The Human Body

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Joint in anatomy is a structure that separates two or more adjacent elements of the skeletal system. Depending on the type of joint, such separated elements may or may not move on one another. This article discusses the joints of the human body—particularly their structure but also their ligaments, nerve and blood supply, and nutrition. Although the discussion focuses on human joints, its content is applicable to joints of vertebrates in general and mammals in particular. For information about the disorders and injuries that commonly affect human joints, see joint disease.

JOINT MOVEMENTS
In order to describe the main types of joint structures, it is helpful first to summarize the motions made possible by joints. These motions include spinning, swinging, gliding, rolling, and approximation.

Spin is a movement of a bone around its own long axis; it is denoted by the anatomical term rotation. An important example of spin is provided by the radius (outer bone of the forearm); this bone can spin upon the lower end of the humerus (upper arm) in all positions of the elbow. When an individual presses the back of the hand against the mouth, the forearm is pronated, or twisted; when the palm of the hand is pressed against the mouth, the forearm is supinated, or untwisted. Pronation is caused by medial (inward) rotation of the radius and supination by lateral (outward) rotation.
Swing, or angular movement, brings about a change in the angle between the long axis of the moving bone and some reference line in the fixed bone. Flexion (bending) and extension (straightening) of the elbow are examples of swing. A swing (to the right or left) of one bone away from another is called abduction; the reverse, adduction.
Approximation denotes the movement caused by pressing or pulling one bone directly toward another—i.e., by a "translation" in the physical sense. The reverse of approximation is separation. Gliding and rolling movements occur only within synovial joints and cause a moving bone to swing.
TYPES OF JOINT
Joints can be classified in two ways: temporally and structurally. Each classification is associated with joint function.
Considered temporally, joints are either transient or permanent. The bones of a transient joint fuse together sooner or later, but always after birth. All the joints of the skull, for example, are transient except those of the middle ear and those between the lower jaw and the braincase. The bones of a permanent joint do not fuse except as the result of disease or surgery. Such fusion is called arthrodesis. All permanent and some transient joints permit movement. Movement of the latter may be temporary, as with the roof bones of an infant's skull during birth, or long-term, as with the joints of the base of the skull during postnatal development.
There are two basic structural types of joint: diarthrosis, in which fluid is present, and synarthrosis, in which there is no fluid. All the diarthroses (commonly called synovial joints) are permanent. Some of the synarthroses are transient; others are permanent.
A . SYNARTHROSES
Synarthroses are divided into three classes: fibrous, symphysis, and cartilaginous.
Fibrous joints
In fibrous joints the articulating parts are separated by white connective tissue (collagen) fibres, which pass from one part to the other. There are two types of fibrous joints: suture and gomphosis.
A suture is formed by the fibrous covering, or periosteum, of two bones passing between them. In the adult, sutures are found only in the roof and sides of the braincase and in the upper part of the face. In the infant, however, the two halves of the frontal bone are separated by a suture (the metopic suture), as are the two halves of the mandible at the chin. Excepting those of the fetus and newborn infant, all sutures are narrow. In the late fetus and the newborn child, the sagittal suture, which separates the right and left halves of the roof of the skull, is quite wide and markedly so at its anterior and posterior ends. This enables one of the halves to glide over the other during the passage of the child through the mother's pelvis during birth, thus reducing the width of its skull, a process called molding. (The effects of molding usually disappear quickly.) After birth, all sutures become immobile joints. The expanded anterior and posterior ends of the sagittal suture are called fontanels; they lie immediately above a large blood channel (superior sagittal sinus).
Sutures are transient; they are unossified parts of the skeleton that become fused at various times from childhood to old age. The fusion is effected by direct conversion of the sutures into bone. Until maturity the sutures are active sites of growth of the bones they separate.
A gomphosis is a fibrous mobile peg-and-socket joint. The roots of the teeth (the pegs) fit into their sockets in the mandible and maxilla and are the only examples of this type of joint. Bundles of collagen fibres pass from the wall of the socket to the root; they are part of the circumdental, or periodontal, membrane. There is just enough space between the root and its socket to permit the root to be pressed a little farther into the socket during biting or chewing. Gomphoses are permanent joints in the sense that they last as long as do the roots of the teeth—unless, of course, they are damaged by disease.
The movement of the root within a gomphosis has a threefold effect. It lessens some of the impact between the upper and lower teeth in biting; it pumps blood and lymph from the periodontal membrane into the dental veins and lymph channels; and it stimulates sensory nerve terminals in the membrane to send signals to the brain centres that control the muscles of mastication.
Symphyses

A symphysis (fibrocartilaginous joint) is a joint in which the body (physis) of one bone meets the body of another. All but two of the symphyses lie in the vertebral (spinal) column, and all but one contain fibrocartilage as a constituent tissue. The short-lived suture between the two halves of the mandible is called the symphysis menti (from the Latin mentum, meaning "chin") and is the only symphysis devoid of fibrocartilage. All of the other symphyses are permanent.
The symphysis pubis joins the bodies of the two pubic bones of the pelvis. The adjacent sides of these bodies are covered by cartilage through which collagen fibres run from one pubis to the other. On their way they traverse a plate of cartilage, which in some instances (especially in the female) may contain a small cavity filled with fluid. Surrounding the joint and attached to the bones is a coat of fibrous tissue, particularly thick below (the subpubic ligament). The joint is flexible enough to act as a hinge that allows each of the two hip bones to swing a little upward and outward, as the ribs do during inspiration of air. This slight movement is increased in a woman during childbirth because of the infiltration of the joint and its fibrous coat by fluid toward the end of pregnancy; the fluid makes the joint even more flexible. In both sexes the joint acts as a buffer against shock transmitted to the pelvic bones from the legs in running and jumping.
The symphysis between the bodies of two adjacent vertebrae is called an intervertebral disk. It is composed of two parts: a soft centre (nucleus pulposus) and a tough flexible ring (anulus fibrosus) around it. The centre is a jellylike (mucoid) material containing a few cells derived from the precursor of the spine (notochord) of the embryo. The ring consists of collagen fibres arranged in concentric layers like those of an onion bulb. These fibres reach the adjacent parts of the vertebral bodies and are attached firmly to them.
There are 23 intervertebral disks, one between each pair of vertebrae below the first cervical vertebra, or atlas, and above the second sacral vertrebra (just above the tailbone). The lumbar (lower back) disks are thickest, the thoracic (chest or upper back) are thinnest, and the cervical are of intermediate size. These differences are associated with the function of the disks. In general, these disks have two functions: to allow movement between pairs of vertebrae and to act as buffers against shock caused by running, jumping, and other stresses applied to the spine.
If an intervertebral disk were the only joint between a pair of vertebrae, then one of these could move on the other in any direction; but each pair of vertebrae with an intervertebral disk also has a pair of synovial joints, one on each side of the vertebral (neural) arch. These joints limit the kinds of independent movement possible, so that the thoracic vertebrae move in only two directions and the lumbar in only three; only the cervical vertebrae below the atlas have full freedom of movement.
All intervertebral disks allow approximation and separation of their adjacent vertebrae. This is caused partly by movement brought about by muscle action and partly by the weight of the head and the trunk transmitted to the pelvis when a person is upright. The effect of weight is of special importance. The mucoid substance in the centre of the disk behaves like a fluid. It is acted upon by the person's weight and any other pressure forces transmitted along the spine. Therefore, the disk flattens from above downward and expands in all other directions. After arising in the morning and as the day progresses, a person decreases in height because of this compression of the disks. An average decrease of one millimetre in the height of each disk would mean an overall shortening of 2.3 centimetres, or about an inch. The spine lengthens again, of course, during sleep.
In the infant the greater part of the disk consists of the soft centre. Later the fibrous ring becomes relatively thicker in such a way that the soft part is nearer to the back of the disk. As middle age approaches, there is an increase in the fibrous element, the soft centre is reduced in size, and the amount of cartilage is increased. There is a tendency for the posterior part of the fibrous ring to degenerate in such a way that a sudden violent pressure may rupture the disk and allow the central part to protrude backward against the spinal cord; this condition is commonly referred to as slipped disk.
Cartilaginous joints
These joints, also called synchondroses, are the unossified masses between bones or parts of bones that pass through a cartilaginous stage before ossification. Examples are the synchondroses between the occipital and sphenoid bones and between the sphenoid and ethmoid bones of the floor of the skull. As already stated, these permit growth of the adjacent bones and act as virtual hinges at which the ethmoid and occipital bones swing upward upon the sphenoid; this allows backward growth of the nose and jaws during postnatal life. The juxta-epiphyseal plates separating the ossifying parts of a bone are also an example. Growth of the whole bone takes place at these plates when they appear, usually after birth. All synchondroses are transient, and all normally have vanished by the age of 25.
B. DIARTHROSES
Structure and elements of synovial joints
The synovial bursas are closed, thin-walled sacs, lined with synovial membrane. Bursas are found between structures that glide upon each other, and all motion at diarthroses entails some gliding, the amount varying from one joint to another. The bursal fluid, exuded by the synovial membrane, is called synovia, hence the common name for this class of joints. Two or more parts of the bursal wall become cartilage (chondrify) during prenatal life. These are the parts of the bursa that are attached to the articulating bones, and they constitute the articular cartilage of the bones.
A synovial joint consists of a wall enclosing a joint cavity that is wholly filled with synovial fluid. The wall consists of two layers: an outer complete fibrous layer and an inner incomplete synovial layer. Parts of the outer layer are either chondrified as articular cartilages or partly ossified as sesamoid bones (small, flat bones developed in tendons that move over bony surfaces). Parts of the synovial layer project into the cavity to form fatty pads. In a few diarthroses the fibrous layer also projects inward to become intra-articular disks, or menisci. These various structures will be discussed in connection with the layer to which they belong.
The fibrous layer: The fibrous layer is composed of collagen. The part that is visible in an unopened joint cavity is referred to as the investing ligament or joint capsule. At the point where it reaches the articulating bones, it attaches to the periosteum lining the outer surface of the cortex.
Articular cartilage: Articular cartilage (cartilage that covers the articulating part of a bone) is of the type called hyaline (glasslike) because thin sections of it are translucent, even transparent. Unlike bone, it is easily cut by a sharp knife. It is deformable but elastic, and it recovers its shape quickly when the deforming stress is removed. These properties are important for its function.
The surface of articular cartilage is smooth to the finger, like that of a billiard ball. Images obtained by a scanning electron microscope have shown, however, that the surface is actually irregular, more like that of a golf ball. The part of the cartilage nearest to the bone is impregnated with calcium salts. This calcified layer appears to be a barrier to the passage of oxygen and nutrients to the cartilage from the bone, such that the cartilage is largely dependent upon the synovial fluid for its nourishment.
Every articular cartilage has two parts: a central articulating part and a marginal nonarticulating part. The marginal part is much smaller than the central and is covered by a synovial membrane. It will be described later in connection with that membrane.
The central part is either single, if only two bones are included in the joint, or divided into clearly distinct portions by sharp ridges, if more than two bones are included. Thus, the upper articular surface of the arm bone (humerus) is single, for only this bone and the shoulder blade (scapula) are included in the shoulder joint. The lower articular surface of the humerus is subdivided into two parts, one for articulation with the radius and one for articulation with the ulna, both being included in the elbow joint. There is a functional reason for the subdivision, or partition, of articular cartilage when it does occur.
Within a diarthrosis joint, bones articulate in pairs, each pair being distinguished by its own pair of conarticular surfaces. Conarticular surfaces constitute "mating pairs." Each mating pair consists of a "male" surface and a "female" surface; the reasoning for these terms is explained below. As previously stated, there is only one such pair of bones within the shoulder joint; hence, there is only one pair of conarticular surfaces. There are two such pairs within the elbow joint—the humeroradial and humeroulnar. The radius moves on one of the two subdivisions of the lower humeral articular cartilage; the ulna moves on the other subdivision. There are then two pairs of conarticular surfaces within the elbow joint, even though there are only three bones in it.
Articular surfaces are divisible into two primary classes: ovoid and sellar. An ovoid surface is either convex in all directions or concave in all directions; in this respect it is like one or other of the two sides of a piece of eggshell, hence the name (ovum, egg). A sellar surface is convex in one direction and concave in the direction at right angles to the first; in this respect it is like the whole or part of a horse saddle (sella, saddle). There are no flat articular surfaces, although slightly curved ovoid or sellar surfaces may be classified as flat. Following an engineering convention, an ovoid surface is called "male" if it is convex, "female" if it is concave. In any diarthrosis having ovoid conarticular surfaces, the male surface is always of larger area than the female. For this reason the larger of two sellar conarticular surfaces is called male and the smaller female. The larger the difference in size between conarticular surfaces, the greater the possible amount of motion at the joint.
In all positions of a diarthrosis, except one, the conarticular surfaces fit imperfectly. This incongruence may not be large and may be lessened by mutual deformation of the opposed parts of the surfaces, a consequence of the deformability of articular cartilage. The exceptional position is called the close-packed position; in it the whole of the articulating portion of the female surface is in complete contact with the apposed part of the male surface, and the joint functionally is no longer a diarthrosis but is instead called a synchondrosis. Every joint has its close-packed position brought about by the action of the main ligaments of the joint. A good example is that of the wrist when the hand is fully bent backward (dorsiflexed) on the forearm. In closed-packed positions two bones in series are converted temporarily into a functionally single, but longer, unit that is more likely to be injured by sudden torsional stresses. Thus, a sprained or even fractured wrist usually occurs when that joint, when close packed, is suddenly and violently bent.
No articular surface is of uniform curvature; neither is it a "surface of revolution" such as a cylinder is. That part of a male conarticular surface that comes into contact with the female in close pack is both wider and of lesser curvature than is the remainder. Inspection of two articulating bones is enough to establish their position of close pack, flexion, extension, or whatever it may be.
Intra-articular fibrocartilages: Intra-articular fibrocartilages are complete or incomplete plates of fibrocartilage that are attached to the joint capsule (the investing ligament) and that stretch across the joint cavity between a pair of conarticular surfaces. When complete they are called disks; when incomplete they are called menisci. Disks are found in the temporomandibular joint of the lower jaw, the sternoclavicular (breastbone and collarbone) joint, and the ulnocarpal (inner forearm bone and wrist) joint. A pair of menisci is found in each knee joint, one between each femoral condyle and its female tibial counterpart. A small meniscus is found in the upper part of the acromioclavicular joint at the top of the shoulder. These fibrocartilages are really parts of the fibrous layer of the diarthrosis in which they occur, and they effect a complete or partial division of the articular bursa into two parts, depending upon whether they are disks or menisci, respectively. When the division is complete, there are really two synovial joints—e.g., the sternodiskal and the discoclavicular.
A disk or meniscus is mostly fibrocartilage, the chondrification being slight and the fibrous element predominating, especially in the part nearest to the investing ligament. Both animal experiments and surgical experience have shown that a meniscus of the knee can regrow if removed. The function of these intra-articular plates is to assist the gliding movements of the bones at the joints that contain them.
The synovial layer: The inner layer of the articular joint capsule is called the synovial layer (stratum synoviale) because it is in contact with the synovial fluid. Unlike the fibrous layer, it is incomplete and does not extend over the articulating parts of the articular cartilages and the central parts of articular disks and menisci.
The layer, commonly called the synovial membrane, is itself divisible into two strata, the intima and the subintima. The intima is smooth and moist on its free (synovial) surface. It could be described as an elastic plastic in which cells are embedded. Its elasticity allows it to stretch when one of the articulating bones either spins or swings to the opposite side and to return to its original size when the movement of the bone is reversed.
The cells of a synovial membrane can be divided into two classes: synovial lining cells and protective cells. The synovial lining cells are responsible for the generation and maintenance of the matrix. Their form depends upon their location. They are flattened and rounded at or near the internal surface of the membrane, more elongated and spindle-shaped elsewhere. They appear to be quite mobile and able to make their way to the free surface of the membrane. Excepting the regions in which the synovial membrane passes from the investing ligament (fibrous capsule) to the synovial periostea, these cells are scattered and do not form a continuous surface layer as do, for example, the cells lining the inner surface of the gut or of a blood vessel. In this respect they resemble the cells of other connective tissues, such as bone and cartilage. Apart from the generation and maintenance of the matrix of the membrane, they also can ingest foreign material and thus have a phagocytic function. They seem to be the only cells capable of secreting hyaluronic acid, the characteristic component of synovial fluid.
The protective cells are scattered through the depths of the membrane. They are of two kinds: mast cells and phagocytes. The mast cells secrete heparin and play the same part in synovial membrane as they do elsewhere—for example, in the skin and the gums. The phagocytes ingest unwanted particles, even such large ones as those of injected India ink; they are, in short, scavengers here as elsewhere.
The subintima is the connective tissue base on which the intima lies; it may be fibrous, fatty, or areolar (loose). In it are found the blood vessels and nerves that have penetrated the fibrous layer. Both the blood vessels and the nerves form plexuses, to be described later. The areolar subintima forms folds (synovial fringes) or minute fingerlike projections (villi) that project into the synovial fluid. The villi become more abundant in middle and old age. The fatty parts of the subintima may be quite thin, but in all joints there are places where they project into the bursal cavity as fatty pads (plicae adiposae); these are wedge-shaped in section, like a meniscus, with the base of the wedge against the fibrous capsule. The fatty pads are large in the elbow, knee, and ankle joints.
The function of fatty pads depends upon the fact that fat is liquid in a living body and that, therefore, a mass of fat cells is easily deformable. When a joint is moved, the synovial fluid is thrown into motion because it is adhesive to the articular cartilages, the motion of the fluid being in the direction of motion of the moving part. The fatty pads project into those parts of the synovial space in which there would be a likelihood of an eddying (vortical) motion of the fluid if those parts were filled with fluid. In short, the pads contribute to the "internal streamlining" of the joint cavity. Their deformability enables them to do this effectively. Of equal importance is the fact that the fatty pads by their very presence keep the synovial fluid between the immediately neighbouring parts of the male and female surfaces sufficiently thin, with proper elasticity as well as viscosity, to lubricate the joint.
Fatty pads are well provided with elastic fibres that bring about recovery from the deformation caused by pressure across a moving joint and that prevent the pads from being squeezed between two conarticular surfaces at rest. Such squeezing can happen, however, as the result of an accident and is very painful because of the large number of pain nerve fibres in these pads.
The synovial fluid: The main features of synovial fluid are: (1) Chemically, it is a dialyzate (a material subjected to dialysis) of blood plasma—that is, the portion of the plasma that has filtered through a membrane—but it contains a larger amount of hyaluronic acid than other plasma dialyzates. (2) Physically, it is a markedly thixotropic fluid—that is, one that is both viscous and elastic. Its viscosity decreases with an increase in the speed of the fluid when it is in motion. Its elasticity, on the other hand, increases with an increase in the speed of the fluid. Its thixotropy is due to the hyaluronic acid in it. (3) Functionally, it has two parts to play: nutrition and lubrication. It has been established that synovial fluid alone, by virtue of its being a blood-plasma dialyzate, can nourish the articulating parts of the articular cartilages. Its thixotropic properties make it suitable for forming what are called elastohydrodynamic lubricant films between the moving and the fixed conarticular surfaces of any mating pair. The motion of the synovial fluid, referred to earlier in connection with the fatty pads, assists its nutritional function by distributing it over the articular surfaces, from which it slowly passes into the interior of the cartilage. The source of the hyaluronic acid is the synovial lining cells.

Types of synovial joints
Recognition of the bursal nature of synovial joints makes it possible to describe them simply in terms of the bursal wall and to group together a number of types of structures. There are seven types of synovial joints: plane, hinge, pivot, sellar, ellipsoid, spheroidal (ball-and-socket), and bicondylar (two articulating surfaces). This classification is based on the anatomical form of the articular surfaces.
Plane joint: The plane, or arthrodial, joint has mating surfaces that are slightly curved and may be either ovoid or sellar. Only a small amount of gliding movement is found. Examples are the joints between the metacarpal bones of the hand and those between the cuneiform bones of the foot.
Hinge joint: The hinge, or ginglymus, joint is a modified sellar joint with each mating surface ovoid on its right and left sides. This modification reduces movement to a backward-forward swing like that allowed by the hinge of a box or a door. The swing of the joint, however, differs from that of a hinge in that it is accompanied by a slight spin (rotation) of the moving bone around its long axis. This brings the joint either into or out of its close-packed position, which is always that of extension. The joints between the bones of the fingers (phalanges) and that between the ulna (inner bone of the forearm) and the humerus at the elbow are classic examples.
Pivot joint: The pivot, or trochoid, joints are of two forms: in one a pivot rotates within a ring; in the other a ring moves around a pivot. In each case the ring is composed of fibrous tissue, part of which is converted into cartilage to form a female surface; the remainder may be ossified. Similarly, only part of the pivot is covered by a male articular cartilage. Pivot joints are always of the ovoid class; from a functional aspect, they are the ovoid counterparts of hinge joints. The joint between the atlas and the axis (first and second cervical vertebrae), directly under the skull, allows for turning of the head from side to side. Pivot joints also provide for the twisting movement of the bones of the forearm (radius and ulna) against the upper arm, a movement used, for instance, in unscrewing the lid of a jar.
Sellar joint: The sellar joint has already been described in the section Articular cartilage. It has two types of movement, both swings: flexion-extension and abduction-adduction. In addition to these it allows movements combining these two—that is, swings accompanied by rotation of the moving bone. An example of a sellar joint is the carpometacarpal joint of the thumb. The thumb can be swung from side to side or from behind forward, but the most frequent movement is that in which the thumb swings so that it comes "face to face" with one or another of the fingers, as in grasping a needle or a ball. This movement is called opposition (i.e., of thumb to fingers). During opposition the thumb is rotated around its long axis; it has been said that human civilization depends upon the opposition of the thumb.
Ellipsoid joint: The ellipsoid joint also has two types of movement but allows opposition movement only to a small degree. Its surfaces are ovoid and vary in both length and curvature as they are traced from front to back or from side to side, just as the diameter and curvature of an ellipse vary in directions at right angles to each other (hence the name). The joint between the second metacarpal and the first phalanx of the second finger is a good example. It allows the finger to flex and extend, to swing toward or away from its neighbouring finger, and to swing forward with a slight amount of rotation.
Ball and socket joint: The ball-and-socket joint, also known as a spheroidal joint, is the only one with three types of movement. It is an ovoid joint the male element of which could be described as a portion of a slightly deformed sphere. The rounded surface of the bone moves within a depression on another bone, thus allowing greater freedom of movement than any other kind of joint. It is most highly developed in the large hip and shoulder joints of mammals, including humans, in which it provides swing for the arms and legs in various directions and also spin of those limbs upon the more stationary bones.
Bicondylar joint: The condylar joint is better called bicondylar, for in it two distinct surfaces on one bone articulate with corresponding distinct surfaces on another bone. The two male surfaces are on one and the same bone and are of the same type (ovoid or sellar). These joints have two types of movement: one is always a swing, and the other is either another swing or a spin. Bicondylar joints are quite common. The largest is the tibiofemoral joint, in which both pairs of mating surfaces are within a single joint. At this joint, flexion and extension are the main movements; but active rotation of the leg on the femur is possible in most people when the leg and thigh are at right angles to each other. Every vertebra of the cervical, thoracic, and lumbar series is connected to (or separated from) the one below it by a pair of synovial joints as well as by an intervertebral disk. This pair of joints constitutes a bicondylar joint, the shape of whose articular surfaces determines the amount of movement permitted between the vertebra. The atlanto-occipital joint, between the skull and the vertebral column, is also a bicondylar joint. Finally, the right and left temporomandibular joints, between the lower jaw and the skull, are really two parts of a bicondylar joint, not only by definition—if the base of the skull is considered as a single bone—but also functionally, for one mandibular condyle cannot move without the other moving also.
JOINT LIGAMENTS
Any set of collagen fibres joining one bone of an articulating pair to the other is called a ligament. Thus, the articular bursal wall is a ligament, called either the fibrous capsule or the joint capsule.
There are two types of these sets: capsular and noncapsular. Capsular ligaments are simply thickenings of the fibrous capsule itself that take the form of either elongated bands or triangles, the fibres of which radiate from a small area of one articulating bone to a line upon its mating fellow. The iliofemoral ligament of the hip joint is an example of a triangular ligament. Capsular ligaments are found on the outer surface of the capsule. There is one exception to this rule: ligaments of the shoulder joint (glenohumeral ligaments) are found on the inner surface.
Noncapsular ligaments are free from the capsule and are of two kinds: internal and external. The internal type is found in the knee, wrist, and foot. In the knee there are two, both arising from the upper surface of the tibia; each passes to one of the two femoral condyles and lies within the joint cavity, surrounded by synovial membrane. They are called cruciate ligaments because they cross each other X-wise. At the wrist most of the articulations of the carpal bones share a common joint cavity, and neighbouring bones are connected sideways by short internal ligaments. The same is true of the tarsal bones that lie in front of the talus and the calcaneus.

The external noncapsular ligaments are of two kinds: proximate and remote. The proximate ligaments pass over at least two joints and are near the capsules of these joints. They are found only on the outer side of the lower limb. Examples are the outer (fibular) ligament of the knee, which passes from the femur to the upper part of the fibula over both the knee and tibiofibular joints, and the middle part of the outer ligament of the ankle joint, which passes from the lowest part of the fibula to the heel bone. These two ligaments, particularly that passing over the ankle, are especially liable to damage (sprain).
The remote ligaments are so called because they are far from, rather than close to, the joint capsule. A notable example is that of the ligaments that pass between the back parts (spines and laminae) of neighbouring vertebrae in the cervical, thoracic, and lumbar parts of the spinal column. These are the chief ligaments of the pairs of synovial joints between the vertebrae of these regions. Unlike most ligaments, they contain a high proportion of elastic fibres that assist the spinal column to return to its normal shape after it has been bent forward or sideways.
Contrary to the opinion of earlier anatomists, ligaments are not normally responsible for holding joint surfaces together. This is because a set of collagen fibres, like a string, can exert a reactive force only if stretched and tightened by some tensile stress. Normally, the bones at a joint are pressed together (when at rest) by the action of muscles or by gravity. An individual ligament can stop a movement that tightens it. Such a movement will loosen the ligaments that would be tightened by the opposite movement. The one exception to this case is the movement that brings a joint into the close-packed position. This movement is brought about by a combination of a swing with a spin of the moving bone. Experiments show that the combination of movement screws the articular surfaces firmly together so that they cannot be separated by traction and that the capsule and most of the ligaments are in simultaneous maximal tautness.
NERVE SUPPLY AND BLOOD SUPPLY OF JOINT
The nerve and blood supply of synovial joints follows the general rule for the body: "Ubi nervus ibi arteria" ("Where there is a nerve, there also is an artery").
Articular nerves
The sources of nerve fibres to a joint conform well to Hilton's law—the nerves to the muscles acting on a joint give branches to that joint as well as to the skin over the area of action of these muscles. Thus, the knee joint is supplied by branches from the femoral, sciatic, and obturator nerves, which among them supply the various muscles moving the joint. Some of these nerves go to the fibrous capsule and ligaments; others innervate this capsule and reach the synovial membrane. Some of these nerves are sensory; others give both motor and sensory fibres to the arteries that accompany them.
The sensory fibres to the fibrous capsule are of two kinds: (1) algesic, responsible for painful sensation, particularly when the capsule or other ligaments are overstretched or torn, and (2) proprioceptive, which terminate in various forms of specialized structures and convey information to all parts of the central nervous system, including the cerebellum and the cerebrum. It has been established that this information includes the posture of a resting joint and both the rate and extent of motion at a moving joint. The latter is supplemented by impulses conveyed by the nerves from the muscles acting and the skin affected by the movement.

The sensory fibres to the synovial membrane reach it by innervating the fibrous capsule at various points and form wide-meshed networks in the subsynovial layer. They are mainly algesic in function, and stimulation of them gives rise to diffused rather than localized pain (unlike the corresponding fibres to the fibrous capsule). They are found wherever the synovial membrane is, being especially abundant in the fatty pads, and are also present over the peripheral (nonarticulating) parts of the articular cartilage, disks, and menisci. This fact accounts for the excruciating pain that accompanies injury of these latter structures. The articulating part of the articular cartilage has no nerve supply.
Articular blood and lymph vessels
The joints are surrounded by a rich network of arteries and veins. The arteries in the vicinity of a synovial joint give off subdivisions that join (anastomose) freely on its outer surface. From the network of vessels so formed, branches lead to the fibrous capsule and ligaments and to the synovial membrane. Blood vessels to the synovial membrane are accompanied by nerves, and, when these vessels reach the subsynovial membrane, they proliferate to form another anastomotic network from which capillaries go to all parts of the membrane. These subsynovial arteries also ramify to the fatty pads and the nonarticulating parts of the articular cartilage, disks, and menisci. None, however, go to the articulating part of an articular cartilage, which therefore depends upon the synovial fluid for its nourishment.
Veins align with the arteries. In addition, a joint has a well-developed set of lymphatic vessels, the ultimate channels of which join those of the neighbouring parts of the limb or body wall.
NUTRITION AND METABOLISM OF JOINT
The metabolism and nutrition of the fibrous capsule and ligaments are similar to that of fibrous tissues elsewhere. Their blood supply is relatively small, indicating a low rate of metabolic changes. Unlike skin, for example, they heal slowly if injured.
The metabolism of articular cartilage is primarily dependent upon that of its cells (chondrocytes). Carbohydrate metabolism in these cells is similar to that of cells elsewhere and is unaffected by age. The oxygen consumption of the chondrocytes, on the other hand, decreases with age once the cells have matured. All the evidence suggests that the intracellular combustion is of glucose and protein, in that order of preference, rather than of fat. Sulfur passes from the blood to the synovial fluid and from there to the chondrocytes. From these it is transferred to the matrix to help to form chondroitin sulfate and keratosulfate molecules, the main constituents of the cartilaginous material. Chondroitin sulfate could be described as a sulfonated form of hyaluronic acid, the characteristic constituent of synovial fluid. Its presence in the matrix of the cartilage, but not in the synovial fluid, shows that the chondrocytes are necessary for its formation. After the second decade of life, the proportion of chondroitin sulfate falls and that of keratosulfate rises, as would be expected in view of the corresponding diminution of metabolic activity of the cells.
Excepting the articular cartilages, disks, and menisci, all other tissues of synovial joints are nourished directly by the blood vessels. The excepted parts are nourished indirectly by the synovial fluid. This is distributed over the surface of the articulating cartilage by the movements of the joint. The need for keeping joints healthy by frequent exercise of all of them is thus apparent.

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