Module 21: Bone Tissue and the Skeletal System

Lesson 4: Bone Formation and Development

Quá Trình Hình Thành Và Phát Triển Xương

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Mỗi bài học (lesson) bao gồm 4 phần chính: Thuật ngữ, Luyện Đọc, Luyện Nghe, và Bàn Luận.
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Medical Terminology: Bone Tissue and the Skeletal System

articular cartilage
thin layer of cartilage covering an epiphysis; reduces friction and acts as a shock absorber
where two bone surfaces meet
hard, dense connective tissue that forms the structural elements of the skeleton
(singular = canaliculus) channels within the bone matrix that house one of an osteocyte’s many cytoplasmic extensions that it uses to communicate and receive nutrients
semi-rigid connective tissue found on the skeleton in areas where flexibility and smooth surfaces support movement
central canal
longitudinal channel in the center of each osteon; contains blood vessels, nerves, and lymphatic vessels; also known as the Haversian canal
closed reduction
manual manipulation of a broken bone to set it into its natural position without surgery
compact bone
dense osseous tissue that can withstand compressive forces
tubular shaft that runs between the proximal and distal ends of a long bone
layer of spongy bone, that is sandwiched between two the layers of compact bone found in flat bones
endochondral ossification
process in which bone forms by replacing hyaline cartilage
delicate membranous lining of a bone’s medullary cavity
epiphyseal line
completely ossified remnant of the epiphyseal plate
epiphyseal plate
(also, growth plate) sheet of hyaline cartilage in the metaphysis of an immature bone; replaced by bone tissue as the organ grows in length
wide section at each end of a long bone; filled with spongy bone and red marrow
external callus
collar of hyaline cartilage and bone that forms around the outside of a fracture
flat bone
thin and curved bone; serves as a point of attachment for muscles and protects internal organs
broken bone
fracture hematoma
blood clot that forms at the site of a broken bone
production of blood cells, which occurs in the red marrow of the bones
opening or depression in a bone
condition characterized by abnormally high levels of calcium
condition characterized by abnormally low levels of calcium
internal callus
fibrocartilaginous matrix, in the endosteal region, between the two ends of a broken bone
intramembranous ossification
process by which bone forms directly from mesenchymal tissue
irregular bone
bone of complex shape; protects internal organs from compressive forces
(singular = lacuna) spaces in a bone that house an osteocyte
long bone
cylinder-shaped bone that is longer than it is wide; functions as a lever
medullary cavity
hollow region of the diaphysis; filled with yellow marrow
process, during bone growth, by which bone is resorbed on one surface of a bone and deposited on another
nutrient foramen
small opening in the middle of the external surface of the diaphysis, through which an artery enters the bone to provide nourishment
open reduction
surgical exposure of a bone to reset a fracture
doctor who specializes in diagnosing and treating musculoskeletal disorders and injuries
osseous tissue
bone tissue; a hard, dense connective tissue that forms the structural elements of the skeleton
(also, osteogenesis) bone formation
ossification center
cluster of osteoblasts found in the early stages of intramembranous ossification
cell responsible for forming new bone
cell responsible for resorbing bone
primary cell in mature bone; responsible for maintaining the matrix
osteogenic cell
undifferentiated cell with high mitotic activity; the only bone cells that divide; they differentiate and develop into osteoblasts
uncalcified bone matrix secreted by osteoblasts
(also, Haversian system) basic structural unit of compact bone; made of concentric layers of calcified matrix
disease characterized by a decrease in bone mass; occurs when the rate of bone resorption exceeds the rate of bone formation, a common occurrence as the body ages
perforating canal
(also, Volkmann’s canal) channel that branches off from the central canal and houses vessels and nerves that extend to the periosteum and endosteum
membrane that covers cartilage
fibrous membrane covering the outer surface of bone and continuous with ligaments
primary ossification center
region, deep in the periosteal collar, where bone development starts during endochondral ossification
bone markings where part of the surface sticks out above the rest of the surface, where tendons and ligaments attach
proliferative zone
region of the epiphyseal plate that makes new chondrocytes to replace those that die at the diaphyseal end of the plate and contributes to longitudinal growth of the epiphyseal plate
red marrow
connective tissue in the interior cavity of a bone where hematopoiesis takes place
process by which osteoclasts resorb old or damaged bone at the same time as and on the same surface where osteoblasts form new bone to replace that which is resorbed
reserve zone
region of the epiphyseal plate that anchors the plate to the osseous tissue of the epiphysis
secondary ossification center
region of bone development in the epiphyses
sesamoid bone
small, round bone embedded in a tendon; protects the tendon from compressive forces
short bone
cube-shaped bone that is approximately equal in length, width, and thickness; provides limited motion
skeletal system
organ system composed of bones and cartilage that provides for movement, support, and protection
spongy bone
(also, cancellous bone) trabeculated osseous tissue that supports shifts in weight distribution
(singular = trabecula) spikes or sections of the lattice-like matrix in spongy bone
yellow marrow
connective tissue in the interior cavity of a bone where fat is stored
zone of calcified matrix
region of the epiphyseal plate closest to the diaphyseal end; functions to connect the epiphyseal plate to the diaphysis
zone of maturation and hypertrophy
region of the epiphyseal plate where chondrocytes from the proliferative zone grow and mature and contribute to the longitudinal growth of the epiphyseal plate
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In the early stages of embryonic development, the embryo’s skeleton consists of fibrous membranes and hyaline cartilage. By the sixth or seventh week of embryonic life, the actual process of bone development, ossification (osteogenesis), begins. There are two osteogenic pathways—intramembranous ossification and endochondral ossification—but bone is the same regardless of the pathway that produces it.
Bone is a replacement tissue; that is, it uses a model tissue on which to lay down its mineral matrix. For skeletal development, the most common template is cartilage. During fetal development, a framework is laid down that determines where bones will form. This framework is a flexible, semi-solid matrix produced by chondroblasts and consists of hyaluronic acid, chondroitin sulfate, collagen fibers, and water. As the matrix surrounds and isolates chondroblasts, they are called chondrocytes. Unlike most connective tissues, cartilage is avascular, meaning that it has no blood vessels supplying nutrients and removing metabolic wastes. All of these functions are carried on by diffusion through the matrix. This is why damaged cartilage does not repair itself as readily as most tissues do.

Throughout fetal development and into childhood growth and development, bone forms on the cartilaginous matrix. By the time a fetus is born, most of the cartilage has been replaced with bone. Some additional cartilage will be replaced throughout childhood, and some cartilage remains in the adult skeleton.
During intramembranous ossification, compact and spongy bone develops directly from sheets of mesenchymal (undifferentiated) connective tissue. The flat bones of the face, most of the cranial bones, and the clavicles (collarbones) are formed via intramembranous ossification.

The process begins when mesenchymal cells in the embryonic skeleton gather together and begin to differentiate into specialized cells (Figure 1a). Some of these cells will differentiate into capillaries, while others will become osteogenic cells and then osteoblasts. Although they will ultimately be spread out by the formation of bone tissue, early osteoblasts appear in a cluster called an ossification center.

The osteoblasts secrete osteoid, uncalcified matrix, which calcifies (hardens) within a few days as mineral salts are deposited on it, thereby entrapping the osteoblasts within. Once entrapped, the osteoblasts become osteocytes (Figure 1b). As osteoblasts transform into osteocytes, osteogenic cells in the surrounding connective tissue differentiate into new osteoblasts.

Osteoid (unmineralized bone matrix) secreted around the capillaries results in a trabecular matrix, while osteoblasts on the surface of the spongy bone become the periosteum (Figure 1c). The periosteum then creates a protective layer of compact bone superficial to the trabecular bone. The trabecular bone crowds nearby blood vessels, which eventually condense into red marrow (Figure 1d).

Intramembranous ossification begins in utero during fetal development and continues on into adolescence. At birth, the skull and clavicles are not fully ossified nor are the sutures of the skull closed. This allows the skull and shoulders to deform during passage through the birth canal. The last bones to ossify via intramembranous ossification are the flat bones of the face, which reach their adult size at the end of the adolescent growth spurt.
In endochondral ossification, bone develops by replacing hyaline cartilage. Cartilage does not become bone. Instead, cartilage serves as a template to be completely replaced by new bone. Endochondral ossification takes much longer than intramembranous ossification. Bones at the base of the skull and long bones form via endochondral ossification.

In a long bone, for example, at about 6 to 8 weeks after conception, some of the mesenchymal cells differentiate into chondrocytes (cartilage cells) that form the cartilaginous skeletal precursor of the bones (Figure 2a). Soon after, the perichondrium, a membrane that covers the cartilage, appears (Figure 2b).

As more matrix is produced, the chondrocytes in the center of the cartilaginous model grow in size. As the matrix calcifies, nutrients can no longer reach the chondrocytes. This results in their death and the disintegration of the surrounding cartilage. Blood vessels invade the resulting spaces, not only enlarging the cavities but also carrying osteogenic cells with them, many of which will become osteoblasts. These enlarging spaces eventually combine to become the medullary cavity.

As the cartilage grows, capillaries penetrate it. This penetration initiates the transformation of the perichondrium into the bone-producing periosteum. Here, the osteoblasts form a periosteal collar of compact bone around the cartilage of the diaphysis. By the second or third month of fetal life, bone cell development and ossification ramps up and creates the primary ossification center, a region deep in the periosteal collar where ossification begins (Figure 2c).

While these deep changes are occurring, chondrocytes and cartilage continue to grow at the ends of the bone (the future epiphyses), which increases the bone’s length at the same time bone is replacing cartilage in the diaphyses. By the time the fetal skeleton is fully formed, cartilage only remains at the joint surface as articular cartilage and between the diaphysis and epiphysis as the epiphyseal plate, the latter of which is responsible for the longitudinal growth of bones. After birth, this same sequence of events (matrix mineralization, death of chondrocytes, invasion of blood vessels from the periosteum, and seeding with osteogenic cells that become osteoblasts) occurs in the epiphyseal regions, and each of these centers of activity is referred to as a secondary ossification center (Figure 2e).
The epiphyseal plate is the area of growth in a long bone. It is a layer of hyaline cartilage where ossification occurs in immature bones. On the epiphyseal side of the epiphyseal plate, cartilage is formed. On the diaphyseal side, cartilage is ossified, and the diaphysis grows in length. The epiphyseal plate is composed of four zones of cells and activity (Figure 3). The reserve zone is the region closest to the epiphyseal end of the plate and contains small chondrocytes within the matrix. These chondrocytes do not participate in bone growth but secure the epiphyseal plate to the osseous tissue of the epiphysis.

The proliferative zone is the next layer toward the diaphysis and contains stacks of slightly larger chondrocytes. It makes new chondrocytes (via mitosis) to replace those that die at the diaphyseal end of the plate. Chondrocytes in the next layer, the zone of maturation and hypertrophy, are older and larger than those in the proliferative zone. The more mature cells are situated closer to the diaphyseal end of the plate. The longitudinal growth of bone is a result of cellular division in the proliferative zone and the maturation of cells in the zone of maturation and hypertrophy.

Most of the chondrocytes in the zone of calcified matrix, the zone closest to the diaphysis, are dead because the matrix around them has calcified. Capillaries and osteoblasts from the diaphysis penetrate this zone, and the osteoblasts secrete bone tissue on the remaining calcified cartilage. Thus, the zone of calcified matrix connects the epiphyseal plate to the diaphysis. A bone grows in length when osseous tissue is added to the diaphysis.

Bones continue to grow in length until early adulthood. The rate of growth is controlled by hormones, which will be discussed later. When the chondrocytes in the epiphyseal plate cease their proliferation and bone replaces the cartilage, longitudinal growth stops. All that remains of the epiphyseal plate is the epiphyseal line (Figure 4).
While bones are increasing in length, they are also increasing in diameter; growth in diameter can continue even after longitudinal growth ceases. This is called appositional growth. Osteoclasts resorb old bone that lines the medullary cavity, while osteoblasts, via intramembranous ossification, produce new bone tissue beneath the periosteum. The erosion of old bone along the medullary cavity and the deposition of new bone beneath the periosteum not only increase the diameter of the diaphysis but also increase the diameter of the medullary cavity. This process is called modeling.
The process in which matrix is resorbed on one surface of a bone and deposited on another is known as bone modeling. Modeling primarily takes place during a bone’s growth. However, in adult life, bone undergoes remodeling, in which resorption of old or damaged bone takes place on the same surface where osteoblasts lay new bone to replace that which is resorbed. Injury, exercise, and other activities lead to remodeling. Those influences are discussed later in the chapter, but even without injury or exercise, about 5 to 10 percent of the skeleton is remodeled annually just by destroying old bone and renewing it with fresh bone.

OpenStax. (2022). Anatomy and Physiology 2e. Rice University. Retrieved June 15, 2023. ISBN-13: 978-1-711494-06-7 (Hardcover) ISBN-13: 978-1-711494-05-0 (Paperback) ISBN-13: 978-1-951693-42-8 (Digital). License: Attribution 4.0 International (CC BY 4.0). Access for free at

Intramembranous ossification follows four steps. (a) Mesenchymal cells group into clusters, and ossification centers form. (b) Secreted osteoid traps osteoblasts, which then become osteocytes. (c) Trabecular matrix and periosteum form. (d) Compact bone develops superficial to the trabecular bone, and crowded blood vessels condense into red marrow.

Endochondral ossification follows five steps. (a) Mesenchymal cells differentiate into chondrocytes. (b) The cartilage model of the future bony skeleton and the perichondrium form. (c) Capillaries penetrate cartilage. Perichondrium transforms into periosteum. Periosteal collar develops. Primary ossification center develops. (d) Cartilage and chondrocytes continue to grow at ends of the bone. (e) Secondary ossification centers develop. (f) Cartilage remains at epiphyseal (growth) plate and at joint surface as articular cartilage.

The epiphyseal plate is responsible for longitudinal bone growth.

As a bone matures, the epiphyseal plate progresses to an epiphyseal line. (a) Epiphyseal plates are visible in a growing bone. (b) Epiphyseal lines are the remnants of epiphyseal plates in a mature bone.

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  1. All bone formation is a replacement process.
  2. During embryonic development, a cartilaginous skeleton and various membranes are initially formed.
  3. As development progresses, these structures are replaced by bone through the ossification process.
  4. In intramembranous ossification, bone develops directly from sheets of mesenchymal connective tissue.
  5. On the other hand, in endochondral ossification, bone forms by replacing hyaline cartilage.
  6. The epiphyseal plate is responsible for the lengthening of bones during growth, while modeling allows bones to increase in diameter.
  7. Remodeling takes place as bone is resorbed and replaced by new bone.
  8. Osteogenesis imperfecta is a genetic disease which is characterized by altered collagen production, leading to the development of fragile and brittle bones.
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