Muscle

Muscle is the contractile tissue of the body and is derived from the mesodermal layer of embryonic germ cells. Its function is to produce force and cause motion, either locomotion or movement within internal organs. Much of muscle contraction occurs without conscious thought and is necessary for survival, like the contraction of the heart or peristalsis, which pushes food through the digestive system. Voluntary muscle contraction is used to move the body and can be finely controlled, such as movements of the finger or gross movements like the quadriceps muscle of the thigh.

Types

There are three general types of muscle:

• Smooth muscle or "involuntary muscle" is found within structures such as the intestines, throat and blood vessels. It does not get tired.
• Cardiac muscle is also an "involuntary muscle" but it's a specialized kind of muscle found only within the heart. It does not get tired.
• Skeletal muscle or "voluntary muscle" is anchored by tendons to bone and is used to affect skeletal movement such as locomotion. It gets tired.

Cardiac and skeletal muscle are "striated" in that they contain sarcomeres and are packed into highly-regular arrangements of bundles; smooth muscle has neither. Striated muscle is often used 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 colour. It can carry more oxygen and sustain aerobic activity.
• Type II, fast twitch, muscle has three major kinds (Larsson et al. 1991. Am J Physiol. 261:C93-101) that are, in order of increasing contractile speed:
• a) Type IIa, which, like slow muscle, is aerobic, rich in mitochondria and capillaries and appears red.
• b) 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 a build-up of lactic acid in tissue begins to interfere with muscular contraction and causes pain. N.B. in some books and articles this muscle in humans was, confusingly, called type IIB (Smerdu et al. 1994. Am J Physiol. 267:C1723-8).
• c) Type IIb, which is anaerobic, glycolytic, "white" muscle that is even less dense in mitochondria and myoglobin. In small animals like rodents or rabbits this is the major fast muscle type, explaining the pale colour of their meat.

Anatomy

Muscle is composed of muscle cells (sometimes known as " muscle fibers"). Within the cells are myofibrils; myofibrils contain sarcomeres, which are composed of actin and myosin. Individual muscle cells are lined with endomysium. Muscle cells are bound together by perimysium into bundles called fascicles; the bundles are then grouped together to form muscle, which is lined by epimysium. Muscle spindles are distributed throughout the muscles and provide sensory feedback information to the central nervous system.

Skeletal muscle is arranged in discrete groups, examples of which include the biceps brachii. It is connected by tendons to processes of the skeleton. In contrast, smooth muscle occurs at various scales in almost every organ, from the skin (in which it controls erection of body hair) to the blood vessels and digestive tract (in which it controls the caliber of a lumen and peristalsis).

There are approximately 650 skeletal muscles in the human body (see list of muscles of the human body). Contrary to popular belief, the number of muscles cannot be increased through exercise; instead the muscle cells simply get bigger.

Physiology

The three types of muscle have significant differences, but all use the movement of actin against myosin to produce contraction and relaxation. In skeletal muscle, contraction is stimulated by electrical impulses transmitted by the nerves, the motor nerves and motoneurons in particular. All skeletal muscle and many smooth muscle contractions are facilitated by the neurotransmitter acetylcholine.

Muscles and muscular activity account for most of the body's energy consumption. Muscles store energy for their own use in the form of glycogen, which represents about 1% of their mass. This can be rapidly converted to glucose when more energy is necessary. Muscle can produce 3.35kW (4 1/2 horsepower) at full rate.

Nervous control

Efferent leg

Vertebrates move muscles in response to voluntary and autonomic signals from the brain. Deep muscles, superficial muscles, muscles of the face and internal muscles all correspond with dedicated regions in the brain.

In addition, muscles react to reflexive nerve stimuli that do not always send signals all the way to the brain, but most muscle activity is 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 loops 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

Sometimes known as muscle memory, 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 nucleus ruber in particular continuously sample position against movement and make minor corrections to assure a smooth projection.

Role in health and disease

Exercise

Exercise is often recommended as a means of improving motor skills, fitness and muscle strength. Exercise has several effects upon muscles, connective tissue and bone, and the nerves that stimulate the muscles.

Disease

Symptoms of muscle disease may include weakness or spasticity/rigidity, myoclonus (twitching) and myalgia (muscle pain). Diagnostic procedures that may reveal muscular disorders include testing creatine kinase levels in the blood and electromyography (measuring electrical activity in muscles).

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.

Diseases of the motor end plate include myasthenia gravis, a form of muscle weakness due to antibodies to the acetylcholine receptor, and its related condition Lambert-Eaton myasthenic syndrome (LEMS). Tetanus and botulism are bacterial infections in which bacterial toxins cause increased or decreased muscle tone, respectively.

The myopathies are all diseases affecting the muscle itself, rather than its nervous control.

Muscular dystrophy is a large group of diseases, many of them hereditary, where the muscle integrity is disrupted. It leads to progressive loss of strength, high dependence and decreased life span.

Inflammatory muscle disorders:

• Polymyalgia rheumatica (or "muscle rheumatism") is an inflammatory condition that mainly occurs in the elderly; it is associated with giant-cell arteritis. It often responds dramatically to glucocorticoids (e.g. prednisolone).
• Polymyositis, dermatomyositis and inclusion body myositis are autoimmune conditions in which the muscle is affected.

Rhabdomyolysis is the breakdown of muscular tissue due to any cause. While it may not lead to any muscular symptoms at all, the myoglobin thus released may cause acute renal failure.

Tumors of muscle include:

• Smooth muscle: leiomyoma (benign, very common in the uterus), leiomyosarcoma (malignant, very rare)
• Striated muscle: rhabdomyoma (benign) and rhabdomyosarcoma (malignant) - both very rare
• Metastasis from elsewhere (e.g. lung cancer)

Smooth muscle has been implicated to play a role in a large number of diseases affecting blood vessels, the respiratory tract (e.g., asthma), the digestive system (e.g. irritable bowel syndrome) and the urinary tract (e.g., urinary incontinence). These disease processes are not usually confined to the muscular tissue.

Injuries of muscles include wounds and strains.

The strongest human muscle

Depending on what definition of "strongest" is used, many different muscles in the human body can be characterized as being the "strongest."

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 975 lbf (4337 N) for two 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 at their belly. This is because the tension exerted by an individual skeletal (striated) muscle fiber does not vary much, either from muscle to muscle, or with length. Each fiber can exert a force on the order of 0.3 micronewtons. By this definition, the strongest muscle of the body is usually said to be the Quadriceps femoris or the Gluteus maximus.

Again taking strength to mean only "force" (in the physicist's sense, and as contrasted with " energy" or " power"), then a shorter muscle will be stronger "pound for pound" (i.e., by weight) than a longer muscle. The uterus may be the strongest muscle by weight in the human body. At the time when an infant is delivered, the human uterus weighs about 40 oz (1.1 kg). During childbirth, the uterus exerts 25 to 100 lbf (100 to 400 N) 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." Eye movements, however, probably do "need" to be exceptionally fast.

The unexplained 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 tongue may possibly be the strongest muscle at birth.

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 can "outwork" other muscles. An output of one watt continuously for seventy years yields a total work output of 2 to 3 ×10⁹ joules.

Efficiency

The efficiency of human muscle has been measured (in the context of rowing and cycling) at 14% to 27%. The efficiency is defined as the ratio of mechanical work done to the total energy output (heat plus work).

Muscle evolution

According to a recent study published in 1999 [1], specialized forms of skeletal and cardiac muscles predated the divergence of the vertebrate/ arthropod evolutionary line. This indicates that these types of muscle developed in a common ancestor sometime before 700 million years ago (mya). Vertebrate smooth muscle (smooth muscle found in humans) was found to have evolved independently from the skeletal and cardiac muscles.