Vertebra

A diagram of a human thoracic vertebra. Notice the articulations for the ribs

A vertebra (plural: vertebrae) is an individual bone in the flexible column that defines vertebrate animals, e.g., humans. The vertebral column encases and protects the spinal cord, which runs from the base of the cranium down the dorsal side of the animal until reaching the pelvis. From there, vertebrae continue into the tail.

Vertebrae are defined by the regions of the vertebral column they occur in. Cervical vertebrae are those in the neck area. With exception of two sloth species (Choleopus and Bradypus) and the manatee (Trichechus), all mammals have seven cervical vertebrae^[1]. In other vertebrates it can range from a single vertebra in amphibians, to as many as 25 in swans or 76 in the extinct plesiosaur Elasmosaurus. The dorsal vertebrae range from the bottom of the neck to the top of the pelvis. Dorsal vertebrae attached to ribs are called thoracic vertebrae, while those without ribs are called lumbar vertebrae. The sacral vertebrae are those in the pelvic region, and range from one in amphibians, to two in most birds and modern reptiles, or up to 3 to 5 in mammals. When multiple sacral vertebrae are fused into a single structure, it is called the sacrum. The synsacrum is a similar fused structure found in birds that is composed of the sacral, lumbar, and some of the thoracic and caudal vertebra, as well as the pelvic girdle. Caudal vertebrae compose the tail, and the final few can be fused into the pygostyle in birds, or into the coccygeal or tail bone in chimpanzees and humans.

Structure

Individual vertebrae are composed of a centrum (body), arches protruding from the top and bottom of the centrum, and various processes projecting from the centrum and/or arches. An arch extending from the top of the centrum is called a neural arch, while the hemal arch or chevron is found underneath the centrum in the caudal (tail) vertebrae of fish, most reptiles, some birds, and some mammals with long tails. The vertebral processes can either give the structure rigidity, help them articulate with ribs, or serve as muscle attachment points. Common types are tranverse process, diapophyses, parapophyses, and zygapophyses (both the cranial zygapophyses and the caudal zygapophyses).

Amphicelous refers to a centrum that is concave at both ends, similar to those found in most fish. Opisthocoelous centra are convex in the front and concave in the back, similar to those of most salamanders. In contrast, procelous centra are concave in the front and convex in the back, as found in most frogs and modern reptiles. Centra with flat ends are acelous, like those in mammals. Birds have heterocelous centra, shaped like saddles at both ends.

In humans

There are normally thirty-three (33) vertebrae in humans, including the five that are fused to form the sacrum (the others are separated by intervertebral discs) and the four coccygeal bones that form the tailbone. The upper three regions comprise the remaining 24, and are grouped under the names cervical (7 vertebrae), thoracic (12 vertebrae) and lumbar (5 vertebrae), according to the regions they occupy. This number is sometimes increased by an additional vertebra in one region, or it may be diminished in one region, the deficiency often being supplied by an additional vertebra in another. The number of cervical vertebrae is, however, very rarely increased or diminished.

With the exception of the first and second cervical, the true or movable vertebrae (the upper three regions) present certain common characteristics that are best studied by examining one from the middle of the thoracic region.

General structure

Oblique view of cervical vertebrae

A typical vertebra consists of two essential parts: an anterior (front) segment, which is the vertebral body; and a posterior part – the vertebral (neural) arch – which encloses the vertebral foramen. The vertebral arch is formed by a pair of pedicles and a pair of laminae, and supports seven processes, four articular, two transverse, and one spinous, the latter also being known as the neural spine.

When the vertebrae are articulated with each other, the bodies form a strong pillar for the support of the head and trunk, and the vertebral foramina constitute a canal for the protection of the medulla spinalis (spinal cord). In between every pair of vertebrae are two apertures, the intervertebral foramina, one on either side, for the transmission of the spinal nerves and vessels.

Two transverse processes and one spinous process are posterior to (behind) the vertebral body. The spinous process comes out the back, one transverse process comes out the left, and one on the right. The spinous processes of the cervical and lumbar regions can be felt through the skin. Superior and inferior articular facets on each vertebra act to restrict the range of movement possible. These facets are joined by a thin portion of the neural arch called the pars interarticularis.

Classification

The centra of the vertebra can be classified based upon the fusion of its elements. In aspidospondyly, bones such as the neural spine, the pleurocentrum and the intercentrum are separate ossifications. Fused elements, however, classify a vertebra as having holospondyly.

A vertebra can also be described in terms of the shape of the ends of the centra. Humans are said to be acoelous, or with flat ends. These flat ends of the centra are especially good at supporting and distributing compressive forces. Amphicoelus vertebra is represented by both ends of the centra being concave. This shape is common in fish, where most motion is limited. Amphicoelus centra often are integrated with a full notochord. Procoelus vertebrae are anteriorly concave, and posteriorly convex. An opisthocoelus vertebra however, possess anterior convexity, and posterior concavity. Heterocoelous vertebrae are saddle-shaped at each end of the centra. This type of configuration is seen in turtles that retract their necks, and birds, because it permits extensive lateral and vertical flexion motion without stretching the nerve cord too extensively or wringing it about its long axis.

Regions

Orientation of vertebral column on surface. T3 is at level of medial part of spine of scapula. T7 is at inferior angle of the scapula. L4 is at highest point of iliac crest. S2 is at the level of posterior superior iliac spine. Furthermore, C7 is easily localized as a prominence at the lower part of the neck.^[2]

Cervical

There are seven (7) cervical bones (but 8 cervical spinal nerves) and these bones are, in general, small and delicate. Their spinous processes are short (with the exception of C2 and C7, which have palpable spinous processes). Numbered top-to-bottom from C1-C7, atlas (C1) and axis (C2), are the vertebrae that allow the neck and head so much movement. For the most part, the atlanto-occipital joint allows the skull to move up and down, while the atlanto-axial joint allows the upper neck to twist left and right. The axis also sits upon the first intervertebral disk of the spinal column. All mammals except manatees and sloths have seven cervical vertebrae, whatever the length of the neck.

Cervical vertebrae possess transverse foramina to allow for the vertebral arteries to pass through on their way to the foramen magnum to end in the circle of Willis. These are the smallest, lightest vertebrae and the vertebral foramina are triangular in shape. The spinous processes are short and often bifurcated (the spinous process of C7, however, is not bifurcated, and is substantially longer than that of the other cervical spinous processes).

Thoracic

The twelve (12) thoracic bones and their transverse processes have surfaces that articulate with the ribs. Some rotation can occur between the thoracic vertebrae, but their connection with the rib cage prevents much flexion or other excursion. They may also be known as 'dorsal vertebrae', in the human context.

Bodies are roughly heart-shaped and are about as wide anterio-posterioly as they are in the transverse dimension. Vertebral foramina are roughly circular in shape.

Lumbar

These five (5) vertebrae are very robust in construction, as they must support more weight than other vertebrae. They allow significant flexion and extension, moderate lateral flexion (sidebending), and a small degree of rotation. The discs between these vertebrae create a lumbar lordosis (curvature that is concave posteriorly) in the human spine.

Sacral

There are five (5) vertebrae (S1-S5) and they are fused in maturity, with no intervertebral discs.

Coccygeal

There are usually four (4) and rarely 3-5 vertebrae (Co1-Co5), with no intervertebral discs. Many animals have a greater number of "tail vertebrae," and, in animals, they are more commonly known as "caudal vertebrae." Pain at the coccyx (tailbone) is known as coccydynia.

Development

The striking segmented pattern of the human spine is established during embryogenesis when the precursor of the vertebrae, the somites, are rhythmically added to the forming posterior part of the embryo. In humans, somite formation begins around the third week post-fertilization and continues until a total of around 52 somites are formed. The somites are epithelial spheres that contain the precursors of the vertebrae, the ribs, the skeletal muscles of the body wall and limbs, and the dermis of the back. The periodicity of somite distribution and production is thought to be imposed by a molecular oscillator or clock acting in cells of the presomitic mesoderm (PSM). Somites form soon after the beginning of gastrulation, on both sides of the neural tube from a tissue called the presomitic mesoderm (PSM). The PSM is part of the paraxial mesoderm and is generated caudally by gastrulation when cells ingress through the primitive streak, and later, through the tail bud. Soon after their formation, somites become subdivided into the dermomyotome dorsally, which gives rise to the muscles and dermis, and the sclerotome ventrally, which will form the spine components. Sclerotomes become subvidided into an anterior and a posterior compartment. This subdivision plays a key role in the definitive patterning of vertebrae that form when the posterior part of one somite fuses to the anterior part of the consecutive somite during a process termed resegmentation. Disruption of the somitogenesis process in humans results in diseases such as congenital scoliosis. So far, the human homologues of three genes associated to the mouse segmentation clock (MESP2, DLL3 and LFNG) have been shown to be mutated in human patients with human congenital scoliosis suggesting that the mechanisms involved in vertebral segmentation are conserved across vertebrates. In humans the first four somites are incorporated in the basi-occipital bone of the skull and the next 33 somites will form the vertebrae. The remaining posterior somites degenerate. During the fourth week of embryonic development, the sclerotomes shift their position to surround the spinal cord and the notochord. The sclerotome is made of mesoderm and originates from the ventromedial part of the somites. This column of tissue has a segmented appearance, with alternating areas of dense and less dense areas.

As the sclerotome develops, it condenses further eventually developing into the vertebral body. Development of the appropriate shapes of the vertebral bodies is regulated by HOX genes.

The less dense tissue that separates the sclerotome segments develop into the intervertebral discs.

The notochord disappears in the sclerotome (vertebral body) segments, but persists in the region of the intervertebral discs as the nucleus pulposus. The nucleus pulposus and the fibers of the annulus fibrosus make up the intervertebral disc.

The primary curves (thoracic and sacral curvatures) form during fetal development. The secondary curves develop after birth. The cervical curvature forms as a result of lifting the head and the lumbar curvature forms as a result of walking.

Unfused arch of C1 at CT.

There are various defects associated with vertebral development. Scoliosis will result in improper fusion of the vertebrae. In Klippel-Feil anomaly patients have two or more cervical vertebrae that are fused together, along with other associated birth defects. One of the most serious defects is failure of the vertebral arches to fuse. This results in a condition called spina bifida. There are several variations of spina bifida that reflect the severity of the defect.

In other animals

Fish and amphibians

The vertebrae of lobe-finned fishes consist of three discrete bony elements. The vertebral arch surrounds the spinal cord, and is of broadly similar form to that found in most other vertebrates. Just beneath the arch lies a small plate-like pleurocentrum, which protects the upper surface of the notochord, and below that, a larger arch-shaped intercentrum to protect the lower border. Both of these structures are embedded within a single cylindrical mass of cartilage. A similar arrangement was found in the primitive Labyrinthodonts, but in the evolutionary line that led to reptiles (and hence, also to mammals and birds), the intercentrum became partially or wholly replaced by an enlarged pleurocentrum, which in turn became the bony vertebral body.^[3]

In most ray-finned fishes, including all teleosts, these two structures are fused with, and embedded within, a solid piece of bone superficially resembling the vertebral body of mammals. In living amphibians, there is simply a cylindrical piece of bone below the vertebral arch, with no trace of the separate elements present in the early tetrapods.^[3]

In cartilagenous fish, such as sharks, the vertebrae consist of two cartilagenous tubes. The upper tube is formed from the vertebral arches, but also includes additional cartilagenous structures filling in the gaps between the vertebrae, and so enclosing the spinal cord in an essentially continuous sheath. The lower tube surrounds the notochord, and has a complex structure, often including multiple layers of calcification.^[3]

Lampreys have vertebral arches, but nothing resembling the vertebral bodies found in all higher vertebrates. Even the arches are discontinuous, consisting of separate pieces of arch-shaped cartilage around the spinal cord in most parts of the body, changing to long strips of cartilage above and below in the tail region. Hagfishes lack a true vertebral column, and are therefore not properly considered vertebrates, but a few tiny neural arches are present in the tail.^[3]

Amniotes

The general structure of human vertebrae is fairly typical of that found in mammals, reptiles, and birds. The shape of the vertebral body does, however, vary somewhat between different groups. In mammals, such as humans, it typically has flat upper and lower surfaces, while in reptiles the anterior surface commonly has a concave socket into which the expanded convex face of the next vertebral body fits. Even these patterns are only generalisations, however, and there may be variation in form of the vertebrae along the length of the spine even within a single species. Some unusual variations include the saddle-shaped sockets between the cervical vertebrae of birds and the presence of a narrow hollow canal running down the centre of the vertebral bodies of geckos and tuataras, containing a remnant of the notochord.^[3]

Reptiles often retain the primitive intercentra, which are present as small crescent-shaped bony elements lying between the bodies of adjacent vertebrae; similar structures are often found in the caudal vertebrae of mammals. In the tail, these are attached to chevron-shaped bones called haemal arches, which attach below the base of the spine, and help to support the musculature. These latter bones are probably homologous with the ventral ribs of fish. The number of vertebrae in the spines of reptiles is highly variable, and may be several hundred in some species of snake.^[3]

In birds, there is a variable number of cervical vertebrae, which often form the only truly flexible part of the spine. The thoracic vertebrae are partially fused, providing a solid brace for the wings during flight. The sacral vertebrae are fused with the lumbar vertebrae, and some thoracic and caudal vertebrae, to form a single structure, the synsacrum, which is thus of greater relative length than the sacrum of mammals. In living birds, the remaining caudal vertebrae are fused into a further bone, the pygostyle, for attachment of the tail feathers.^[3]

Aside from the tail, the number of vertebrae in mammals is generally fairly constant. There are almost always seven cervical vertebrae (sloths and manatees are among the few exceptions), followed by around twenty or so further vertebrae, divided between the thoracic and lumbar forms, depending on the number of ribs. There are generally three to five vertebrae with the sacrum, and anything up to fifty caudal vertebrae.^[3]

See also

• Copenhagen disease

Footnotes

References

• Gray's Anatomy: The Vertebral column - The 1917 Gray's Anatomy is available via the Bartleby project. It is available with full colour diagrams, and provides an excellent starting point in anatomy, as well as a relatively complete source for gross anatomy. Some of this article was copied and pasted from the 1917 Gray's Anatomy, which is in the public domain.