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Structure and Function

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Structure and Function SECTION I Basic Concepts 1 Structure and Function Alan M. Rosenberg, Ross E. Petty Rheumatic diseases involve abnormalities of multiple organs and response to fluctuating mechanical loads, and as a normal physiolog- systems, but it is predominantly the musculoskeletal system that is ical process throughout life. It is continuously generating new blood affected. The musculoskeletal system consists of bones, joints, carti- cells and contributing to the maintenance of the body’s biochemical lage, tendons, ligaments, muscles, and associated connective tissues. and energy balance. This chapter is an overview of normal musculoskeletal biology. In other sections, pathology of the musculoskeletal system and other sys- Bone tems pertinent to rheumatic diseases, including immune, vascular, and Bone is a composite of organic and inorganic materials. The hardness neurological systems, are discussed in relevant contexts. of bone is a consequence of the interaction of organic constituents such as collagen, proteoglycans, and matrix proteins, which confer ten- THE SKELETON sile strength and permit flexibility to withstand stress, and inorganic materials such as hydroxylapatite and other calcium and phosphate The musculoskeletal system is derived from embryonic mesoderm, a salts, which account for bone’s compressive strength. layer that first appears at approximately the fourth week of gestation. As the embryonic neural crest closes to create the notochord, cells from Classification of Bones the dorsal margins migrate to form the teeth and some facial bones, Bones are categorized based on either their respective shapes and cartilage, and connective tissue of the anterior skull.1 Mesodermal cells attendant functions or the process by which they become ossified. arising from the prechordal region (located rostral to the notochord) Bone classification based on shape and function.All bones form cartilage and bone for the posterior skull, paraxial mesodermal share similar ultrastructural characteristics, but gross anatomical shapes cells (somites) generate the axial skeleton, and lateral plate mesoder- differ, reflecting adaptations to their respective functions. Long bones, mal cells form the appendicular skeleton.1 with a greater length than width, have a thick wall of cortical bone to As with all large organisms, the human endoskeleton provides a withstand and adapt to fluctuating stresses of weight-bearing and to rigid framework, allowing movement to occur with power and pre- allow for rotation and leverage. Short, cuboid-shaped bones, such as cision. The axial skeleton comprises the skull, mandible, auditory carpal bones, support and facilitate movement. Broad flat bones, such as ossicles, hyoid bone, vertebral column, sternum, and ribs, and the cranial bones, sternum, and ribs, are thin and curvilinear; they serve as appendicular skeleton includes the limbs and limb girdles. Ossification attachment sites for musculature and they shield vital organs. Irregular- of the fibrous membranes and hyaline cartilage that constitute the shaped bones protect the face and spinal cord. Sesamoid bones develop primordial skeletal scaffold begins at approximately the seventh gesta- as smooth, round ossifications to counteract compressive forces within tional week, and by birth it is almost, but not entirely, complete. tendons; patellae are the only sesamoid bones consistently present The skeleton is the framework that maintains body shape; sup- in humans, although the hallux sesamoid bone of the foot and the ports muscles and connective tissues, neurovascular networks, and pisiform of the wrist are usually present as well. skin; protects the body’s vital organs; and makes movement possible Bone classification based on ossification process.Bone is through articulations and muscle and tendon attachments. The skel- formed by either intramembranous or endochondral ossification. eton is the repository for hematopoietic and mesenchymal progeni- Intramembranous ossification occurs in facial, cranial, and clavicular tor cells, a source of energy derived from yellow marrow adipocytes, bones as mesenchymal cells within the unossified skeleton coalesce and a mediator of the immune response, and functions as an endocrine then differentiate into either capillaries or osteoblasts. The osteoblasts organ contributing to calcium, phosphate, and glucose homeosta- secrete unmineralized osteoid matrix into which mineral salts (notably sis.2-7 Further, the musculoskeletal system aids other bodily functions, calcium and phosphorus) are then deposited. When ossified, osteoid including digestion by mastication, respiration by moistening inspired produced around capillaries becomes honeycomb-like trabecular bone air passing through bone sinuses, and communication by way of audi- (also referred to as cancellous or spongy bone), which encroaches on the tory ossicles for hearing and the larynx for vocalization. vasculature, compacting it into red marrow. The skeleton is a dynamic organ both anatomically and physio- Endochondral ossification occurs in most appendicular and axial logically. It is modified structurally during growth and maturation, in skeleton bones. Endochondral ossification involves the progressive 1 2 SECTION I Basic Concepts Circumferential lamellae Concentric lamellae of osteon Osteon Osteocytes Central (haversian) canal Neurovascular bundle Perforating (Volkmann’s) canal Periosteum Compact bone Spongy bone Fig. 1.1 A depiction of the ultrastructural characteristics of bone. For clarity, 2 or 3 concentric lamellae are shown to represent the osteon; normally, the osteon is composed of 5 to 20 concentric lamellae. (Drawing by Alexandru Margarit and labeling by Alan Rosenberg. Printed with permission. All rights reserved.) replacement of the primordial cartilaginous scaffold with bone through as the bones of the leg, tend to be cylindrical with a thick collar of cor- a sequence in which chondrocytes proliferate, mature, and enlarge. By tical bone and a marrow cavity. 8 weeks gestational age some mesenchymal cells have differentiated Type I collagen constitutes 90% of bone matrix. The triple heli- into cartilage-producing chondrocytes that create the cartilaginous cal strands of collagen, when densely packed into parallel alignments skeletal framework. Building the skeletal scaffold begins as mesen- (lamellae), confer optimal tensile strength to accommodate mechani- chymal stem cells form condensations at the site of future bone. Cells cal loading. Other organic constituents of bone contribute to regulat- within the condensations do not differentiate directly into bone-pro- ing mineralization, growth, and energy metabolism. ducing osteoblasts; instead, they become proliferating chondrocytes, The circular osteon is the fundamental physiological unit of most which synthesize the extracellular matrix comprising predominantly mammalian compact bone (Fig. 1.1). The microstructure of the osteon type II collagen, some type IX and XI collagens, and proteoglycans. is characterized by 5 to 20 concentric layers of compact (cortical) bone Then, as mature, hypertrophied chondrocytes, they secrete a matrix that encircle a central haversian canal (first identified by 17th-century rich in type X collagen, which provides the supporting framework for English physician Clopton Havers), through which neurovascular bun- endochondral ossification. dles traverse to supply the adjacent osteocytes (Fig. 1.1). Osteons, mea- As the cartilage matrix ossifies, blood flow to the chondrocytes is suring a few millimeters in length and less than 0.25 mm in diameter, impeded, the cells die, and adjacent cartilage breaks down, leaving a tend to orient to the long axis of the bone. Osteons are characteristically medullary cavity into which new vessels invade to deliver bone-pro- present in mature bone that has undergone new bone growth, remodel- ducing osteoblasts. While ossification continues from the primary ing, or repair. Osteons and their associated intra-haversian blood vessels ossification center in the diaphysis, cartilage continues to be produced communicate by interconnecting transverse Volkmann canals (named at the ends of bones as the epiphyses. At birth, cartilage remains at the for 19th-century German physiologist Alfred Volkmann). articular surface and the growth plate (the physis) situated between the Deeper layers of bone are made of spongy bone, the normal physio- epiphysis and the diaphysis. A mature long bone includes the epiphysis logical consequence of bone breakdown induced by osteoclasts. at the end of the bone, the diaphysis (the shaft of the bone), and the interposing metaphysis, which represents the region of the previous Bone Cells cartilaginous growth plate. Apophyses, such as the tibial tuberosity, Bone contains four cell types: mesenchymal stem cells, osteoblasts, posterior calcaneus, greater and lesser trochanters, and iliac crests, like osteocytes, and osteoclasts. Mesenchymal stem cells, located in the epiphyses, are the sites of new bone formation but do not contribute to intertrabecular loose connective tissue adjacent to vascular channels bone length; instead, they generate new bone in response to tendinous and in the periosteum, have the potential to differentiate into cells of or ligamentous traction. bone (osteoblasts, osteocytes, and osteoclasts), cartilage (chondro- cytes), muscle (myocytes), or fat (adipocytes). The Morphology and Physiology of Bone The differentiation of mesenchymal stem cells into osteoclasts is Bone includes an outer layer of thick, compact,
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