted by the muscle itself, e.g., on the place where it inserts into a bone, then the strongest muscles are those with the largest cross-sectional area. This is because the tension exerted by an individual skeletal muscle fiber does not vary much. Each fiber can exert a force on the order of 0.3 micronewton. By this definition, the strongest muscle of the body is usually said to be the quadriceps femoris or the gluteus maximus.
• A shorter muscle will be stronger "pound for pound" (i.e., by weight) than a longer muscle. The myometrial layer of the uterus may be the strongest muscle by weight in the human body. At the time when an infant is delivered, the entire human uterus weighs about 1.1 kg (40 oz). During childbirth, the uterus exerts 100 to 400 N (25 to 100 lbf) of downward force with each contraction.
• The external muscles of the eye are conspicuously large and strong in relation to the small size and weight of the eyeball. It is frequently said that they are "the strongest muscles for the job they have to do" and are sometimes claimed to be "100 times stronger than they need to be." However, eye movements (particularly saccades used on facial scanning and reading) do require high speed movements, and eye muscles are exercised nightly during rapid eye movement sleep.
• The statement that "the tongue is the strongest muscle in the body" appears frequently in lists of surprising facts, but it is difficult to find any definition of "strength" that would make this statement true. Note that the tongue consists of sixteen muscles, not one.
• The heart has a claim to being the muscle that performs the largest quantity of physical work in the course of a lifetime. Estimates of the power output of the human heart range from 1 to 5 watts. This is much less than the maximum power output of other muscles; for example, the quadriceps can produce over 100 watts, but only for a few minutes. The heart does its work continuously over an entire lifetime without pause, and thus does "outwork" other muscles. An output of one watt continuously for eighty years yields a total work output of two and a half gigajoules.

Efficiency

The efficiency of human muscle has been measured (in the context of rowing and cycling) at 18% to 26%.^[15] The efficiency is defined as the ratio of mechanical work output to the total metabolic cost, as can be calculated from oxygen consumption. This low efficiency is the result of about 40% effiency of generating ATP from food energy, losses in converting energy from ATP into mechanical work inside the muscle, and mechanical losses inside the body. The latter two losses are dependent on the type of exercise and the type of muscle fibers being used (fast-twitch or slow-twitch). For an overal efficiency of 20 percent, one watt of mechanical power is equivalent to 4.3 kcal per hour. For example, a manufacturer of rowing equipment shows burned calories as four times the actual mechanical work, plus 300 kcal per hour,^[16] which amounts to about 20 percent efficiency at 250 watts of mechanical output.

Density of muscle tissue compared to adipose tissue

The density of mammalian skeletal muscle tissue is about 1.06 kg/liter^[17]. This can be contrasted with the density of adipose tissue (fat), which is 0.9196 kg/liter^[18]. This makes muscle tissue approximately 15% denser than fat tissue.

Resting energy expenditure of muscle

At rest, skeletal muscle consumes 54.4 kJ/kg (13.0 kcal/kg) per day. This is larger than adipose tissue (fat) at 18.8 kJ/kg (4.5 kcal/kg), and bone at 9.6 kJ/kg (2.3 kcal/kg).^[19]

Muscle evolution

Evolutionarily, specialized forms of skeletal and cardiac muscles predated the divergence of the vertebrate/arthropod evolutionary line.^[20] This indicates that these types of muscle developed in a common ancestor sometime before 700 million years ago (mya). Vertebrate smooth muscle was found to have evolved independently from the skeletal and cardiac muscles.

See also

Wikimedia Commons has media related to: muscles

• Atrophy
• Bodybuilding
• Cross education
• Electroactive polymers (materials that behave like muscles, used in robotics research)
• Fascia
• Hand strength
• List of muscles of the human body
• List of weight training exercises
• Muscle atrophy
• Muscle memory
• Muscle tone (residual muscle tension)
• Musculoskeletal system
• Muscular system
• Myopathy (pathology of muscle cells)
• Myotomy
• Phonomyography
• Preflexes
• Rapid plant movement
• Rohmert's law
• Soft tissue

References

External links

Look up muscle in Wiktionary, the free dictionary.

• Muslumova, Irada (2003). "Power of a Human Heart". The Physics Factbook. http://hypertextbook.com/facts/2003/IradaMuslumova.shtml.  (Heart output 1.3 to 5 watts, lifetime output 2 to 3 ×10⁹ joules)
• University of Dundee article on performing neurological examinations (Quadriceps "strongest")
• Muscle efficiency in rowing
• Human Muscle Tutorial (clear pictures of main human muscles and their Latin names, good for orientation)
• Microscopic stains of skeletal and cardiac muscular fibers to show striations. Note the differences in myofibrilar arrangements.