Muscle is one of the four primary tissue types of the body, and the body contains three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle (Figure). All three muscle tissues have some properties in common; they all exhibit a quality called excitability as their plasma membranes can change their electrical states (from polarized to depolarized) and send an electrical wave called an action potential along the entire length of the membrane. While the nervous system can influence the excitability of cardiac and smooth muscle to some degree, skeletal muscle completely depends on signaling from the nervous system to work properly. On the other hand, both cardiac muscle and smooth muscle can respond to other stimuli, such as hormones and local stimuli.

The muscles all begin the actual process of contracting (shortening) when a protein called actin is pulled by a protein called myosin. This occurs in striated muscle (skeletal and cardiac) after specific binding sites on the actin have been exposed in response to the interaction between calcium ions (Ca⁺⁺) and proteins (troponin and tropomyosin) that “shield” the actin-binding sites. Ca⁺⁺ also is required for the contraction of smooth muscle, although its role is different: here Ca⁺⁺ activates enzymes, which in turn activate myosin heads. All muscles require adenosine triphosphate (ATP) to continue the process of contracting, and they all relax when the Ca⁺⁺ is removed and the actin-binding sites are re-shielded.

A muscle can return to its original length when relaxed due to a quality of muscle tissue called elasticity. It can recoil back to its original length due to elastic fibers. Muscle tissue also has the quality of extensibility; it can stretch or extend. Contractility allows muscle tissue to pull on its attachment points and shorten with force.

Differences among the three muscle types include the microscopic organization of their contractile proteins—actin and myosin. The actin and myosin proteins are arranged very regularly in the cytoplasm of individual muscle cells (referred to as fibers) in both skeletal muscle and cardiac muscle, which creates a pattern, or stripes, called striations. The striations are visible with a light microscope under high magnification (Figure). Skeletal muscle fibers are multinucleated structures that compose the skeletal muscle. Cardiac muscle fibers each have one to two nuclei and are physically and electrically connected to each other so that the entire heart contracts as one unit (called a syncytium).

Because the actin and myosin are not arranged in such regular fashion in smooth muscle, the cytoplasm of a smooth muscle fiber (which has only a single nucleus) has a uniform, nonstriated appearance (resulting in the name smooth muscle). However, the less organized appearance of smooth muscle should not be interpreted as less efficient. Smooth muscle in the walls of arteries is a critical component that regulates blood pressure necessary to push blood through the circulatory system; and smooth muscle in the skin, visceral organs, and internal passageways is essential for moving all materials through the body.

Muscle is the tissue in animals that allows for active movement of the body or materials within the body. There are three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle. Most of the body’s skeletal muscle produces movement by acting on the skeleton. Cardiac muscle is found in the wall of the heart and pumps blood through the circulatory system.

Smooth muscle is found in the skin, where it is associated with hair follicles; it also is found in the walls of internal organs, blood vessels, and internal passageways, where it assists in moving materials.

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

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

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

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

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

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

Overview

Skeletal muscles contain connective tissue, blood vessels, and nerves. There are three layers of connective tissue: epimysium, perimysium, and endomysium. Skeletal muscle fibers are organized into groups called fascicles. Blood vessels and nerves enter the connective tissue and branch in the cell. Muscles attach to bones directly or through tendons or aponeuroses. Skeletal muscles maintain posture, stabilize bones and joints, control internal movement, and generate heat.

Skeletal muscle fibers are long, multinucleated cells. The membrane of the cell is the sarcolemma; the cytoplasm of the cell is the sarcoplasm. The sarcoplasmic reticulum (SR) is a form of endoplasmic reticulum. Muscle fibers are composed of myofibrils. The striations are created by the organization of actin and myosin resulting in the banding pattern of myofibrils.

The Greeks and Romans conducted the first studies done on the human body in Western culture. The educated class of subsequent societies studied Latin and Greek, and therefore the early pioneers of anatomy continued to apply Latin and Greek terminology or roots when they named the skeletal muscles. The large number of muscles in the body and unfamiliar words can make learning the names of the muscles in the body seem daunting, but understanding the etymology can help. Etymology is the study of how the root of a particular word entered a language and how the use of the word evolved over time. Taking the time to learn the root of the words is crucial to understanding the vocabulary of anatomy and physiology. When you understand the names of muscles it will help you remember where the muscles are located and what they do (Figure 1,2). Pronunciation of words and terms will take a bit of time to master, but after you have some basic information; the correct names and pronunciations will become easier.

Anatomists name the skeletal muscles according to a number of criteria, each of which describes the muscle in some way. These include naming the muscle after its shape, its size compared to other muscles in the area, its location in the body or the location of its attachments to the skeleton, how many origins it has, or its action.

The skeletal muscle’s anatomical location or its relationship to a particular bone often determines its name. For example, the frontalis muscle is located on top of the frontal bone of the skull. Similarly, the shapes of some muscles are very distinctive and the names, such as orbicularis, reflect the shape. For the buttocks, the size of the muscles influences the names: gluteus maximus (largest), gluteus medius (medium), and the gluteus minimus (smallest). Names were given to indicate length—brevis (short), longus (long)—and to identify position relative to the midline: lateralis (to the outside away from the midline), and medialis (toward the midline). The direction of the muscle fibers and fascicles are used to describe muscles relative to the midline, such as the rectus (straight) abdominis, or the oblique (at an angle) muscles of the abdomen.

Some muscle names indicate the number of muscles in a group. One example of this is the quadriceps, a group of four muscles located on the anterior (front) thigh. Other muscle names can provide information as to how many origins a particular muscle has, such as the biceps brachii. The prefix bi indicates that the muscle has two origins and tri indicates three origins.

The location of a muscle’s attachment can also appear in its name. When the name of a muscle is based on the attachments, the origin is always named first. For instance, the sternocleidomastoid muscle of the neck has a dual origin on the sternum (sterno) and clavicle (cleido), and it inserts on the mastoid process of the temporal bone. The last feature by which to name a muscle is its action. When muscles are named for the movement they produce, one can find action words in their name. Some examples are flexor (decreases the angle at the joint), extensor (increases the angle at the joint), abductor (moves the bone away from the midline), or adductor (moves the bone toward the midline).

Overview

Muscle names are based on many characteristics. The location of a muscle in the body is important. Some muscles are named based on their size and location, such as the gluteal muscles of the buttocks. Other muscle names can indicate the location in the body or bones with which the muscle is associated, such as the tibialis anterior. The shapes of some muscles are distinctive; for example, the direction of the muscle fibers is used to describe muscles of the body midline. The origin and/or insertion can also be features used to name a muscle; examples are the biceps brachii, triceps brachii, and the pectoralis major.

Muscles of the Posterior Neck and the Back

The posterior muscles of the neck are primarily concerned with head movements, like extension. The back muscles stabilize and move the vertebral column, and are grouped according to the lengths and direction of the fascicles.

The splenius muscles originate at the midline and run laterally and superiorly to their insertions. From the sides and the back of the neck, the splenius capitis inserts onto the head region, and the splenius cervicis extends onto the cervical region. These muscles can extend the head, laterally flex it, and rotate it (image).

The erector spinae group forms the majority of the muscle mass of the back and it is the primary extensor of the vertebral column. It controls flexion, lateral flexion, and rotation of the vertebral column, and maintains the lumbar curve. The erector spinae comprises the iliocostalis (laterally placed) group, the longissimus (intermediately placed) group, and the spinalis (medially placed) group.

The iliocostalis group includes the iliocostalis cervicis, associated with the cervical region; the iliocostalis thoracis, associated with the thoracic region; and the iliocostalis lumborum, associated with the lumbar region. The three muscles of the longissimus group are the longissimus capitis, associated with the head region; the longissimus cervicis, associated with the cervical region; and the longissimus thoracis, associated with the thoracic region. The third group, the spinalis group, comprises the spinalis capitis (head region), the spinalis cervicis (cervical region), and the spinalis thoracis (thoracic region).

The transversospinales muscles run from the transverse processes to the spinous processes of the vertebrae. Similar to the erector spinae muscles, the semispinalis muscles in this group are named for the areas of the body with which they are associated. The semispinalis muscles include the semispinalis capitis, the semispinalis cervicis, and the semispinalis thoracis. The multifidus muscle of the lumbar region helps extend and laterally flex the vertebral column.

Important in the stabilization of the vertebral column is the segmental muscle group, which includes the interspinales and intertransversarii muscles. These muscles bring together the spinous and transverse processes of each consecutive vertebra. Finally, the scalene muscles work together to flex, laterally flex, and rotate the head. They also contribute to deep inhalation. The scalene muscles include the anterior scalene muscle (anterior to the middle scalene), the middle scalene muscle (the longest, intermediate between the anterior and posterior scalenes), and the posterior scalene muscle (the smallest, posterior to the middle scalene).

Muscles of the Anterior Neck

The muscles of the anterior neck assist in deglutition (swallowing) and speech by controlling the positions of the larynx (voice box), and the hyoid bone, a horseshoe-shaped bone that functions as a solid foundation on which the tongue can move. The muscles of the neck are categorized according to their position relative to the hyoid bone (image). Suprahyoid muscles are superior to it, and the infrahyoid muscles are located inferiorly.

Figure 11.13 Muscles of the Anterior Neck The anterior muscles of the neck facilitate swallowing and speech. The suprahyoid muscles originate from above the hyoid bone in the chin region. The infrahyoid muscles originate below the hyoid bone in the lower neck.

The suprahyoid muscles raise the hyoid bone, the floor of the mouth, and the larynx during deglutition. These include the digastric muscle, which has anterior and posterior bellies that work to elevate the hyoid bone and larynx when one swallows; it also depresses the mandible. The stylohyoid muscle moves the hyoid bone posteriorly, elevating the larynx, and the mylohyoid muscle lifts it and helps press the tongue to the top of the mouth. The geniohyoid depresses the mandible in addition to raising and pulling the hyoid bone anteriorly.

The strap-like infrahyoid muscles generally depress the hyoid bone and control the position of the larynx. The omohyoid muscle, which has superior and inferior bellies, depresses the hyoid bone in conjunction with the sternohyoid and thyrohyoid muscles. The thyrohyoid muscle also elevates the larynx’s thyroid cartilage, whereas the sternothyroid depresses it to create different tones of voice.

Muscles That Move the Head

The head, attached to the top of the vertebral column, is balanced, moved, and rotated by the neck muscles (image). When these muscles act unilaterally, the head rotates. When they contract bilaterally, the head flexes or extends. The major muscle that laterally flexes and rotates the head is the sternocleidomastoid. In addition, both muscles working together are the flexors of the head. Place your fingers on both sides of the neck and turn your head to the left and to the right. You will feel the movement originate there. This muscle divides the neck into anterior and posterior triangles when viewed from the side (image).

Figure 11.14 Posterior and Lateral Views of the Neck The superficial and deep muscles of the neck are responsible for moving the head, cervical vertebrae, and scapulas.

Muscles That Move the Head

Table11.5

The movement of the eyeball is under the control of the extrinsic eye muscles, which originate outside the eye and insert onto the outer surface of the white of the eye. These muscles are located inside the eye socket and cannot be seen on any part of the visible eyeball (image and image). If you have ever been to a doctor who held up a finger and asked you to follow it up, down, and to both sides, he or she is checking to make sure your eye muscles are acting in a coordinated pattern.

Muscles That Move the Lower Jaw

In anatomical terminology, chewing is called mastication. Muscles involved in chewing must be able to exert enough pressure to bite through and then chew food before it is swallowed (image and image). The masseter muscle is the main muscle used for chewing because it elevates the mandible (lower jaw) to close the mouth, and it is assisted by the temporalis muscle, which retracts the mandible. You can feel the temporalis move by putting your fingers to your temple as you chew.

Although the masseter and temporalis are responsible for elevating and closing the jaw to break food into digestible pieces, the medial pterygoid and lateral pterygoid muscles provide assistance in chewing and moving food within the mouth.

The skeletal muscles are divided into axial (muscles of the trunk and head) and appendicular (muscles of the arms and legs) categories. This system reflects the bones of the skeleton system, which are also arranged in this manner. The axial muscles are grouped based on location, function, or both. Some of the axial muscles may seem to blur the boundaries because they cross over to the appendicular skeleton. The first grouping of the axial muscles you will review includes the muscles of the head and neck, then you will review the muscles of the vertebral column, and finally you will review the oblique and rectus muscles.

Overview

Muscles are either axial muscles or appendicular. The axial muscles are grouped based on location, function, or both. Some axial muscles cross over to the appendicular skeleton. The muscles of the head and neck are all axial. The muscles in the face create facial expression by inserting into the skin rather than onto bone. Muscles that move the eyeballs are extrinsic, meaning they originate outside of the eye and insert onto it. Tongue muscles are both extrinsic and intrinsic. The genioglossus depresses the tongue and moves it anteriorly; the styloglossus lifts the tongue and retracts it; the palatoglossus elevates the back of the tongue; and the hyoglossus depresses and flattens it. The muscles of the anterior neck facilitate swallowing and speech, stabilize the hyoid bone and position the larynx. The muscles of the neck stabilize and move the head. The sternocleidomastoid divides the neck into anterior and posterior triangles.

The muscles of the back and neck that move the vertebral column are complex, overlapping, and can be divided into five groups. The splenius group includes the splenius capitis and the splenius cervicis. The erector spinae has three subgroups. The iliocostalis group includes the iliocostalis cervicis, the iliocostalis thoracis, and the iliocostalis lumborum. The longissimus group includes the longissimus capitis, the longissimus cervicis, and the longissimus thoracis. The spinalis group includes the spinalis capitis, the spinalis cervicis, and the spinalis thoracis. The transversospinales include the semispinalis capitis, semispinalis cervicis, semispinalis thoracis, multifidus, and rotatores. The segmental muscles include the interspinales and intertransversarii. Finally, the scalenes include the anterior scalene, middle scalene, and posterior scalene.

Muscles That Create Facial Expression

The origins of the muscles of facial expression are on the surface of the skull (remember, the origin of a muscle does not move). The insertions of these muscles have fibers intertwined with connective tissue and the dermis of the skin. Because the muscles insert in the skin rather than on bone, when they contract, the skin moves to create facial expression (image).

The orbicularis oris is a circular muscle that moves the lips, and the orbicularis oculi is a circular muscle that closes the eye. The occipitofrontalis muscle moves up the scalp and eyebrows. The muscle has a frontal belly and an occipital (near the occipital bone on the posterior part of the skull) belly. In other words, there is a muscle on the forehead (frontalis) and one on the back of the head (occipitalis), but there is no muscle across the top of the head. Instead, the two bellies are connected by a broad tendon called the epicranial aponeurosis, or galea aponeurosis (galea = “apple”). The physicians originally studying human anatomy thought the skull looked like an apple.

A large portion of the face is composed of the buccinator muscle, which compresses the cheek. This muscle allows you to whistle, blow, and suck; and it contributes to the action of chewing. There are several small facial muscles, one of which is the corrugator supercilii, which is the prime mover of the eyebrows. Place your finger on your eyebrows at the point of the bridge of the nose. Raise your eyebrows as if you were surprised and lower your eyebrows as if you were frowning. With these movements, you can feel the action of the corrugator supercilli. Additional muscles of facial expression are presented in image.

Muscles That Move the Tongue

Although the tongue is obviously important for tasting food, it is also necessary for mastication, deglutition (swallowing), and speech (image and image). Because it is so moveable, the tongue facilitates complex speech patterns and sounds.

Figure 11.11 Muscles that Move the Tongue

Figure 11.12 Muscles for Tongue Movement, Swallowing, and Speech

Tongue muscles can be extrinsic or intrinsic. Extrinsic tongue muscles insert into the tongue from outside origins, and the intrinsic tongue muscles insert into the tongue from origins within it. The extrinsic muscles move the whole tongue in different directions, whereas the intrinsic muscles allow the tongue to change its shape (such as, curling the tongue in a loop or flattening it).

The extrinsic muscles all include the word root glossus (glossus = “tongue”), and the muscle names are derived from where the muscle originates. The genioglossus (genio = “chin”) originates on the mandible and allows the tongue to move downward and forward. The styloglossus originates on the styloid bone, and allows upward and backward motion. The palatoglossus originates on the soft palate to elevate the back of the tongue, and the hyoglossus originates on the hyoid bone to move the tongue downward and flatten it.

Everyday Connections

Anesthesia and the Tongue Muscles
Before surgery, a patient must be made ready for general anesthesia. The normal homeostatic controls of the body are put “on hold” so that the patient can be prepped for surgery. Control of respiration must be switched from the patient’s homeostatic control to the control of the anesthesiologist. The drugs used for anesthesia relax a majority of the body’s muscles.

Among the muscles affected during general anesthesia are those that are necessary for breathing and moving the tongue. Under anesthesia, the tongue can relax and partially or fully block the airway, and the muscles of respiration may not move the diaphragm or chest wall. To avoid possible complications, the safest procedure to use on a patient is called endotracheal intubation. Placing a tube into the trachea allows the doctors to maintain a patient’s (open) airway to the lungs and seal the airway off from the oropharynx. Post-surgery, the anesthesiologist gradually changes the mixture of the gases that keep the patient unconscious, and when the muscles of respiration begin to function, the tube is removed. It still takes about 30 minutes for a patient to wake up, and for breathing muscles to regain control of respiration. After surgery, most people have a sore or scratchy throat for a few days.

It is a complex job to balance the body on two feet and walk upright. The muscles of the vertebral column, thorax, and abdominal wall extend, flex, and stabilize different parts of the body’s trunk. The deep muscles of the core of the body help maintain posture as well as carry out other functions. The brain sends out electrical impulses to these various muscle groups to control posture by alternate contraction and relaxation. This is necessary so that no single muscle group becomes fatigued too quickly. If any one group fails to function, body posture will be compromised.

Overview

Made of skin, fascia, and four pairs of muscle, the anterior abdominal wall protects the organs located in the abdomen and moves the vertebral column. These muscles include the rectus abdominis, which extends through the entire length of the trunk, the external oblique, the internal oblique, and the transversus abdominus. The quadratus lumborum forms the posterior abdominal wall.

The muscles of the thorax play a large role in breathing, especially the dome-shaped diaphragm. When it contracts and flattens, the volume inside the pleural cavities increases, which decreases the pressure within them. As a result, air will flow into the lungs. The external and internal intercostal muscles span the space between the ribs and help change the shape of the rib cage and the volume-pressure ratio inside the pleural cavities during inspiration and expiration.

The perineum muscles play roles in urination in both sexes, ejaculation in men, and vaginal contraction in women. The pelvic floor muscles support the pelvic organs, resist intra-abdominal pressure, and work as sphincters for the urethra, rectum, and vagina.

Muscles of the Abdomen

There are four pairs of abdominal muscles that cover the anterior and lateral abdominal region and meet at the anterior midline. These muscles of the anterolateral abdominal wall can be divided into four groups: the external obliques, the internal obliques, the transversus abdominis, and the rectus abdominis (image and image).

The linea alba is a white, fibrous band that is made of the bilateral rectus sheaths that join at the anterior midline of the body. These enclose the rectus abdominis muscles (a pair of long, linear muscles, commonly called the “sit-up” muscles) that originate at the pubic crest and symphysis, and extend the length of the body’s trunk. Each muscle is segmented by three transverse bands of collagen fibers called the tendinous intersections. This results in the look of “six-pack abs,” as each segment hypertrophies on individuals at the gym who do many sit-ups.There are three flat skeletal muscles in the antero-lateral wall of the abdomen. The external oblique, closest to the surface, extend inferiorly and medially, in the direction of sliding one’s four fingers into pants pockets. Perpendicular to it is the intermediate internal oblique, extending superiorly and medially, the direction the thumbs usually go when the other fingers are in the pants pocket. The deep muscle, the transversus abdominis, is arranged transversely around the abdomen, similar to the front of a belt on a pair of pants. This arrangement of three bands of muscles in different orientations allows various movements and rotations of the trunk. The three layers of muscle also help to protect the internal abdominal organs in an area where there is no bone.

The posterior abdominal wall is formed by the lumbar vertebrae, parts of the ilia of the hip bones, psoas major and iliacus muscles, and quadratus lumborum muscle. This part of the core plays a key role in stabilizing the rest of the body and maintaining posture.

Career Connections

Physical Therapists
Those who have a muscle or joint injury will most likely be sent to a physical therapist (PT) after seeing their regular doctor. PTs have a master’s degree or doctorate, and are highly trained experts in the mechanics of body movements. Many PTs also specialize in sports injuries.

If you injured your shoulder while you were kayaking, the first thing a physical therapist would do during your first visit is to assess the functionality of the joint. The range of motion of a particular joint refers to the normal movements the joint performs. The PT will ask you to abduct and adduct, circumduct, and flex and extend the arm. The PT will note the shoulder’s degree of function, and based on the assessment of the injury, will create an appropriate physical therapy plan.

The first step in physical therapy will probably be applying a heat pack to the injured site, which acts much like a warm-up to draw blood to the area, to enhance healing. You will be instructed to do a series of exercises to continue the therapy at home, followed by icing, to decrease inflammation and swelling, which will continue for several weeks. When physical therapy is complete, the PT will do an exit exam and send a detailed report on the improved range of motion and return of normal limb function to your doctor. Gradually, as the injury heals, the shoulder will begin to function correctly. A PT works closely with patients to help them get back to their normal level of physical activity.

The pelvic floor is a muscular sheet that defines the inferior portion of the pelvic cavity. The pelvic diaphragm, spanning anteriorly to posteriorly from the pubis to the coccyx, comprises the levator ani and the ischiococcygeus. Its openings include the anal canal and urethra, and the vagina in women.

The large levator ani consists of two skeletal muscles, the pubococcygeus and the iliococcygeus (image). The levator ani is considered the most important muscle of the pelvic floor because it supports the pelvic viscera. It resists the pressure produced by contraction of the abdominal muscles so that the pressure is applied to the colon to aid in defecation and to the uterus to aid in childbirth (assisted by the ischiococcygeus, which pulls the coccyx anteriorly). This muscle also creates skeletal muscle sphincters at the urethra and anus.

Muscles of the Thorax

The muscles of the chest serve to facilitate breathing by changing the size of the thoracic cavity (image). When you inhale, your chest rises because the cavity expands. Alternately, when you exhale, your chest falls because the thoracic cavity decreases in size.

The Diaphragm

The change in volume of the thoracic cavity during breathing is due to the alternate contraction and relaxation of the diaphragm (image). It separates the thoracic and abdominal cavities, and is dome-shaped at rest. The superior surface of the diaphragm is convex, creating the elevated floor of the thoracic cavity. The inferior surface is concave, creating the curved roof of the abdominal cavity.

Defecating, urination, and even childbirth involve cooperation between the diaphragm and abdominal muscles (this cooperation is referred to as the “Valsalva maneuver”). You hold your breath by a steady contraction of the diaphragm; this stabilizes the volume and pressure of the peritoneal cavity. When the abdominal muscles contract, the pressure cannot push the diaphragm up, so it increases pressure on the intestinal tract (defecation), urinary tract (urination), or reproductive tract (childbirth).

The inferior surface of the pericardial sac and the inferior surfaces of the pleural membranes (parietal pleura) fuse onto the central tendon of the diaphragm. To the sides of the tendon are the skeletal muscle portions of the diaphragm, which insert into the tendon while having a number of origins including the xiphoid process of the sternum anteriorly, the inferior six ribs and their cartilages laterally, and the lumbar vertebrae and 12th ribs posteriorly.

The diaphragm also includes three openings for the passage of structures between the thorax and the abdomen. The inferior vena cava passes through the caval opening, and the esophagus and attached nerves pass through the esophageal hiatus. The aorta, thoracic duct, and azygous vein pass through the aortic hiatus of the posterior diaphragm.

The Intercostal Muscles

There are three sets of muscles, called intercostal muscles, which span each of the intercostal spaces. The principal role of the intercostal muscles is to assist in breathing by changing the dimensions of the rib cage (image).

The 11 pairs of superficial external intercostal muscles aid in inspiration of air during breathing because when they contract, they raise the rib cage, which expands it. The 11 pairs of internal intercostal muscles, just under the externals, are used for expiration because they draw the ribs together to constrict the rib cage. The innermost intercostal muscles are the deepest, and they act as synergists for the action of the internal intercostals.

Muscles of the shoulder and upper limb can be divided into four groups: muscles that stabilize and position the pectoral girdle, muscles that move the arm, muscles that move the forearm, and muscles that move the wrists, hands, and fingers. The pectoral girdle, or shoulder girdle, consists of the lateral ends of the clavicle and scapula, along with the proximal end of the humerus, and the muscles covering these three bones to stabilize the shoulder joint. The girdle creates a base from which the head of the humerus, in its ball-and-socket joint with the glenoid fossa of the scapula, can move the arm in multiple directions.

Overview

The clavicle and scapula make up the pectoral girdle, which provides a stable origin for the muscles that move the humerus. The muscles that position and stabilize the pectoral girdle are located on the thorax. The anterior thoracic muscles are the subclavius, pectoralis minor, and the serratus anterior. The posterior thoracic muscles are the trapezius, levator scapulae, rhomboid major, and rhomboid minor. Nine muscles cross the shoulder joint to move the humerus. The ones that originate on the axial skeleton are the pectoralis major and the latissimus dorsi. The deltoid, subscapularis, supraspinatus, infraspinatus, teres major, teres minor, and coracobrachialis originate on the scapula.

The forearm flexors include the biceps brachii, brachialis, and brachioradialis. The extensors are the triceps brachii and anconeus. The pronators are the pronator teres and the pronator quadratus. The supinator is the only one that turns the forearm anteriorly.

The extrinsic muscles of the hands originate along the forearm and insert into the hand in order to facilitate crude movements of the wrists, hands, and fingers. The superficial anterior compartment of the forearm produces flexion. These muscles are the flexor carpi radialis, palmaris longus, flexor carpi ulnaris, and the flexor digitorum superficialis. The deep anterior compartment produces flexion as well. These are the flexor pollicis longus and the flexor digitorum profundus. The rest of the compartments produce extension. The extensor carpi radialis longus, extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, and extensor carpi ulnaris are the muscles found in the superficial posterior compartment. The deep posterior compartment includes the abductor longus, extensor pollicis brevis, extensor pollicis longus, and the extensor indicis.

Finally, the intrinsic muscles of the hands allow our fingers to make precise movements, such as typing and writing. They both originate and insert within the hand. The thenar muscles, which are located on the lateral part of the palm, are the abductor pollicis brevis, opponens pollicis, flexor pollicis brevis, and adductor pollicis. The hypothenar muscles, which are located on the medial part of the palm, are the abductor digiti minimi, flexor digiti minimi brevis, and opponens digiti minimi. The intermediate muscles, located in the middle of the palm, are the lumbricals, palmar interossei, and dorsal interossei.

Muscles That Position the Pectoral Girdle

Muscles that position the pectoral girdle are located either on the anterior thorax or on the posterior thorax (image and image). The anterior muscles include the subclavius, pectoralis minor, and serratus anterior. The posterior muscles include the trapezius, rhomboid major, and rhomboid minor. When the rhomboids are contracted, your scapula moves medially, which can pull the shoulder and upper limb posteriorly.

Muscles That Move the Humerus

Similar to the muscles that position the pectoral girdle, muscles that cross the shoulder joint and move the humerus bone of the arm include both axial and scapular muscles (image and image). The two axial muscles are the pectoralis major and the latissimus dorsi. The pectoralis major is thick and fan-shaped, covering much of the superior portion of the anterior thorax. The broad, triangular latissimus dorsi is located on the inferior part of the back, where it inserts into a thick connective tissue shealth called an aponeurosis.

The rest of the shoulder muscles originate on the scapula. The anatomical and ligamental structure of the shoulder joint and the arrangements of the muscles covering it, allows the arm to carry out different types of movements. The deltoid, the thick muscle that creates the rounded lines of the shoulder is the major abductor of the arm, but it also facilitates flexing and medial rotation, as well as extension and lateral rotation. The subscapularis originates on the anterior scapula and medially rotates the arm. Named for their locations, the supraspinatus (superior to the spine of the scapula) and the infraspinatus (inferior to the spine of the scapula) abduct the arm, and laterally rotate the arm, respectively. The thick and flat teres major is inferior to the teres minor and extends the arm, and assists in adduction and medial rotation of it. The long teres minor laterally rotates and extends the arm. Finally, the coracobrachialis flexes and adducts the arm.

The tendons of the deep subscapularis, supraspinatus, infraspinatus, and teres minor connect the scapula to the humerus, forming the rotator cuff (musculotendinous cuff), the circle of tendons around the shoulder joint. When baseball pitchers undergo shoulder surgery it is usually on the rotator cuff, which becomes pinched and inflamed, and may tear away from the bone due to the repetitive motion of bring the arm overhead to throw a fast pitch.

Muscles That Move the Forearm

The forearm, made of the radius and ulna bones, has four main types of action at the hinge of the elbow joint: flexion, extension, pronation, and supination. The forearm flexors include the biceps brachii, brachialis, and brachioradialis. The extensors are the triceps brachii and anconeus. The pronators are the pronator teres and the pronator quadratus, and the supinator is the only one that turns the forearm anteriorly. When the forearm faces anteriorly, it is supinated. When the forearm faces posteriorly, it is pronated.

The biceps brachii, brachialis, and brachioradialis flex the forearm. The two-headed biceps brachii crosses the shoulder and elbow joints to flex the forearm, also taking part in supinating the forearm at the radioulnar joints and flexing the arm at the shoulder joint. Deep to the biceps brachii, the brachialis provides additional power in flexing the forearm. Finally, the brachioradialis can flex the forearm quickly or help lift a load slowly. These muscles and their associated blood vessels and nerves form the anterior compartment of the arm (anterior flexor compartment of the arm) (image and image).

Muscles That Move the Wrist, Hand, and Fingers

Wrist, hand, and finger movements are facilitated by two groups of muscles. The forearm is the origin of the extrinsic muscles of the hand. The palm is the origin of the intrinsic muscles of the hand.

Muscles of the Arm That Move the Wrists, Hands, and Fingers

The muscles in the anterior compartment of the forearm (anterior flexor compartment of the forearm) originate on the humerus and insert onto different parts of the hand. These make up the bulk of the forearm. From lateral to medial, the superficial anterior compartment of the forearm includes the flexor carpi radialis, palmaris longus, flexor carpi ulnaris, and flexor digitorum superficialis. The flexor digitorum superficialis flexes the hand as well as the digits at the knuckles, which allows for rapid finger movements, as in typing or playing a musical instrument (see image and image). However, poor ergonomics can irritate the tendons of these muscles as they slide back and forth with the carpal tunnel of the anterior wrist and pinch the median nerve, which also travels through the tunnel, causing Carpal Tunnel Syndrome. The deep anterior compartment produces flexion and bends fingers to make a fist. These are the flexor pollicis longus and the flexor digitorum profundus.

The muscles in the superficial posterior compartment of the forearm (superficial posterior extensor compartment of the forearm) originate on the humerus. These are the extensor radialis longus, extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, and the extensor carpi ulnaris.

The muscles of the deep posterior compartment of the forearm (deep posterior extensor compartment of the forearm) originate on the radius and ulna. These include the abductor pollicis longus, extensor pollicis brevis, extensor pollicis longus, and extensor indicis (see image).

The tendons of the forearm muscles attach to the wrist and extend into the hand. Fibrous bands called retinacula sheath the tendons at the wrist. The flexor retinaculum extends over the palmar surface of the hand while the extensor retinaculum extends over the dorsal surface of the hand.

Intrinsic Muscles of the Hand

The intrinsic muscles of the hand both originate and insert within it (image). These muscles allow your fingers to also make precise movements for actions, such as typing or writing. These muscles are divided into three groups. The thenar muscles are on the radial aspect of the palm. The hypothenar muscles are on the medial aspect of the palm, and the intermediate muscles are midpalmar.

The thenar muscles include the abductor pollicis brevis, opponens pollicis, flexor pollicis brevis, and the adductor pollicis. These muscles form the thenar eminence, the rounded contour of the base of the thumb, and all act on the thumb. The movements of the thumb play an integral role in most precise movements of the hand.

The hypothenar muscles include the abductor digiti minimi, flexor digiti minimi brevis, and the opponens digiti minimi. These muscles form the hypothenar eminence, the rounded contour of the little finger, and as such, they all act on the little finger. Finally, the intermediate muscles act on all the fingers and include the lumbrical, the palmar interossei, and the dorsal interossei.

The appendicular muscles of the lower body position and stabilize the pelvic girdle, which serves as a foundation for the lower limbs. Comparatively, there is much more movement at the pectoral girdle than at the pelvic girdle. There is very little movement of the pelvic girdle because of its connection with the sacrum at the base of the axial skeleton. The pelvic girdle is less range of motion because it was designed to stabilize and support the body.

Overview

The pelvic girdle attaches the legs to the axial skeleton. The hip joint is where the pelvic girdle and the leg come together. The hip is joined to the pelvic girdle by many muscles. In the gluteal region, the psoas major and iliacus form the iliopsoas. The large and strong gluteus maximus, gluteus medius, and gluteus minimus extend and abduct the femur. Along with the gluteus maximus, the tensor fascia lata muscle forms the iliotibial tract. The lateral rotators of the femur at the hip are the piriformis, obturator internus, obturator externus, superior gemellus, inferior gemellus, and quadratus femoris. On the medial part of the thigh, the adductor longus, adductor brevis, and adductor magnus adduct the thigh and medially rotate it. The pectineus muscle adducts and flexes the femur at the hip.

The thigh muscles that move the femur, tibia, and fibula are divided into medial, anterior, and posterior compartments. The medial compartment includes the adductors, pectineus, and the gracilis. The anterior compartment comprises the quadriceps femoris, quadriceps tendon, patellar ligament, and the sartorius. The quadriceps femoris is made of four muscles: the rectus femoris, the vastus lateralis, the vastus medius, and the vastus intermedius, which together extend the knee. The posterior compartment of the thigh includes the hamstrings: the biceps femoris, semitendinosus, and the semimembranosus, which all flex the knee.

The muscles of the leg that move the foot and toes are divided into anterior, lateral, superficial- and deep-posterior compartments. The anterior compartment includes the tibialis anterior, the extensor hallucis longus, the extensor digitorum longus, and the fibularis (peroneus) tertius. The lateral compartment houses the fibularis (peroneus) longus and the fibularis (peroneus) brevis. The superficial posterior compartment has the gastrocnemius, soleus, and plantaris; and the deep posterior compartment has the popliteus, tibialis posterior, flexor digitorum longus, and flexor hallucis longus.

Muscles of the Thigh

What would happen if the pelvic girdle, which attaches the lower limbs to the torso, were capable of the same range of motion as the pectoral girdle? For one thing, walking would expend more energy if the heads of the femurs were not secured in the acetabula of the pelvis. The body’s center of gravity is in the area of the pelvis. If the center of gravity were not to remain fixed, standing up would be difficult as well. Therefore, what the leg muscles lack in range of motion and versatility, they make up for in size and power, facilitating the body’s stabilization, posture, and movement.

Gluteal Region Muscles That Move the Femur

Most muscles that insert on the femur (the thigh bone) and move it, originate on the pelvic girdle. The psoas major and iliacus make up the iliopsoas group. Some of the largest and most powerful muscles in the body are the gluteal muscles or gluteal group. The gluteus maximus is the largest; deep to the gluteus maximus is the gluteus medius, and deep to the gluteus medius is the gluteus minimus, the smallest of the trio (image and image).

The tensor fascia latae is a thick, squarish muscle in the superior aspect of the lateral thigh. It acts as a synergist of the gluteus medius and iliopsoas in flexing and abducting the thigh. It also helps stabilize the lateral aspect of the knee by pulling on the iliotibial tract (band), making it taut. Deep to the gluteus maximus, the piriformis, obturator internus, obturator externus, superior gemellus, inferior gemellus, and quadratus femoris laterally rotate the femur at the hip.

The adductor longus, adductor brevis, and adductor magnus can both medially and laterally rotate the thigh depending on the placement of the foot. The adductor longus flexes the thigh, whereas the adductor magnus extends it. The pectineus adducts and flexes the femur at the hip as well. The pectineus is located in the femoral triangle, which is formed at the junction between the hip and the leg and also includes the femoral nerve, the femoral artery, the femoral vein, and the deep inguinal lymph nodes.

Thigh Muscles That Move the Femur, Tibia, and Fibula

Deep fascia in the thigh separates it into medial, anterior, and posterior compartments (see image and image). The muscles in the medial compartment of the thigh are responsible for adducting the femur at the hip. Along with the adductor longus, adductor brevis, adductor magnus, and pectineus, the strap-like gracilis adducts the thigh in addition to flexing the leg at the knee.

The muscles of the anterior compartment of the thigh flex the thigh and extend the leg. This compartment contains the quadriceps femoris group, which actually comprises four muscles that extend and stabilize the knee. The rectus femoris is on the anterior aspect of the thigh, the vastus lateralis is on the lateral aspect of the thigh, the vastus medialis is on the medial aspect of the thigh, and the vastus intermedius is between the vastus lateralis and vastus medialis and deep to the rectus femoris. The tendon common to all four is the quadriceps tendon (patellar tendon), which inserts into the patella and continues below it as the patellar ligament. The patellar ligament attaches to the tibial tuberosity. In addition to the quadriceps femoris, the sartorius is a band-like muscle that extends from the anterior superior iliac spine to the medial side of the proximal tibia. This versatile muscle flexes the leg at the knee and flexes, abducts, and laterally rotates the leg at the hip. This muscle allows us to sit cross-legged.

The posterior compartment of the thigh includes muscles that flex the leg and extend the thigh. The three long muscles on the back of the knee are the hamstring group, which flexes the knee. These are the biceps femoris, semitendinosus, and semimembranosus. The tendons of these muscles form the popliteal fossa, the diamond-shaped space at the back of the knee.

Muscles That Move the Feet and Toes

Similar to the thigh muscles, the muscles of the leg are divided by deep fascia into compartments, although the leg has three: anterior, lateral, and posterior (image and image).

The muscles in the anterior compartment of the leg: the tibialis anterior, a long and thick muscle on the lateral surface of the tibia, the extensor hallucis longus, deep under it, and the extensor digitorum longus, lateral to it, all contribute to raising the front of the foot when they contract. The fibularis tertius, a small muscle that originates on the anterior surface of the fibula, is associated with the extensor digitorum longus and sometimes fused to it, but is not present in all people. Thick bands of connective tissue called the superior extensor retinaculum (transverse ligament of the ankle) and the inferior extensor retinaculum, hold the tendons of these muscles in place during dorsiflexion.

The lateral compartment of the leg includes two muscles: the fibularis longus (peroneus longus) and the fibularis brevis (peroneus brevis). The superficial muscles in the posterior compartment of the leg all insert onto the calcaneal tendon (Achilles tendon), a strong tendon that inserts into the calcaneal bone of the ankle. The muscles in this compartment are large and strong and keep humans upright. The most superficial and visible muscle of the calf is the gastrocnemius. Deep to the gastrocnemius is the wide, flat soleus. The plantaris runs obliquely between the two; some people may have two of these muscles, whereas no plantaris is observed in about seven percent of other cadaver dissections. The plantaris tendon is a desirable substitute for the fascia lata in hernia repair, tendon transplants, and repair of ligaments. There are four deep muscles in the posterior compartment of the leg as well: the popliteus, flexor digitorum longus, flexor hallucis longus, and tibialis posterior.

The foot also has intrinsic muscles, which originate and insert within it (similar to the intrinsic muscles of the hand). These muscles primarily provide support for the foot and its arch, and contribute to movements of the toes (image and image). The principal support for the longitudinal arch of the foot is a deep fascia called plantar aponeurosis, which runs from the calcaneus bone to the toes (inflammation of this tissue is the cause of “plantar fasciitis,” which can affect runners. The intrinsic muscles of the foot consist of two groups. The dorsal group includes only one muscle, the extensor digitorum brevis. The second group is the plantar group, which consists of four layers, starting with the most superficial.

Two criteria to consider when classifying the types of muscle fibers are how fast some fibers contract relative to others, and how fibers produce ATP. Using these criteria, there are three main types of skeletal muscle fibers. Slow oxidative (SO) fibers contract relatively slowly and use aerobic respiration (oxygen and glucose) to produce ATP. Fast oxidative (FO) fibers have fast contractions and primarily use aerobic respiration, but because they may switch to anaerobic respiration (glycolysis), can fatigue more quickly than SO fibers. Lastly, fast glycolytic (FG) fibers have fast contractions and primarily use anaerobic glycolysis. The FG fibers fatigue more quickly than the others. Most skeletal muscles in a human contain(s) all three types, although in varying proportions.

The speed of contraction is dependent on how quickly myosin’s ATPase hydrolyzes ATP to produce cross-bridge action. Fast fibers hydrolyze ATP approximately twice as quickly as slow fibers, resulting in much quicker cross-bridge cycling (which pulls the thin filaments toward the center of the sarcomeres at a faster rate). The primary metabolic pathway used by a muscle fiber determines whether the fiber is classified as oxidative or glycolytic. If a fiber primarily produces ATP through aerobic pathways it is oxidative. More ATP can be produced during each metabolic cycle, making the fiber more resistant to fatigue. Glycolytic fibers primarily create ATP through anaerobic glycolysis, which produces less ATP per cycle. As a result, glycolytic fibers fatigue at a quicker rate.

The oxidative fibers contain many more mitochondria than the glycolytic fibers, because aerobic metabolism, which uses oxygen (O₂) in the metabolic pathway, occurs in the mitochondria. The SO fibers possess a large number of mitochondria and are capable of contracting for longer periods because of the large amount of ATP they can produce, but they have a relatively small diameter and do not produce a large amount of tension. SO fibers are extensively supplied with blood capillaries to supply O₂ from the red blood cells in the bloodstream. The SO fibers also possess myoglobin, an O₂-carrying molecule similar to O₂-carrying hemoglobin in the red blood cells. The myoglobin stores some of the needed O₂ within the fibers themselves (and gives SO fibers their red color). All of these features allow SO fibers to produce large quantities of ATP, which can sustain muscle activity without fatiguing for long periods of time.

The fact that SO fibers can function for long periods without fatiguing makes them useful in maintaining posture, producing isometric contractions, stabilizing bones and joints, and making small movements that happen often but do not require large amounts of energy. They do not produce high tension, and thus they are not used for powerful, fast movements that require high amounts of energy and rapid cross-bridge cycling.

FO fibers are sometimes called intermediate fibers because they possess characteristics that are intermediate between fast fibers and slow fibers. They produce ATP relatively quickly, more quickly than SO fibers, and thus can produce relatively high amounts of tension. They are oxidative because they produce ATP aerobically, possess high amounts of mitochondria, and do not fatigue quickly. However, FO fibers do not possess significant myoglobin, giving them a lighter color than the red SO fibers. FO fibers are used primarily for movements, such as walking, that require more energy than postural control but less energy than an explosive movement, such as sprinting. FO fibers are useful for this type of movement because they produce more tension than SO fibers but they are more fatigue-resistant than FG fibers.

FG fibers primarily use anaerobic glycolysis as their ATP source. They have a large diameter and possess high amounts of glycogen, which is used in glycolysis to generate ATP quickly to produce high levels of tension. Because they do not primarily use aerobic metabolism, they do not possess substantial numbers of mitochondria or significant amounts of myoglobin and therefore have a white color. FG fibers are used to produce rapid, forceful contractions to make quick, powerful movements. These fibers fatigue quickly, permitting them to only be used for short periods. Most muscles possess a mixture of each fiber type. The predominant fiber type in a muscle is determined by the primary function of the muscle.

Overview

ATP provides the energy for muscle contraction. The three mechanisms for ATP regeneration are creatine phosphate, anaerobic glycolysis, and aerobic metabolism. Creatine phosphate provides about the first 15 seconds of ATP at the beginning of muscle contraction. Anaerobic glycolysis produces small amounts of ATP in the absence of oxygen for a short period. Aerobic metabolism utilizes oxygen to produce much more ATP, allowing a muscle to work for longer periods. Muscle fatigue, which has many contributing factors, occurs when muscle can no longer contract. An oxygen debt is created as a result of muscle use. The three types of muscle fiber are slow oxidative (SO), fast oxidative (FO) and fast glycolytic (FG). SO fibers use aerobic metabolism to produce low power contractions over long periods and are slow to fatigue. FO fibers use aerobic metabolism to produce ATP but produce higher tension contractions than SO fibers. FG fibers use anaerobic metabolism to produce powerful, high-tension contractions but fatigue quickly.

The Sarcomere

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

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

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

Excitation-Contraction Coupling

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

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

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

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

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

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

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

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

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

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

The sequence of events that result in the contraction of an individual muscle fiber begins with a signal—the neurotransmitter, ACh—from the motor neuron innervating that fiber. The local membrane of the fiber will depolarize as positively charged sodium ions (Na⁺) enter, triggering an action potential that spreads to the rest of the membrane will depolarize, including the T-tubules. This triggers the release of calcium ions (Ca⁺⁺) from storage in the sarcoplasmic reticulum (SR). The Ca⁺⁺ then initiates contraction, which is sustained by ATP (image). As long as Ca⁺⁺ ions remain in the sarcoplasm to bind to troponin, which keeps the actin-binding sites “unshielded,” and as long as ATP is available to drive the cross-bridge cycling and the pulling of actin strands by myosin, the muscle fiber will continue to shorten to an anatomical limit.

Muscle contraction usually stops when signaling from the motor neuron ends, which repolarizes the sarcolemma and T-tubules, and closes the voltage-gated calcium channels in the SR. Ca⁺⁺ ions are then pumped back into the SR, which causes the tropomyosin to reshield (or re-cover) the binding sites on the actin strands. A muscle also can stop contracting when it runs out of ATP and becomes fatigued (image).

The molecular events of muscle fiber shortening occur within the fiber’s sarcomeres (see image). The contraction of a striated muscle fiber occurs as the sarcomeres, linearly arranged within myofibrils, shorten as myosin heads pull on the actin filaments.

The region where thick and thin filaments overlap has a dense appearance, as there is little space between the filaments. This zone where thin and thick filaments overlap is very important to muscle contraction, as it is the site where filament movement starts. Thin filaments, anchored at their ends by the Z-discs, do not extend completely into the central region that only contains thick filaments, anchored at their bases at a spot called the M-line. A myofibril is composed of many sarcomeres running along its length; thus, myofibrils and muscle cells contract as the sarcomeres contract.

Overview

A sarcomere is the smallest contractile portion of a muscle. Myofibrils are composed of thick and thin filaments. Thick filaments are composed of the protein myosin; thin filaments are composed of the protein actin. Troponin and tropomyosin are regulatory proteins.

Muscle contraction is described by the sliding filament model of contraction. ACh is the neurotransmitter that binds at the neuromuscular junction (NMJ) to trigger depolarization, and an action potential travels along the sarcolemma to trigger calcium release from SR. The actin sites are exposed after Ca⁺⁺ enters the sarcoplasm from its SR storage to activate the troponin-tropomyosin complex so that the tropomyosin shifts away from the sites. The cross-bridging of myposin heads docking into actin-binding sites is followed by the “power stroke”—the sliding of the thin filaments by thick filaments. The power strokes are powered by ATP. Ultimately, the sarcomeres, myofibrils, and muscle fibers shorten to produce movement.

To move an object, referred to as load, the sarcomeres in the muscle fibers of the skeletal muscle must shorten. The force generated by the contraction of the muscle (or shortening of the sarcomeres) is called muscle tension. However, muscle tension also is generated when the muscle is contracting against a load that does not move, resulting in two main types of skeletal muscle contractions: isotonic contractions and isometric contractions.

In isotonic contractions, where the tension in the muscle stays constant, a load is moved as the length of the muscle changes (shortens). There are two types of isotonic contractions: concentric and eccentric. A concentric contraction involves the muscle shortening to move a load. An example of this is the biceps brachii muscle contracting when a hand weight is brought upward with increasing muscle tension. As the biceps brachii contract, the angle of the elbow joint decreases as the forearm is brought toward the body. Here, the biceps brachii contracts as sarcomeres in its muscle fibers are shortening and cross-bridges form; the myosin heads pull the actin. An eccentric contraction occurs as the muscle tension diminishes and the muscle lengthens. In this case, the hand weight is lowered in a slow and controlled manner as the amount of cross-bridges being activated by nervous system stimulation decreases. In this case, as tension is released from the biceps brachii, the angle of the elbow joint increases. Eccentric contractions are also used for movement and balance of the body.

An isometric contraction occurs as the muscle produces tension without changing the angle of a skeletal joint. Isometric contractions involve sarcomere shortening and increasing muscle tension, but do not move a load, as the force produced cannot overcome the resistance provided by the load. For example, if one attempts to lift a hand weight that is too heavy, there will be sarcomere activation and shortening to a point, and ever-increasing muscle tension, but no change in the angle of the elbow joint. In everyday living, isometric contractions are active in maintaining posture and maintaining bone and joint stability. However, holding your head in an upright position occurs not because the muscles cannot move the head, but because the goal is to remain stationary and not produce movement. Most actions of the body are the result of a combination of isotonic and isometric contractions working together to produce a wide range of outcomes (image).

All of these muscle activities are under the exquisite control of the nervous system. Neural control regulates concentric, eccentric and isometric contractions, muscle fiber recruitment, and muscle tone. A crucial aspect of nervous system control of skeletal muscles is the role of motor units.

Overview

The number of cross-bridges formed between actin and myosin determines the amount of tension produced by a muscle. The length of a sarcomere is optimal when the zone of overlap between thin and thick filaments is greatest. Muscles that are stretched or compressed too greatly do not produce maximal amounts of power. A motor unit is formed by a motor neuron and all of the muscle fibers that are innervated by that same motor neuron. A single contraction is called a twitch. A muscle twitch has a latent period, a contraction phase, and a relaxation phase. A graded muscle response allows variation in muscle tension. Summation occurs as successive stimuli are added together to produce a stronger muscle contraction. Tetanus is the fusion of contractions to produce a continuous contraction. Increasing the number of motor neurons involved increases the amount of motor units activated in a muscle, which is called recruitment. Muscle tone is the constant low-level contractions that allow for posture and stability.

Skeletal Muscle Fibers

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

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

Physical training alters the appearance of skeletal muscles and can produce changes in muscle performance. Conversely, a lack of use can result in decreased performance and muscle appearance. Although muscle cells can change in size, new cells are not formed when muscles grow. Instead, structural proteins are added to muscle fibers in a process called hypertrophy , so cell diameter increases. The reverse, when structural proteins are lost and muscle mass decreases, is called atrophy . Age-related muscle atrophy is called sarcopenia . Cellular components of muscles can also undergo changes in response to changes in muscle use.

Hypertrophy is an increase in muscle mass due to the addition of structural proteins. The opposite of hypertrophy is atrophy, the loss of muscle mass due to the breakdown of structural proteins. Endurance exercise causes an increase in cellular mitochondria, myoglobin, and capillary networks in SO fibers. Endurance athletes have a high level of SO fibers relative to the other fiber types. Resistance exercise causes hypertrophy. Power-producing muscles have a higher number of FG fibers than of slow fibers. Strenuous exercise causes muscle cell damage that requires time to heal. Some athletes use performance-enhancing substances to enhance muscle performance. Muscle atrophy due to age is called sarcopenia and occurs as muscle fibers die and are replaced by connective and adipose tissue.

Cardiac muscle tissue is only found in the heart. Highly coordinated contractions of cardiac muscle pump blood into the vessels of the circulatory system. Similar to skeletal muscle, cardiac muscle is striated and organized into sarcomeres, possessing the same banding organization as skeletal muscle (image). However, cardiac muscle fibers are shorter than skeletal muscle fibers and usually contain only one nucleus, which is located in the central region of the cell. Cardiac muscle fibers also possess many mitochondria and myoglobin, as ATP is produced primarily through aerobic metabolism. Cardiac muscle fibers cells also are extensively branched and are connected to one another at their ends by intercalated discs. An intercalated disc allows the cardiac muscle cells to contract in a wave-like pattern so that the heart can work as a pump.

Intercalated discs are part of the sarcolemma and contain two structures important in cardiac muscle contraction: gap junctions and desmosomes. A gap junction forms channels between adjacent cardiac muscle fibers that allow the depolarizing current produced by cations to flow from one cardiac muscle cell to the next. This joining is called electric coupling, and in cardiac muscle it allows the quick transmission of action potentials and the coordinated contraction of the entire heart. This network of electrically connected cardiac muscle cells creates a functional unit of contraction called a syncytium. The remainder of the intercalated disc is composed of desmosomes. A desmosome is a cell structure that anchors the ends of cardiac muscle fibers together so the cells do not pull apart during the stress of individual fibers contracting (image).

Contractions of the heart (heartbeats) are controlled by specialized cardiac muscle cells called pacemaker cells that directly control heart rate. Although cardiac muscle cannot be consciously controlled, the pacemaker cells respond to signals from the autonomic nervous system (ANS) to speed up or slow down the heart rate. The pacemaker cells can also respond to various hormones that modulate heart rate to control blood pressure.

The wave of contraction that allows the heart to work as a unit, called a functional syncytium, begins with the pacemaker cells. This group of cells is self-excitable and able to depolarize to threshold and fire action potentials on their own, a feature called autorhythmicity ; they do this at set intervals which determine heart rate. Because they are connected with gap junctions to surrounding muscle fibers and the specialized fibers of the heart’s conduction system, the pacemaker cells are able to transfer the depolarization to the other cardiac muscle fibers in a manner that allows the heart to contract in a coordinated manner.

Another feature of cardiac muscle is its relatively long action potentials in its fibers, having a sustained depolarization “plateau.” The plateau is produced by Ca⁺⁺ entry though voltage-gated calcium channels in the sarcolemma of cardiac muscle fibers. This sustained depolarization (and Ca⁺⁺ entry) provides for a longer contraction than is produced by an action potential in skeletal muscle. Unlike skeletal muscle, a large percentage of the Ca⁺⁺ that initiates contraction in cardiac muscles comes from outside the cell rather than from the SR.

Overview

Cardiac muscle is striated muscle that is present only in the heart. Cardiac muscle fibers have a single nucleus, are branched, and joined to one another by intercalated discs that contain gap junctions for depolarization between cells and desmosomes to hold the fibers together when the heart contracts. Contraction in each cardiac muscle fiber is triggered by Ca⁺⁺ ions in a similar manner as skeletal muscle, but here the Ca⁺⁺ ions come from SR and through voltage-gated calcium channels in the sarcolemma. Pacemaker cells stimulate the spontaneous contraction of cardiac muscle as a functional unit, called a syncytium.

Smooth muscle (so-named because the cells do not have striations) is present in the walls of hollow organs like the urinary bladder, uterus, stomach, intestines, and in the walls of passageways, such as the arteries and veins of the circulatory system, and the tracts of the respiratory, urinary, and reproductive systems (imageab). Smooth muscle is also present in the eyes, where it functions to change the size of the iris and alter the shape of the lens; and in the skin where it causes hair to stand erect in response to cold temperature or fear.

Smooth muscle fibers are spindle-shaped (wide in the middle and tapered at both ends, somewhat like a football) and have a single nucleus; they range from about 30 to 200 μm (thousands of times shorter than skeletal muscle fibers), and they produce their own connective tissue, endomysium. Although they do not have striations and sarcomeres, smooth muscle fibers do have actin and myosin contractile proteins, and thick and thin filaments. These thin filaments are anchored by dense bodies. A dense body is analogous to the Z-discs of skeletal and cardiac muscle fibers and is fastened to the sarcolemma. Calcium ions are supplied by the SR in the fibers and by sequestration from the extracellular fluid through membrane indentations called calveoli.

Because smooth muscle cells do not contain troponin, cross-bridge formation is not regulated by the troponin-tropomyosin complex but instead by the regulatory protein calmodulin . In a smooth muscle fiber, external Ca⁺⁺ ions passing through opened calcium channels in the sarcolemma, and additional Ca⁺⁺ released from SR, bind to calmodulin. The Ca⁺⁺-calmodulin complex then activates an enzyme called myosin (light chain) kinase, which, in turn, activates the myosin heads by phosphorylating them (converting ATP to ADP and P_i, with the P_i attaching to the head). The heads can then attach to actin-binding sites and pull on the thin filaments. The thin filaments also are anchored to the dense bodies; the structures invested in the inner membrane of the sarcolemma (at adherens junctions) that also have cord-like intermediate filaments attached to them. When the thin filaments slide past the thick filaments, they pull on the dense bodies, structures tethered to the sarcolemma, which then pull on the intermediate filaments networks throughout the sarcoplasm. This arrangement causes the entire muscle fiber to contract in a manner whereby the ends are pulled toward the center, causing the midsection to bulge in a corkscrew motion (image).

Although smooth muscle contraction relies on the presence of Ca⁺⁺ ions, smooth muscle fibers have a much smaller diameter than skeletal muscle cells. T-tubules are not required to reach the interior of the cell and therefore not necessary to transmit an action potential deep into the fiber. Smooth muscle fibers have a limited calcium-storing SR but have calcium channels in the sarcolemma (similar to cardiac muscle fibers) that open during the action potential along the sarcolemma. The influx of extracellular Ca⁺⁺ ions, which diffuse into the sarcoplasm to reach the calmodulin, accounts for most of the Ca⁺⁺ that triggers contraction of a smooth muscle cell.

Muscle contraction continues until ATP-dependent calcium pumps actively transport Ca⁺⁺ ions back into the SR and out of the cell. However, a low concentration of calcium remains in the sarcoplasm to maintain muscle tone. This remaining calcium keeps the muscle slightly contracted, which is important in certain tracts and around blood vessels.

Because most smooth muscles must function for long periods without rest, their power output is relatively low, but contractions can continue without using large amounts of energy. Some smooth muscle can also maintain contractions even as Ca⁺⁺ is removed and myosin kinase is inactivated/dephosphorylated. This can happen as a subset of cross-bridges between myosin heads and actin, called latch-bridges , keep the thick and thin filaments linked together for a prolonged period, and without the need for ATP. This allows for the maintaining of muscle “tone” in smooth muscle that lines arterioles and other visceral organs with very little energy expenditure.

Smooth muscle is not under voluntary control; thus, it is called involuntary muscle. The triggers for smooth muscle contraction include hormones, neural stimulation by the ANS, and local factors. In certain locations, such as the walls of visceral organs, stretching the muscle can trigger its contraction (the stress-relaxation response).

Axons of neurons in the ANS do not form the highly organized NMJs with smooth muscle, as seen between motor neurons and skeletal muscle fibers. Instead, there is a series of neurotransmitter-filled bulges called varicosities as an axon courses through smooth muscle, loosely forming motor units (image). A varicosity releases neurotransmitters into the synaptic cleft. Also, visceral muscle in the walls of the hollow organs (except the heart) contains pacesetter cells. A pacesetter cell can spontaneously trigger action potentials and contractions in the muscle.

Smooth muscle is organized in two ways: as single-unit smooth muscle, which is much more common; and as multiunit smooth muscle. The two types have different locations in the body and have different characteristics. Single-unit muscle has its muscle fibers joined by gap junctions so that the muscle contracts as a single unit. This type of smooth muscle is found in the walls of all visceral organs except the heart (which has cardiac muscle in its walls), and so it is commonly called visceral muscle . Because the muscle fibers are not constrained by the organization and stretchability limits of sarcomeres, visceral smooth muscle has a stress-relaxation response . This means that as the muscle of a hollow organ is stretched when it fills, the mechanical stress of the stretching will trigger contraction, but this is immediately followed by relaxation so that the organ does not empty its contents prematurely. This is important for hollow organs, such as the stomach or urinary bladder, which continuously expand as they fill. The smooth muscle around these organs also can maintain a muscle tone when the organ empties and shrinks, a feature that prevents “flabbiness” in the empty organ. In general, visceral smooth muscle produces slow, steady contractions that allow substances, such as food in the digestive tract, to move through the body.

Multiunit smooth muscle cells rarely possess gap junctions, and thus are not electrically coupled. As a result, contraction does not spread from one cell to the next, but is instead confined to the cell that was originally stimulated. Stimuli for multiunit smooth muscles come from autonomic nerves or hormones but not from stretching. This type of tissue is found around large blood vessels, in the respiratory airways, and in the eyes.

Overview

Smooth muscle is found throughout the body around various organs and tracts. Smooth muscle cells have a single nucleus, and are spindle-shaped. Smooth muscle cells can undergo hyperplasia, mitotically dividing to produce new cells. The smooth cells are nonstriated, but their sarcoplasm is filled with actin and myosin, along with dense bodies in the sarcolemma to anchor the thin filaments and a network of intermediate filaments involved in pulling the sarcolemma toward the fiber’s middle, shortening it in the process. Ca⁺⁺ ions trigger contraction when they are released from SR and enter through opened voltage-gated calcium channels. Smooth muscle contraction is initiated when the Ca⁺⁺ binds to intracellular calmodulin, which then activates an enzyme called myosin kinase that phosphorylates myosin heads so they can form the cross-bridges with actin and then pull on the thin filaments. Smooth muscle can be stimulated by pacesetter cells, by the autonomic nervous system, by hormones, spontaneously, or by stretching. The fibers in some smooth muscle have latch-bridges, cross-bridges that cycle slowly without the need for ATP; these muscles can maintain low-level contractions for long periods. Single-unit smooth muscle tissue contains gap junctions to synchronize membrane depolarization and contractions so that the muscle contracts as a single unit. Single-unit smooth muscle in the walls of the viscera, called visceral muscle, has a stress-relaxation response that permits muscle to stretch, contract, and relax as the organ expands. Multiunit smooth muscle cells do not possess gap junctions, and contraction does not spread from one cell to the next.

Most muscle tissue of the body arises from embryonic mesoderm. Paraxial mesodermal cells adjacent to the neural tube form blocks of cells called somites . Skeletal muscles, excluding those of the head and limbs, develop from mesodermal somites, whereas skeletal muscle in the head and limbs develop from general mesoderm. Somites give rise to myoblasts. A myoblast is a muscle-forming stem cell that migrates to different regions in the body and then fuse(s) to form a syncytium, or myotube . As a myotube is formed from many different myoblast cells, it contains many nuclei, but has a continuous cytoplasm. This is why skeletal muscle cells are multinucleate, as the nucleus of each contributing myoblast remains intact in the mature skeletal muscle cell. However, cardiac and smooth muscle cells are not multinucleate because the myoblasts that form their cells do not fuse.

Gap junctions develop in the cardiac and single-unit smooth muscle in the early stages of development. In skeletal muscles, ACh receptors are initially present along most of the surface of the myoblasts, but spinal nerve innervation causes the release of growth factors that stimulate the formation of motor end-plates and NMJs. As neurons become active, electrical signals that are sent through the muscle influence the distribution of slow and fast fibers in the muscle.

Although the number of muscle cells is set during development, satellite cells help to repair skeletal muscle cells. A satellite cell is similar to a myoblast because it is a type of stem cell; however, satellite cells are incorporated into muscle cells and facilitate the protein synthesis required for repair and growth. These cells are located outside the sarcolemma and are stimulated to grow and fuse with muscle cells by growth factors that are released by muscle fibers under certain forms of stress. Satellite cells can regenerate muscle fibers to a very limited extent, but they primarily help to repair damage in living cells. If a cell is damaged to a greater extent than can be repaired by satellite cells, the muscle fibers are replaced by scar tissue in a process called fibrosis . Because scar tissue cannot contract, muscle that has sustained significant damage loses strength and cannot produce the same amount of power or endurance as it could before being damaged.

Smooth muscle tissue can regenerate from a type of stem cell called a pericyte , which is found in some small blood vessels. Pericytes allow smooth muscle cells to regenerate and repair much more readily than skeletal and cardiac muscle tissue. Similar to skeletal muscle tissue, cardiac muscle does not regenerate to a great extent. Dead cardiac muscle tissue is replaced by scar tissue, which cannot contract. As scar tissue accumulates, the heart loses its ability to pump because of the loss of contractile power. However, some minor regeneration may occur due to stem cells found in the blood that occasionally enter cardiac tissue.

Overview

Muscle tissue arises from embryonic mesoderm. Somites give rise to myoblasts and fuse to form a myotube. The nucleus of each contributing myoblast remains intact in the mature skeletal muscle cell, resulting in a mature, multinucleate cell. Satellite cells help to repair skeletal muscle cells. Smooth muscle tissue can regenerate from stem cells called pericytes, whereas dead cardiac muscle tissue is replaced by scar tissue. Aging causes muscle mass to decrease and be replaced by noncontractile connective tissue and adipose tissue.