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Chapter 6 The Skeletal System Copyright 2010, John Wiley & Sons, Inc. Bone Function Support Protection - Skull, rib cage Assist in movements - together with muscles & tendons Mineral homeostasis - calcium, magnesium, phosphate Blood cell production Hemopoiesis in red bone marrow Triglyceride (fat) storage - in yellow bone marrow 2 Million year old hand skeleton of possible early human ancestral hominid Types of Bones Long bones: longer than wide Short bones: almost cube shaped Most wrist and ankle bones Flat bones: thin and extensive surface Such as thigh, leg, arm, forearm, fingers and toes Such as cranial bones sternum, ribs and scapulas Irregular bones: do not fit above categories Such as vertebrae and some facial bones Copyright 2010, John Wiley & Sons, Inc. Macroscopic Structure Parts of a long bone Diaphysis: shaft of long bone; made up mostly of compact bone Epiphysis: broad end of long bone; mostly spongy bone Metaphysis: growth area between diaphysis and epiphysis Articular cartilage: hyaline cartilage at joint Periosteum: fibrous covering over most of bone Medullary cavity (marrow) with fat and blood cells Endosteum: membrane lining medullary cavity Copyright 2010, John Wiley & Sons, Inc. Long Bones Copyright 2010, John Wiley & Sons, Inc. Long Bones Copyright 2010, John Wiley & Sons, Inc. Microscopic Structure of Bone Matrix 25% water, 25% collagen fibers, 50% mineral salts Cells Osteogenic cells in periosteum Osteoblasts Secrete collagen fibers Build matrix and become trapped in lacunae Become Osteocytes that maintain bone Osteoclasts are formed from monocytes Digest bone matrix for normal bone turnover Copyright 2010, John Wiley & Sons, Inc. Histology of Bones Copyright 2010, John Wiley & Sons, Inc. Compact Bone Structure Arranged in osteons (haversian systems) Central canal through center of osteon Cylinders running parallel to long axis of bone Contains blood vessels, nerves, lymphatics Concentric lamellae: layers of matrix Lacunae: “lakes” between lamellae Contain osteocytes (bone cells) Copyright 2010, John Wiley & Sons, Inc. Compact Bone Structure Canaliculi (“little canals”) Contain extensions of osteocytes Permit flow of ECF between central canal and lacunae Compact bone is covered by periosteum Perforating (Volkmann’s) canals Carry blood and lymphatic vessels and nerves from periosteum They supply central (Haversian) canals and also bone marrow Copyright 2010, John Wiley & Sons, Inc. Histology of Bones Copyright 2010, John Wiley & Sons, Inc. Copyright 2010, John Wiley & Sons, Inc. Spongy Bone Not arranged in osteons, but irregular latticework of trabeculae These contain lacunae with osteocytes and canaliculi Spaces between trabeculae may contain red bone marrow Spongy bone is lighter than compact bone, so reduces weight of skeleton Copyright 2010, John Wiley & Sons, Inc. Bone Dynamics and Tissue Interactions Animation Bone Dynamics and Tissue You must be connected to the internet to run this animation. Copyright 2010, John Wiley & Sons, Inc. Bone Formation Known as ossification Timeline Initial bone development in embryo and fetus Growth of bone into adulthood Remodeling: replacement of old bone Repair if fractures occur Mesenchyme (early connective tissue) model This initial “skeleton” model will be replaced by bone tissue beginning at 6 weeks of embryonic life Copyright 2010, John Wiley & Sons, Inc. Bone Formation Two different methods of ossification each result in similar bone tissue 1. Intramembranous: Bone forms within sheets of mesenchyme that resemble membranes Only a few bones form by this process: flat bones of the skull, lower jawbone (mandible), and part of clavicle (collarbone) 2. Endochondrial: Mesenchyme forms hyaline cartilage which then develops into bone All other bones form by this process Copyright 2010, John Wiley & Sons, Inc. Flat bone of skull Blood capillary Ossification center Mesenchymal cell Osteoblast Mandible Collagen fiber 1 Development of ossification center Mesenchyme condenses Osteocyte in lacuna Blood vessel Canaliculus Spongy bone trabeculae Osteoblast Osteoblast Newly calcified bone matrix 2 Calcification 3 Formation of trabeculae Periosteum Spongy bone tissue Compact bone tissue Copyright 2010, John Wiley & Sons, Inc. 4 Development of the periosteum Intramembranous Ossification Four steps 1.Development of ossification center Mesenchyme cells osteogenic osteoblasts Osteoblasts secrete organic matrix 2. Calcification: cells become osteocytes In lacunae they extend cytoplasmic processes to each other Deposit calcium & other mineral salts 3.Formation of trabeculae (spongy bone) Blood vessels grow in and red marrow is formed 4. Periosteum covering the bone forms from mesenchyme Copyright 2010, John Wiley & Sons, Inc. Perichondrium Proximal epiphysis Uncalcified matrix Hyaline cartilage Periosteum Uncalcified matrix Diaphysis Calcified matrix Primary ossification center Nutrient artery Spongy bone Distal epiphysis Calcified matrix Periosteum (covering compact bone) Medullary cavity Nutrient artery and vein 1 Development of cartilage model 2 Growth of cartilage model 3 Development of primary ossification center 4 Development of the medullary cavity Articular cartilage Secondary ossification center Epiphyseal artery and vein Spongy bone Uncalcified matrix Epiphyseal plate Nutrient artery and vein Copyright 2010, John 5 Development of secondary ossification center Wiley6&Formation Sons, Inc.of articular cartilage and epiphyseal plate Endochondrial Ossification Six Steps 1. Formation of cartilage model of the “bone” As mesenchyme cells develop into chondroblasts 2. Growth of cartilage model Cartilage “bone” grows as chondroblasts secrete cartilage matrix Chondrocytes increase in size, matrix around them calcifies Chondrocytes die as they are cut off from nutrients, leaving small spaces (lacunae) Copyright 2010, John Wiley & Sons, Inc. Endochondrial Ossification Six Steps 3. Primary ossification center Perichondrium sends nutrient artery inwards into disintegrating cartilage Osteogenic cells in perichondrium become osteoblasts that deposit bony matrix over remnants of calcified cartilage spongy bone forms in center of the model As perichondrium starts to form bone, the membrane is called periosteum Copyright 2010, John Wiley & Sons, Inc. Endochondrial Ossification Six Steps 4. Medullary (marrow) cavity Spongy bone in center of the model grows towards ends of model Octeoclasts break down some of new spongy bone forming a cavity (marrow) through most of diaphysis Most of the wall of the diaphysis is replaced by a collar of compact bone Copyright 2010, John Wiley & Sons, Inc. Endochondrial Ossification Six Steps 5. Secondary ossification center Similar to step 3 except that nutrient arteries enter ends (epiphyses) of bones and osteoblasts deposit bony matrix spongy bone forms in epiphyses from center outwards Occurs about time of birth 6. Articular cartilage and epiphyseal cartilage Articular cartilage at ends of epiphyses becomes articular cartilage Epiphyseal (growth) plate of cartilage remains between epiphysis and diaphysis until bone growth ceases Copyright 2010, John Wiley & Sons, Inc. Growth in Length Chondrocytes divide and grow more cartilage on epiphyseal side of the epiphyseal plate Chondrocytes on the diaphyseal side die and are replaced by bone Therefore bone grows from diaphyseal side towards epiphyseal side Growth in length stops between 18-25 years; cartilage in epiphyseal plate is completely replaced by bone (epiphyseal line) Copyright 2010, John Wiley & Sons, Inc. Growth in Thickness As bones grow in length, they must also grow in thickness (width) Perichondrial osteoblasts osteoblasts lay down additional lamellae of compact bone Simultaneously, osteoclasts in the endosteum destroy interior bone to increase width of the marrow Copyright 2010, John Wiley & Sons, Inc. Remodeling and Repair Remodeling in response to use Resorption of bone matrix by osteoclasts and Deposition of bone matrix by osteoblasts Repair after a fracture Dead tissue removed Chondroblasts fibrocartilage spongy bone deposited by osteoblasts remodeled to compact bone Copyright 2010, John Wiley & Sons, Inc. Types of Fractures Partial: incomplete break (crack) Complete: bone broken into two or more pieces Closed (simple): not through skin Open (compound): broken ends break skin Copyright 2010, John Wiley & Sons, Inc. Factors Affecting Growth Adequate minerals (Ca, P, Mg) Vitamins A, C, D Hormones Before puberty: hGH + insulin-like growth factors Thyroid hormone and insulin also required Sex hormones contribute to adolescent growth spurt Weight-bearing activity Copyright 2010, John Wiley & Sons, Inc. Copyright 2010, John Wiley & Sons, Inc. Calcium Homeostasis Blood levels of Ca2+ controlled Negative feedback loops Parathyroid hormone (PTH) released by parathyroid gland increases osteoclast activity + decreases loss of Ca2+ in urine Calcitonin decreases osteoclast activity Copyright 2010, John Wiley & Sons, Inc. Negative Feedback Copyright 2010, John Wiley & Sons, Inc. Exercise & Bone Tissue Bone strengthened in response to use Bone resorbed during disuse; examples: During prolonged bed rest Fracture with cast/immobilizer Astronauts without gravity Copyright 2010, John Wiley & Sons, Inc. Divisions of Skeletal System Two divisions: axial and appendicular Axial: bones around body axis Examples: skull bones, hyoid, ribs, sternum, vertebrae Appendicular: bones of upper and lower limbs plus shoulder and hip bones that connect them Examples: collar bone (clavicle), arm (humerus), forearm (radius and ulna), thigh bone (femur) Copyright 2010, John Wiley & Sons, Inc. Divisions of the Skeletal System There are 206 skeletal bones in the human skeleton - 80 bones of the axial skeleton - Skull (22) - Hyoid (1) - Auditory ossicles (6) - Vertebral column (26) - Rib cage (25) - 126 bones make up the appendicular skeleton - Pectoral (shoulder) girdle (4) - Upper limb bones (60) - Pelvic (hip) girdle (2) - Lower limb bones (48) Bone markings 1. Bone Projections 1. Tuberosity - large, rounded projection e.g. Tibial tuberosity 2. Crest - narrow bone ridge, usually prominent e.g. Anterior crest of tibia 3. Trochanter - very large, blunt, irregularly shaped process, e.g. Femur 4. Line - narrow bone ridge, less prominent than crest; e.g. Intertrochanteric line 5. Tubercle - small rounded projection or process e.g. Adductor tubercle (on medial femur) 6. Epicondyle - Raised area on or above a condyle e.g. Medial epicondyle (on femur) 7. Spine - sharp, slender, often pointed projection e.g. Vertebra Copyright 2010, John Wiley & Sons, Inc. Bone markings (Contd.) Bone depressions & Openings 1. Foramen - round or oval bone opening e.g. Optic foramen or Foramen magnum 2. Meatus - canal-like passage way e.g. External acoustic meatus (skull) 3. Fossa - shallow, basin-like depression e.g. Mandibular fossa 4. Groove - slit-like furrow e.g. Mandible 5. Fissure - narrow, slit-like opening 6. Notch - shallow depression e.g. Clavicular notch 7. Body - large, rough-surfaced depression e.g. Body of vertebrae Copyright 2010, John Wiley & Sons, Inc. Bone markings (Contd.) Processes that help to form joints 1. Condyle - large rounded bone prominence e.g. Mandibular condyle 2. Head - bony expansion e.g. Head on rib bones 3. Facet - smooth, nearly flat articular surface e.g. Rib facets, articular facets (vertebrae) Bone cavities 1. Sinus - hollow sections of bones filled with air, lined with mucous membranes e.g. ethmoid or sphenoid sinus Copyright 2010, John Wiley & Sons, Inc. Skull & Hyoid Bone Eight Cranial bones Frontal, 2 parietal, 2 temporal, occipital, sphenoid, and ethmoid Fourteen Facial bones 2 nasal, 2 maxilla, 2 zygomatic, 2 lacrimal 2 palatine, 2 inferior nasal conchae, 1 mandible,1 vomer Copyright 2010, John Wiley & Sons, Inc. Skull Copyright 2010, John Wiley & Sons, Inc. Skull Copyright 2010, John Wiley & Sons, Inc. Skull Copyright 2010, John Wiley & Sons, Inc. Skull Copyright 2010, John Wiley & Sons, Inc. Sphenoid Bone Copyright 2010, John Wiley & Sons, Inc. Ethmoid Bone Copyright 2010, John Wiley & Sons, Inc. Ethmoid Bone Copyright 2010, John Wiley & Sons, Inc. Unique Features of Skull Sutures: immovable joint between skull bones Paranasal sinuses: cavities Coronal, sagittal, lambdoidal, squamous Located in bones near nasal cavity Fontanels: soft spot in fetal skull Allow deformation at birth Calcify to form sutures Copyright 2010, John Wiley & Sons, Inc. Sinuses Found in four skull bones - maxillary, sphenoid, ethmoid and frontal bone - are mucosa-lined air cavities within the skull; - all lead into the nasal passages - help to lighten the weight of the facial bones, - serve as resonance chambers for speech - may also help to insulate the brain from cold effects - largest is the maxillary sinus Paranasal Sinuses Copyright 2010, John Wiley & Sons, Inc. Vertebrae Functions Encloses spinal cord Supports head Point of attachment for muscles of back, ribs and pelvic girdle Regions (from superior to inferior) 7 cervical 12 thoracic 5 lumbar 1 sacrum and 1 coccyx Copyright 2010, John Wiley & Sons, Inc. Normal Curves in Column Four normal curves Cervical and lumbar curves are convex (bulge anteriorly) Thoracic and sacral curves are concave (bulge posteriorly) Curves increase strength, help in balance and absorb shocks Copyright 2010, John Wiley & Sons, Inc. Vertebral Column Copyright 2010, John Wiley & Sons, Inc. Vertebral Column Copyright 2010, John Wiley & Sons, Inc. Structure of Vertebra Body: disc-shaped anterior portion Vertebral arch: posteriorly back from body With the body, creates a hole called vertebral foramen Seven processes from this arch Transverse process extending laterally on each side Spinous process extending dorsally Two each of superior and inferior articular processes that form joints with vertebrae Copyright 2010, John Wiley & Sons, Inc. Structure of Vertebra Copyright 2010, John Wiley & Sons, Inc. Cervical Area Cervical (C1-C7 from superior to inferior) C1: atlas Spinous process often bifid with transverse foramina on transverse processes Articulates with head, specialized to support head Lacks body and spinous process C2: axis Has body and spinous process Called dens (“tooth”) that creates a pivot for head rotation Copyright 2010, John Wiley & Sons, Inc. Cervical Vertebrae Copyright 2010, John Wiley & Sons, Inc. Other Vertebrae Thoracic (T1-T12 ) Lumbar (L1-L5) Largest and strongest; spinous processes short and thick Sacrum (S1-S5 fused into one unit) Larger than cervical Have facets for articulations with ribs Foundation for pelvic girdle Contain sacral foramina for nerves and blood vessels Coccyx: 4 coccygeal vertebrae fused into 1 Copyright 2010, John Wiley & Sons, Inc. Sacrum bone markings One important bony landmark of the sacrum is the sacral cornua which allows the location of the sacral hiatus (located between the cornua); - anesthetic drugs that act on the sacral and coccygeal nerves are often injected through the sacral hiatus; - this procedure is called caudal anesthesia or epidural block; - often used in the clinic to relieve pain during labor and to provide anesthesia to the perineal area - anesthetic drugs may also be injected through the posterior sacral foramina Copyright 2010, John Wiley & Sons, Inc. Lumbar Vertebrae Copyright 2010, John Wiley & Sons, Inc. Sacrum and Coccyx Copyright 2010, John Wiley & Sons, Inc. Thorax Thoracic cage: sternum, costal cartilages, ribs and bodies of T1-T12 Sternum: form by 3 portions fused by about age 25 years: Manubrium, body, xiphoid process Ribs: 12 pairs True ribs are #1-7: articulate with sternum directly by costal cartilages False ribs are #8-12: do not articulate with sternum directly by costal cartilages Copyright 2010, John Wiley & Sons, Inc. Thorax Copyright 2010, John Wiley & Sons, Inc. Pectoral Girdle Function: attach bones of upper limbs to axial skeleton Clavicles (2) and scapulas (2): bilateral Copyright 2010, John Wiley & Sons, Inc. Right Pectoral (Shoulder) Girdle Copyright 2010, John Wiley & Sons, Inc. Upper Limb Humerus (arm bone) - the longest and largest bone of the upper limbs - important bone markings: deltoid tuberosity, anatomical neck, radial fossa Ulna - located on the medial aspect = middle fingerside - important bone markings are: olecranon (forms prominence of the elbow, coronoid process, trochlear notch, radial notch, styloid process Radius - located on the lateral aspect = thumb side - bone markings: head, radial tuberosity (biceps brachii muscle attachment), styloid process Copyright 2010, John Wiley & Sons, Inc. Right Humerus Copyright 2010, John Wiley & Sons, Inc. Right Ulna and Radius Copyright 2010, John Wiley & Sons, Inc. Right Ulna and Radius Copyright 2010, John Wiley & Sons, Inc. Wrist and Hand Carpus (wrist): 8 bones Metacarpals: 5 bones of palm of hand Number 1-5 starting with thumb Phalanges: 14 bones of fingers Numbered 1-5 metacarpals Each finger except the thumb has proximal, middle and distal phalanges; thumb lacks middle phalanx Copyright 2010, John Wiley & Sons, Inc. Carpal bones 8 bones held together by ligaments four carpal bones in each row top row: scaphoid, lunate, triquetrum, pisiform “Scaphoid bone is broken in about 70% of all cases of carpal fractures ..!" lower row: trapezium, trapezoid, capitate, hamate Pisiform and hamate (ulnar side) & scaphoid and trapezium (radial side) form carpal tunnel which allows passage of the long digital and thumb flexor tendons and of the median nerve; narrowing causes the painful disorder carpal tunnel syndrome (CTS).." Right Wrist and Hand Copyright 2010, John Wiley & Sons, Inc. Pelvic (Hip) Girdle Pelvic girdle includes two hip (coxal) bones Joined anteriorly at pubic symphysis Posteriorly attached to sacrum at sacroiliac joint Basin-like pelvis is formed by two hip bones (pelvic girdle) + sacrum and coccyx False (greater) pelvis: broad region superior to pelvic brim; contains abdominal organs True (lesser) pelvis: small region inferior to pelvic brim; contains urinary bladder + internal reproductive organs Copyright 2010, John Wiley & Sons, Inc. Pelvic Girdle (Female) Copyright 2010, John Wiley & Sons, Inc. Pelvic Girdle (Female) Copyright 2010, John Wiley & Sons, Inc. Parts of Each Hip (Coxal) Bone 3 separate bones fuse by age 23 to form a hip bone Ilium: largest and most superior Ischium: lower posterior part Pubis: lower anterior part Bones meet at the acetaulum of the hip bone (socket for head of femur) Copyright 2010, John Wiley & Sons, Inc. Right Hip Bone Copyright 2010, John Wiley & Sons, Inc. Lower Limb Femur (thigh bone): largest bone in the body Patella: kneecap in anterior of knee joint Tibia: shin bone Articulates with hip proximally and with the tibia and patella distally Head (fits into acetabulum) and greater trochanter at proximal end Large medial, weight-bearing bone of leg Fibula: longest, thinnest bone in body Lateral to tibia and smaller Does not articulate with femur Copyright 2010, John Wiley & Sons, Inc. Right Femur Copyright 2010, John Wiley & Sons, Inc. Right Tibia and Fibula Copyright 2010, John Wiley & Sons, Inc. Ankle and Foot Tarsus (ankle) has 7 bones Metatarsals (foot bones) Large talus (ankle bone) and Calcaneus (heel bone) Numbered 1 to 5 from medial to lateral Phalanges (toe bones) Big toe has proximal and distal phalanges while others have proximal, medial and distal phalanges. Numbered like metatarsals from 1-5 Copyright 2010, John Wiley & Sons, Inc. Right Foot Copyright 2010, John Wiley & Sons, Inc. Arches of the Right Foot Copyright 2010, John Wiley & Sons, Inc. Male and Female Differences Males usually have heavier bones Related to muscle size and strength Female pelvis is wider and shallower than male pelvis: allows for birth Aging and Skeletal System Birth through adolescence: more bone formed than lost Young adults: gain and loss about equal As levels of sex steroids decline with age: bone resorption > bone formation Bones become brittle and lose calcium Bone loss can be accelerated through wrong life style choices, e.g. alcoholism, caffeine consumption Copyright 2010, John Wiley & Sons, Inc. Terminology – Disease & disorders affecting skeletal system Osteology - scientific study of bone structure & treatment of bone disorders Bone scan - diagnostic procedure that measures bone blood flow with help of radioactive tracers Giantism - human growth disorder caused by oversecretion of human growth hormone (hGH) by the pituitary gland, e.g. due to a brain tumor; - leads to abnormally strong bone growth and tallness Paget’s disease - world-wide occuring human bone disorder; - marked by increase in osteoclast numbers and activity throughout the skeleton - characterized by excessive breakdown of bone tissue; - increased bone resorption in Paget's disease patients is met by a compensatory increase in bone formation and local bone turn-over; - clinical features involves "woven bones" which are bulky, weak and prone to bowing and fractures; - scientists suspect the role of viruses of the paramyxovirus class (measles virus?, canine distemper virus?); Rheumatoid arthritis (RA) Inflammatory bone disorder which is characterized by the progressive destruction of articular cartilage; Also excessive subchondral bone resorption due to overly active osteoclasts Active macrophages are found in the inflammatory state which accumulate in the (usually swollen) rheumatic synovial joint membrane; Macrophages and other immune cells release digestive enzymes and osteoclast stimulating cytokines (e.g. IL-1, IL-6, IL-11) Treated with anti-inflammatory drugs, such as Vioxx, Ibuprofen, Aspirin; Rheumatoid arthritis (RA) About 1% of the world's population is afflicted by rheumatoid arthritis, women three times more often than men. Onset is most frequent between the ages of 40 and 50 Copyright 2010, John Wiley & Sons, Inc. Fibrodysplasia ossificans progressiva (FOP) very rare, horrifying genetic bone disorder; affected human individuals grow a "second skeleton due to overproduction of bone and connective tissue; bone overgrowth eventually freezes mobility of the neck, spine, hips and even the jaw; patient literally becomes trapped into a "second skeleton"; strong link of FOP to mutations of Chr#2-located ACVR1 gene; Copyright 2010, John Wiley & Sons, Inc. Cleft lip and cleft palate Congenital disorder caused by incomplete fusion of left and right maxillary and palatine bones during early (week 6-8) embryonic development “Cleft lip" often associated with the formation of a cleft palate); Cleft may cause problems with feeding, ear disease, speech and socialization. Cause is not known, but toxic substances, environmental pollutants, and nutritional imbalance during embryogenesis are discussed as risk factors Copyright 2010, John Wiley & Sons, Inc. Osteoporosis Pathological reduction in bone mass and deterioration of the bone architecture after the age of 40 Resulting bone decay leads to an increase in fragility of bone and high susceptibility to bone fractures, especially hip fractures (“broken hip”); Most common cause of osteoporosis in women is the decrease in estrogen levels that accompanies menopause; leads to increased bone resorption and depletion of body calcium; "For every 10% of bone that is lost, the risk of bone fracture doubles." Copyright 2010, John Wiley & Sons, Inc. Osteoporosis Copyright 2010, John Wiley & Sons, Inc. Osteoporosis in numbers "In the U.S., an estimated 16.8 million postmenopausal women have lost more than 10% of their peak adult bone mass, another 9.4 million have lost more than 25% and 4.8 million have already suffered an osteoporotic bone fracture. 25 - 30% of all hip fractures occur in men, which show an increase in osteoporosis in recent years." Copyright 2010, John Wiley & Sons, Inc. Excessive bone loss Pathological or life style conditions that can contribute to excessive bone loss are: 1. Multiple myelomatosis 2. Hyperparathyroidism 3. Hyperthyroidism 4. Chronic glucocorticoid treatment - leads to suppression of osteoblasts - affects 30 million people in the U.S., 80% of them are women; 5. Excessive alcohol and caffeine consumption ? Copyright 2010, John Wiley & Sons, Inc. Spina bifida Congenital defect of the vertebral column Caused by failure of the neural tube to close during the first month of embryonic development (often before the mother knows she is pregnant). Characterized by laminae that failed to unite at the midline Can be surgically closed after birth, but this does not restore normal function to the mostly lumbar or sacral part of the spinal cord Risk factors: lack of folic acid, fever, diabetes, obesity Herniated discs Painful rupture of fibrocartilage of the intervertebral discs (disc ring) due to injury, weakening of ligaments or over weight; Most disc herniations occur in lower back, most often between the fourth and fifth lumbar vertebral bodies or between the fifth and the sacrum. Often emerging spinal nerves or sciatic nerve, may become painfully compressed Disc ring tear may result in release of inflammatory chemicals which may directly cause severe pain Copyright 2010, John Wiley & Sons, Inc.