211 ch 6 bone Bone Tissue Chapter 6 Skeletal Cartilages • All contain chondrocytes in lacunae and extracellular matrix • Three types – Hyaline cartilage • Provides support, flexibility, and resilience • Collagen fibers only; most abundant type • Articular, costal, respiratory, nasal cartilage – Elastic cartilage • Similar to hyaline cartilage, but contains elastic fibers • External ear and epiglottis – Fibrocartilage • Thick collagen fibers—has great tensile strength • Menisci of knee; vertebral discs Classification of Bones • 206 named bones in skeleton • Divided into two groups – Axial skeleton • Long axis of body • Skull, vertebral column, rib cage – Appendicular skeleton • Bones of upper and lower limbs • Girdles attaching limbs to axial skeleton Classification of Bones by Shape • Long bones – Longer than they are wide – Limb, wrist, ankle bones 1 211 ch 6 bone • Short bones – Cube-shaped bones (in wrist and ankle) – Sesamoid bones (within tendons, e.g., Patella) – Vary in size and number in different individuals • Flat bones – Thin, flat, slightly curved – Sternum, scapulae, ribs, most skull bones • Irregular bones – Complicated shapes – Vertebrae, coxal bones Functions of Bones • Support – For body and soft organs • Protection – For brain, spinal cord, and vital organs • Movement – Levers for muscle action • Mineral and growth factor storage – Calcium and phosphorus, and growth factors reservoir • Blood cell formation (hematopoiesis) in red marrow cavities of certain bones • Triglyceride (fat) storage in bone cavities – Energy source • Hormone production – Osteocalcin • Regulates bone formation • Protects against obesity, glucose intolerance, diabetes mellitus Bones • Are organs – Contain different types of tissues • Bone (osseous) tissue, nervous tissue, cartilage, fibrous 2 211 ch 6 bone connective tissue, muscle and epithelial cells in its blood vessels • Three levels of structure – Gross anatomy – Microscopic – Chemical Gross Anatomy • Bone textures – Compact and spongy bone • Compact – Dense outer layer; smooth and solid • Spongy (cancellous or trabecular) – Honeycomb of flat pieces of bone deep to compact called trabeculae Structure of Short, Irregular, and Flat Bones • Thin plates of spongy bone covered by compact bone • Plates sandwiched between connective tissue membranes – Periosteum (outer layer) and endosteum • No shaft or epiphyses • Bone marrow throughout spongy bone; no marrow cavity • Hyaline cartilage covers articular surfaces Structure of Typical Long Bone • Diaphysis – Tubular shaft forms long axis – Compact bone surrounding medullary cavity • Epiphyses – Bone ends – External compact bone; internal spongy bone – Articular cartilage covers articular surfaces 3 211 ch 6 bone – Between is epiphyseal line • Remnant of childhood bone growth at epiphyseal plate Membranes: Periosteum • White, double-layered membrane • Covers external surfaces except joint surfaces • Outer fibrous layer of dense irregular connective tissue – Perforating/Sharpey's fibers secure to bone matrix • Osteogenic layer abuts bone – Contains primitive stem cells – osteogenic cells • Many nerve fibers and blood vessels • Anchoring points for tendons and ligaments Membranes: Endosteum • Delicate connective tissue membrane covering internal bone surface • Covers trabeculae of spongy bone • Lines canals that pass through compact bone • Contains osteogenic cells that can differentiate into other bone cells Hematopoietic Tissue in Bones • Red marrow – Found within trabecular cavities of spongy bone and diploë of flat bones (e.g., Sternum) – In medullary cavities and spongy bone of newborns – Adult long bones have little red marrow • Heads of femur and humerus only – Red marrow in diploë and some irregular bones is most active – Yellow marrow can convert to red, if necessary Bone Markings 4 211 ch 6 bone • Sites of muscle, ligament, and tendon attachment on external • • • • • surfaces Joint surfaces Conduits for blood vessels and nerves Projections Depressions Openings Microscopic Anatomy of Bone: Cells of Bone Tissue • Five major cell types • Each specialized form of same basic cell type – Osteogenic cells – Osteoblasts – Osteocytes – Bone lining cells – Osteoclasts Osteogenic Cells • Also called osteoprogenitor cells – Mitotically active stem cells in periosteum and endosteum – When stimulated differentiate into osteoblasts or bone lining cells • Some persist as osteogenic cells Osteoblasts • Bone-forming cells • Secrete unmineralized bone matrix or osteoid – Includes collagen and calcium-binding proteins • Collagen = 90% of bone protein • Actively mitotic 5 211 ch 6 bone Osteocytes • Mature bone cells in lacunae • Monitor and maintain bone matrix • Act as stress or strain sensors – Respond to and communicate mechanical stimuli to osteoblasts and osteoclasts (cells that destroy bone) so bone remodeling can occur Bone Lining Cells • Flat cells on bone surfaces believed to help maintain matrix • On external bone surface called periosteal cells • Lining internal surfaces called endosteal cells Osteoclasts • Derived from hematopoietic stem cells that become macrophages • Giant, multinucleate cells for bone resorption • When active rest in resorption bay and have ruffled border – Ruffled border increases surface area for enzyme degradation of bone and seals off area from surrounding matrix Microscopic Anatomy of Bone: Compact Bone • Also called lamellar bone • Osteon or haversian system – Structural unit of compact bone – Elongated cylinder parallel to long axis of bone – Hollow tubes of bone matrix called lamellae • Collagen fibers in adjacent rings run in different directions – Withstands stress – resist twisting Microscopic Anatomy of Bone: Compact Bone 6 211 ch 6 bone • Canals and canaliculi – Central (haversian) canal runs through core of osteon • Contains blood vessels and nerve fibers • Perforating (volkmann's) canals – Canals lined with endosteum at right angles to central canal – Connect blood vessels and nerves of periosteum, medullary cavity, and central canal • Lacunae—small cavities that contain osteocytes • Canaliculi—hairlike canals that connect lacunae to each other and central canal Canaliculi Formation • Osteoblasts secreting bone matrix maintain contact with each other and osteocytes via cell projections with gap junctions • When matrix hardens and cells are trapped the canaliculi form – Allow communication – Permit nutrients and wastes to be relayed from one osteocyte to another throughout osteon Lamellae • Interstitial lamellae – Incomplete lamellae not part of complete osteon – Fill gaps between forming osteons – Remnants of osteons cut by bone remodeling • Circumferential lamellae – Just deep to periosteum – Superficial to endosteum – Extend around entire surface of diaphysis – Resist twisting of long bone Microscopic Anatomy of Bone: Spongy Bone • Appears poorly organized • Trabeculae 7 211 ch 6 bone – Align along lines of stress to help resist it – No osteons – Contain irregularly arranged lamellae and osteocytes interconnected by canaliculi – Capillaries in endosteum supply nutrients Chemical Composition of Bone: Organic Components • Includes cells and osteoid – Osteogenic cells, osteoblasts, osteocytes, bone- lining cells, and osteoclasts – Osteoid—1/3 of organic bone matrix secreted by osteoblasts • Made of ground substance (proteoglycans and glycoproteins) • Collagen fibers • Contributes to structure; provides tensile strength and flexibility • Resilience of bone due to (sacrificial) bonds in or between collagen molecules – Stretch and break easily on impact to dissipate energy and prevent fracture – If no addition trauma, bonds re-form Chemical Composition of Bone: Inorganic Components • Hydroxyapatites (mineral salts) – 65% of bone by mass – Mainly of tiny calcium phosphate crystals in and around collagen fibers – Responsible for hardness and resistance to compression Bone • Half as strong as steel in resisting compression • As strong as steel in resisting tension • Last long after death because of mineral composition – Reveal information about ancient people – Can display growth arrest lines 8 211 ch 6 bone • Horizontal lines on bones • Proof of illness - when bones stop growing so nutrients can help fight disease Bone Development • Ossification (osteogenesis) – Process of bone tissue formation – Formation of bony skeleton • Begins in 2nd month of development – Postnatal bone growth • Until early adulthood – Bone remodeling and repair • Lifelong Two Types of Ossification • Endochondral ossification – Bone forms by replacing hyaline cartilage – Bones called cartilage (endochondral) bones – Forms most of skeleton • Intramembranous ossification – Bone develops from fibrous membrane – Bones called membrane bones – Forms flat bones, e.g. clavicles and cranial bones Endochondral Ossification • Forms most all bones inferior to base of skull – Except clavicles • Begins late in 2nd month of development • Uses hyaline cartilage models • Requires breakdown of hyaline cartilage prior to ossification Endochondral Ossification • Begins at primary ossification center in center of shaft 9 211 ch 6 bone – Blood vessel infiltration of perichondrium converts it to • • • • • periosteum underlying cells change to osteoblasts Bone collar forms around diaphysis of cartilage model Central cartilage in diaphysis calcifies, then develops cavities Periosteal bud invades cavities formation of spongy bone Diaphysis elongates & medullary cavity forms Epiphyses ossify Intramembranous Ossification • Forms frontal, parietal, occipital, temporal bones, and clavicles • Begins within fibrous connective tissue membranes formed by mesenchymal cells • Ossification centers appear • Osteoid is secreted • Woven bone and periosteum form • Lamellar bone replaces woven bone & red marrow appears Postnatal Bone Growth • Interstitial (longitudinal) growth – Increase in length of long bones • Appositional growth – Increase in bone thickness Interstitial Growth: Growth in Length of Long Bones • Requires presence of epiphyseal cartilage • Epiphyseal plate maintains constant thickness – Rate of cartilage growth on one side balanced by bone replacement on other • Concurrent remodeling of epiphyseal ends to maintain proportion • Calcification zone – Surrounding cartilage matrix calcifies, chondrocytes die and deteriorate 10 211 ch 6 bone • Ossification zone – Chondrocyte deterioration leaves long spicules of calcified cartilage at epiphysis-diaphysis junction – Spicules eroded by osteoclasts – Covered with new bone by osteoblasts – Ultimately replaced with spongy bone Interstitial Growth: Growth in Length of Long Bones • Near end of adolescence chondroblasts divide less often • Epiphyseal plate thins then is replaced by bone • Epiphyseal plate closure – Bone lengthening ceases • Requires presence of cartilage – Bone of epiphysis and diaphysis fuses – Females – about 18 years – Males – about 21 years Appositional Growth: Growth in Width • Allows lengthening bone to widen • Occurs throughout life • Osteoblasts beneath periosteum secrete bone matrix on external bone • Osteoclasts remove bone on endosteal surface • Usually more building up than breaking down – Thicker, stronger bone but not too heavy Hormonal Regulation of Bone Growth • Growth hormone – Most important in stimulating epiphyseal plate activity in infancy and childhood • Thyroid hormone – Modulates activity of growth hormone 11 211 ch 6 bone Ensures proper proportions • Testosterone (males) and estrogens (females) at puberty – Promote adolescent growth spurts – End growth by inducing epiphyseal plate closure • Excesses or deficits of any cause abnormal skeletal growth – Bone Homeostasis • Recycle 5-7% of bone mass each week – Spongy bone replaced ~ every 3-4 years – Compact bone replaced ~ every 10 years • Older bone becomes more brittle – Calcium salts crystallize – Fractures more easily • Consists of bone remodeling and bone repair Bone Homeostasis: Bone Remodeling • Consists of both bone deposit and bone resorption • Occurs at surfaces of both periosteum and endosteum • Remodeling units – Adjacent osteoblasts and osteoclasts Bone Deposit • Evidence of new matrix deposit by osteoblasts • Trigger not confirmed Bone Resorption • Is function of osteoclasts – Dig depressions or grooves as break down matrix – Secrete lysosomal enzymes that digest matrix and protons (H+) – Acidity converts calcium salts to soluble forms • Osteoclasts also – Phagocytize demineralized matrix and dead osteocytes 12 211 ch 6 bone Transcytosis allow release into interstitial fluid and then into blood – Once resorption complete, osteoclasts undergo apoptosis • Osteoclast activation involves PTH and T cell-secreted proteins • Control of Remodeling • Occurs continuously but regulated by genetic factors and two control loops – Negative feedback hormonal loop for Ca2+ homeostasis • Controls blood Ca2+ levels; Not bone integrity – Responses to mechanical and gravitational forces Importance of Calcium • Functions in – Nerve impulse transmission – Muscle contraction – Blood coagulation – Secretion by glands and nerve cells – Cell division • 1200 – 1400 grams of calcium in body – 99% as bone minerals – Amount in blood tightly regulated (9-11 mg/dl) – Intestinal absorption requires Vitamin D metabolites – Dietary intake required Hormonal Control of Blood Ca2+ • Parathyroid hormone (PTH) – Produced by parathyroid glands – Removes calcium from bone regardless of bone integrity • Calcitonin may be involved – Produced by parafollicular cells of thyroid gland – In high doses lowers blood calcium levels temporarily Negative Feedback Hormonal Loop for blood Ca2+ Homeostasis Controlled by parathyroid hormone (PTH) 13 211 ch 6 bone Blood Ca2+ levels PTH release PTH stimulates osteoclasts to degrade bone matrix, releasing Ca2+ Blood Ca2+ levels PTH release ends Bone Homeostasis: Response to Mechanical Stress • Bones reflect stresses they encounter – Long bones thickest midway along diaphysis where bending stresses greatest • Bones stressed when weight bears on them or muscles pull on them – Usually off center so tends to bend bones – Bending compresses on one side; stretches on other Results of Mechanical Stressors: Wolff's Law • Bones grow or remodel in response to demands placed on it • Explains – Handedness (right or left handed) results in thicker and stronger bone of that upper limb – Curved bones thickest where most likely to buckle – Trabeculae form trusses along lines of stress – Large, bony projections occur where heavy, active muscles attach – Bones of fetus and bedridden featureless 14 211 ch 6 bone How Mechanical Stress Causes Remodeling • Electrical signals produced by deforming bone may cause remodeling – Compressed and stretched regions oppositely charged • Fluid flows within canaliculi appear to provide remodeling stimulus Homeostatic Imbalances • Osteomalacia – Calcium salts not adequate • Rickets (osteomalacia of children) – Cause: Vitamin D deficiency or insufficient dietary calcium –Osteoporosis – Sex hormones maintain normal bone health and density • As secretion wanes with age osteoporosis can develop • Diet poor in calcium and protein • Hormone-related conditions – Hyperthyroidism – Low blood levels of thyroid-stimulating hormone – Diabetes mellitus • Males with prostate cancer taking androgen-suppressing drugs – Electrical stimulation; Daily ultrasound treatments hasten repair 15