Bone
| Bone | |
|---|---|
 | |
 A scanning electronic micrograph of a Wistar rat's bone at 10,000× magnification | |
| Details | |
| Identifiers | |
| Latin | os, ossis |
| Greek | ὀστέον (ostéon) |
| MeSH | D001842 |
| TA98 | A02.0.00.000 |
| TA2 | 366, 377 |
| TH | H3.01.00.0.00001 |
| FMA | 5018 |
| Anatomical terminology | |
A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals.[1] Bones protect the organs of the body, produce red and white blood cells, store minerals, help regulate acid-base homeostasis, provide structure and support for the body, and enable mobility and hearing. Bones come in a variety of shapes and sizes and have complex internal and external structures.[2]
Bone tissue (also known as osseous tissue or bone in the uncountable) is a form of hard tissue, specialised connective tissue that is mineralized and has an intercellular honeycomb-like matrix,[3] which helps to give the bone rigidity. Bone tissue is made up of different types of bone cells: osteoblasts and osteocytes (which form and mineralise bone respectively), osteoclasts (which resorb bone), and modified or flattened osteoblasts (lining cells that form a protective layer on the bone surface). The mineralised matrix of bone tissue has an organic component of mainly ossein, a form of collagen, and an inorganic component of bone mineral, made up of various salts. Bone tissue comprises cortical (compact) bone and cancellous (spongy) bone, although bones may also contain other kinds of tissue including bone marrow, endosteum, periosteum, nerves, blood vessels, and cartilage.
In the human body at birth, approximately 300 bones are present. Many of these fuse together during development, leaving a total of 206 separate bones in the adult, not counting numerous small sesamoid bones.[4][5] The largest bone in the body is the femur or thigh-bone, and the smallest is the stapes in the middle ear.
The Ancient Greek word for bone is ὀστέον ("osteon"). In anatomical terminology, including in the Terminologia Anatomica, the word for a bone is os (for example, os breve, os longum, os sesamoideum). (This is not to be confused with the alternative medical use of os to mean orifice, from the Latin ōs, mouth.)
Gross anatomy
[edit]Five types of bones are found in the human body: long, short, flat, irregular, and sesamoid.[6]
- Long bones are characterized by a shaft, the diaphysis, that is much longer than its width; and by an epiphysis, a rounded head at each end of the shaft. They are made up mostly of compact bone, with lesser amounts of marrow, located within the medullary cavity, and areas of spongy, cancellous bone at the ends of the bones.[7]
- Most bones of the limbs, including those of the fingers and toes, are long bones. The exceptions are the eight carpal bones of the wrist, the seven articulating tarsal bones of the ankle and the sesamoid bone of the kneecap. Long bones such as the clavicle, that have a differently shaped shaft or ends are also called modified long bones.
- Short bones are roughly cube-shaped, and have only a thin layer of compact bone surrounding a spongy interior. Short bones provide stability and support as well as some limited motion.[8]
- The bones of the wrist and ankle are short bones.
- Flat bones are thin and generally curved, with two parallel layers of compact bone sandwiching a layer of spongy bone.
- Sesamoid bones are bones embedded in tendons. Since they act to hold the tendon further away from the joint, the angle of the tendon is increased and thus the leverage of the muscle is increased.
- Irregular bones do not fit into the above categories. They consist of thin layers of compact bone surrounding a spongy interior. As implied by the name, their shapes are irregular and complicated. Often this irregular shape is due to their many centers of ossification or because they contain bony sinuses.
Terminology
[edit]
Anatomists use a number of anatomical terms to describe the appearance, shape and function of bones. Like other anatomical terms, many of these derive from Latin and Greek. Some anatomists still use Latin to refer to bones. The term "osseous", and the prefix "osteo-", referring to things related to bone, are still used commonly today.
Some examples of terms used to describe bones include the term "foramen" to describe a hole through which something passes, and a "canal" or "meatus" to describe a tunnelure. A protrusion from a bone can be called a number of terms, including a "condyle", "crest", "spine", "eminence", "tubercle" or "tuberosity", depending on the protrusion's shape and location. In general, long bones are said to have a "head", "neck", and "body".
When two bones join, they are said to "articulate". If the two bones have a fibrous connection and are relatively immobile, then the joint is called a "suture".
Functions
[edit]Mechanical
[edit]Bones serve a variety of mechanical functions. Together the bones in the body form the skeleton. They provide a frame to keep the body supported, and an attachment point for skeletal muscles, tendons, ligaments and joints, which function together to generate and transfer forces so that individual body parts or the whole body can be manipulated in three-dimensional space (the interaction between bone and muscle is studied in biomechanics).
Bones protect internal organs, such as the skull protecting the brain or the ribs protecting the heart and lungs. Because of the way that bone is formed, bone has a high compressive strength of about 170 MPa (1,700 kgf/cm2),[12] poor tensile strength of 104–121 MPa, and a very low shear stress strength (51.6 MPa).[13][14] This means that bone resists pushing (compressional) stress well, resist pulling (tensional) stress less well, but only poorly resists shear stress (such as due to torsional loads). While bone is essentially brittle, bone does have a significant degree of elasticity, contributed chiefly by collagen.
Mechanically, bones also have a special role in hearing. The ossicles are three small bones in the middle ear which are involved in sound transduction.
Synthetic
[edit]The cancellous part of bones contain bone marrow. Bone marrow produces blood cells in a process called hematopoiesis.[15] Blood cells that are created in bone marrow include red blood cells, platelets and white blood cells.[16] Progenitor cells such as the hematopoietic stem cell divide in a process called mitosis to produce precursor cells. These include precursors which eventually give rise to white blood cells, and erythroblasts which give rise to red blood cells.[17] Unlike red and white blood cells, created by mitosis, platelets are shed from very large cells called megakaryocytes.[18] This process of progressive differentiation occurs within the bone marrow. After the cells are matured, they enter the circulation.[19] Every day, over 2.5 billion red blood cells and platelets, and 50–100 billion granulocytes are produced in this way.[20]
As well as creating cells, bone marrow is also one of the major sites where defective or aged red blood cells are destroyed.[20]
Metabolic
[edit]- Mineral storage – bones act as reserves of minerals important for the body, most notably calcium and phosphorus.[21][22][23]
Determined by the species, age, and the type of bone, bone cells make up to 15 percent of the bone. Growth factor storage—mineralized bone matrix stores important growth factors such as insulin-like growth factors, transforming growth factor, bone morphogenetic proteins and others.[24]
- Fat storage – marrow adipose tissue (MAT) acts as a storage reserve of fatty acids.[25]
- Acid-base balance – bone buffers the blood against excessive pH changes by absorbing or releasing alkaline salts.[26]
- Detoxification – bone tissues can also store heavy metals and other foreign elements, removing them from the blood and reducing their effects on other tissues. These can later be gradually released for excretion.[27]
- Endocrine organ – bone controls phosphate metabolism by releasing fibroblast growth factor 23 (FGF-23), which acts on kidneys to reduce phosphate reabsorption. Bone cells also release a hormone called osteocalcin, which contributes to the regulation of blood sugar (glucose) and fat deposition. Osteocalcin increases both the insulin secretion and sensitivity, in addition to boosting the number of insulin-producing cells and reducing stores of fat.[28]
- Calcium balance – the process of bone resorption by the osteoclasts releases stored calcium into the systemic circulation and is an important process in regulating calcium balance. As bone formation actively fixes circulating calcium in its mineral form, removing it from the bloodstream, resorption actively unfixes it thereby increasing circulating calcium levels. These processes occur in tandem at site-specific locations.[29]
Tissue
[edit]Bone is not uniformly solid, but consists of a flexible matrix (about 30%) and bound minerals (about 70%), which are intricately woven and continuously remodeled by a group of specialized bone cells. Their unique composition and design allows bones to be relatively hard and strong, while remaining lightweight. Bone matrix is 90 to 95% composed of elastic collagen fibers, also known as ossein,[30] and the remainder is ground substance.[31] The elasticity of collagen improves fracture resistance.[12] The matrix is hardened by the binding of inorganic mineral salt, calcium phosphate, in a chemical arrangement known as bone mineral, a form of calcium apatite.[32][33] It is the mineralisation that gives bones rigidity.
Within any single bone, the tissue is woven into two main patterns: cortical and cancellous bone, each with distinct appearances and characteristics. Bone is actively constructed and remodeled throughout life by specialized bone cells known as osteoblasts and osteoclasts.
Cortex
[edit]
The hard outer layer of bones is composed of cortical bone, which is also called compact bone as it is much denser than cancellous bone. It forms the hard exterior (cortex) of bones. The cortical bone gives bone its smooth, white, and solid appearance, and accounts for 80% of the total bone mass of an adult human skeleton.[34] It facilitates bone's main functions—to support the whole body, to protect organs, to provide levers for movement, and to store and release chemical elements, mainly calcium. It consists of multiple microscopic columns, each called an osteon or Haversian system. Each column is multiple layers of osteoblasts and osteocytes around a central canal called the osteonic canal. Volkmann's canals at right angles connect the osteons together. The columns are metabolically active, and as bone is reabsorbed and created the nature and location of the cells within the osteon will change. Cortical bone is covered by a periosteum on its outer surface, and an endosteum on its inner surface. The endosteum is the boundary between the cortical bone and the cancellous bone.[35] The primary anatomical and functional unit of cortical bone is the osteon.
Trabeculae
[edit]
Cancellous bone, spongy bone,[36][35] or trabecular bone is the internal tissue of the skeletal bone and is an open-cell porous network that follows the material properties of biofoams.[37][38] Cancellous bone has a higher surface-area-to-volume ratio than cortical bone and it is less dense. This makes it weaker and more flexible. The greater surface area also makes it suitable for metabolic activities such as the exchange of calcium ions. Cancellous bone is typically found at the ends of long bones, near joints, and in the interior of vertebrae. Cancellous bone is highly vascular and often contains red bone marrow where hematopoiesis, the production of blood cells, occurs. The primary anatomical and functional unit of cancellous bone is the trabecula. The trabeculae are aligned towards the mechanical load distribution that a bone experiences within long bones such as the femur. As far as short bones are concerned, trabecular alignment has been studied in the vertebral pedicle.[39] Thin formations of osteoblasts covered in endosteum create an irregular network of spaces,[40] known as trabeculae. Within these spaces are bone marrow and hematopoietic stem cells that give rise to platelets, red blood cells and white blood cells.[40] Trabecular marrow is composed of a network of rod- and plate-like elements that make the overall organ lighter and allow room for blood vessels and marrow. Trabecular bone accounts for the remaining 20% of total bone mass but has nearly ten times the surface area of compact bone.[41]
The words cancellous and trabecular refer to the tiny lattice-shaped units (trabeculae) that form the tissue. It was first illustrated accurately in the engravings of Crisóstomo Martinez.[42]
Marrow
[edit]Bone marrow, also known as myeloid tissue in red bone marrow, can be found in almost any bone that holds cancellous tissue. In newborns, all such bones are filled exclusively with red marrow or hematopoietic marrow, but as the child ages the hematopoietic fraction decreases in quantity and the fatty/ yellow fraction called marrow adipose tissue (MAT) increases in quantity. In adults, red marrow is mostly found in the bone marrow of the femur, the ribs, the vertebrae and pelvic bones.[43]
Vascular supply
[edit]Bone receives about 10% of cardiac output.[44] Blood enters the endosteum, flows through the marrow, and exits through small vessels in the cortex.[44] In humans, blood oxygen tension in bone marrow is about 6.6%, compared to about 12% in arterial blood, and 5% in venous and capillary blood.[44]
Histology and physiology
[edit]
Bone is metabolically active tissue composed of several types of cells. These cells include osteoblasts, which are involved in the creation and mineralization of bone tissue, osteocytes, and osteoclasts, which are involved in the reabsorption of bone tissue. Osteoblasts and osteocytes are derived from osteoprogenitor cells, but osteoclasts are derived from the same cells that differentiate to form macrophages and monocytes.[45] Within the marrow of the bone there are also hematopoietic stem cells. These cells give rise to other cells, including white blood cells, red blood cells, and platelets.[20]
Osteoblast
[edit]
Osteoblasts are mononucleate bone-forming cells. They are located on the surface of osteon seams and make a protein mixture known as osteoid, which mineralizes to become bone.[46] The osteoid seam is a narrow region of a newly formed organic matrix, not yet mineralized, located on the surface of a bone. Osteoid is primarily composed of Type I collagen. Osteoblasts also manufacture hormones, such as prostaglandins, to act on the bone itself. The osteoblast creates and repairs new bone by actually building around itself. First, the osteoblast puts up collagen fibers. These collagen fibers are used as a framework for the osteoblasts' work. The osteoblast then deposits calcium phosphate which is hardened by hydroxide and bicarbonate ions. The brand-new bone created by the osteoblast is called osteoid.[47] Once the osteoblast is finished working it is actually trapped inside the bone once it hardens. When the osteoblast becomes trapped, it becomes known as an osteocyte. Other osteoblasts remain on the top of the new bone and are used to protect the underlying bone, these become known as bone lining cells.[48]
Osteocyte
[edit]Osteocytes are cells of mesenchymal origin and originate from osteoblasts that have migrated into and become trapped and surrounded by a bone matrix that they themselves produced.[35] The spaces the cell body of osteocytes occupy within the mineralized collagen type I matrix are known as lacunae, while the osteocyte cell processes occupy channels called canaliculi. The many processes of osteocytes reach out to meet osteoblasts, osteoclasts, bone lining cells, and other osteocytes probably for the purposes of communication.[49] Osteocytes remain in contact with other osteocytes in the bone through gap junctions—coupled cell processes which pass through the canalicular channels.
Osteoclast
[edit]Osteoclasts are very large multinucleate cells that are responsible for the breakdown of bones by the process of bone resorption. New bone is then formed by the osteoblasts. Bone is constantly remodeled by the resorption of osteoclasts and created by osteoblasts.[45] Osteoclasts are large cells with multiple nuclei located on bone surfaces in what are called Howship's lacunae (or resorption pits). These lacunae are the result of surrounding bone tissue that has been reabsorbed.[50] Because the osteoclasts are derived from a monocyte stem-cell lineage, they are equipped with phagocytic-like mechanisms similar to circulating macrophages.