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The Skeletal System: Evolution of the Axial Skeleton Skeletal System 205 bones (+/-) Consists of two main

in parts: Axial skeleton: skull, vertebral column, sternum, and ribs Appendicular skeleton: bones of the shoulder and pelvic girdles and the limbs 1

Functions of Bones 1. Protection of vital structures (e.g. ribs and skull) 2. Support for the body (e.g. vertebral column) 3. Mechanical basis for movementprovide attachments for muscles and provide levers (e.g. transverse processes) Bone Structure Calcium carbonate and calcium phosphate make up the majority of the inorganic bone Inorganic components account for 60% of bone weight in adults These salts are deposited in a matrix of collagen fibers Salt crystals give bone hardness and rigidity Collagen fibers give bone toughness like re-rod in concrete 2

FORMATION OF COLLAGEN The role of VITAMIN C The collagen molecule is unique in amino acid content -hydroxyproline and proline together make up about 1/10, glycine 1/3 of all amino acids in collagen. amino acids have C, H, O, and N amino acids are organic & have COOH and NH groups form a polypeptide called protocollagen synthesized in ribosome in fibroblast, osteoblast, and chondroblast high proportion of proline and glycine to other of the 20 amino acids (lysine, cystine, aniline, etc) gives the fiber special characteristics FORMATION OF COLLAGEN The role of VITAMIN C The enzyme collagen hydroxylase adds OH group to proline cofactors are ascorbic acid (Vit C) & the ferrous ion ascorbic is a strong reducing agent (adds OH) used as an antioxidant in foods [oxidation - loss of hydrogen electrons] so ascorbic acid supplies electrons Its a vitamin, animals cannot synthesize it 3

FORMATION OF COLLAGEN The role of VITAMIN C Collagen fibrous protein, linear molecules in long strands polypeptide chain of glycine, hydroxyproline, aniline, proline as major amino acids polypeptide chain is a condensation reaction between the -COOH (carboxyl group) and the amino group (NH2) of adjacent amino acids. It occurs in peptide bonds between acids, through the elimination of a molecule of water other fibrous proteins are myosin (muscle) and fibrin (clot) opposite proteins are globular -coiled into ball (blood proteins) strands are held together by X linkages of disulfide bond Tropocollagen fundamental unit of collagen fiber three helically -arranged polypeptide chains Types of Bone Tissue Cortical or Compact Bone Dense and hard Comprises the outer layer of bones Trabecular or Cancellous Bone Spongelike Found in the expanded ends of long bones 4

Types of Bone Tissue Classification of Bones Long Bones Femur or humerus Short Bones Bones of wrist Flat Bones Skull, ribs Irregular Bones Vertebral bodies Sesamoid Bones Patella 5

Bone Structure Long bones consist of: A diaphysis (shaft) Two epiphyses (expanded ends) that articulate with other bones Growth and Development of Bone Bones develop from embryonic connective tissue Two types Intramembranous bones: form directly in a fibrous membrane Endochondral bones: form by replacement of hyaline cartilage 6

Growth and Development of Bone Intramembranous Flat bones of the cranial vault Are membranous to allow passage through birth canal Ossification begins after birth and continues for several years Growth and Development of Bone Endochondral Form from cartilaginous bones that exist at birth Osteoblasts (bone builder cells) lay down a thin layer of bone around the diaphysis Cartilage is replaced by bone by the process of endochondral ossification 7

Cells in Bone Osteocytes Most abundant Maintain normal bone structure Recycle calcium salts in bone matrix Assist in repair Osteoclasts Bone destroyers Secrete acids and enzymes during resorption Regulates calcium and phosphorous levels in the blood Osteoblasts Bone producers (osteogenesis) Produce bone matrix and promote deposition of calcium salts in matrix Becomes an osteocyte when surrounded by calcified matrix Vertebral Elements 1) Centrum surrounds notochord 2) Dorsal arches protects neural tube 3) Ventral arches protects blood vessel Evolutionary trend: 1) Vertebral Column (VC) replaces notochord 2) vertebral elements fuse into 1 piece 8

General Vertebral Types All elements separate Fish All elements fused Amphibs, reps, birds, mammals Axial Skeleton Dorsal Arches Nerve cord Notochord Centrum (=body) Ventral arches 9

Amphicoelous - centrum hollowed at anterior and posterior end, characteristic of teleosts and early reptiles Opisthocoelous - centrum convex on anterior surface, concave on posterior surface, characteristic of some of the vertebrae of reptiles and mammals Procoelous - centrum concave on anterior surface, convex on posterior surface, characteristic of some reptiles and amphibians Acoelous - centrum flat on posterior and anterior surface, characteristic of some reptiles and mammals 10

Skeletal Evolution The internal skeleton or living scaffolding is an evolutionary new idea -- the living, growing endoskeleton avoids the limitations of size, weight and mobility imposed by the exoskeleton of ancient fishes and arthropods. The ancient fishes with dermal armor couldn't compete with critters featuring this new idea. The changing role of bone is one from protection to support. located in the center of the centra of the vertebrae matrix is intracellular rather than extracellular as with CT, cartilage, and bone surrounded by a sheath, sheath is partly ossified and combined with surrounding bone to make up vert. centrum. Cyclostomes have only a notochord Types In modern fish bony elements restrict and replace the notochord, but some notochord is never missing from modern, adult fish The vertebral centra are ossified, but still biconcave (amphicelous), whereas primitive fish have a notochord like a STRING OF BEADS. Notochord 11

Fish Two vertebral regions only, Trunk, and Caudal (tail); trunk vertebrae with ribs, neural arches and spines, caudal vertebrae with neural and hemal arches. -Centra develop and strengthen the vertebral axis 12

Regionalization of the Vertebral Column 2 types: trunk & caudal 4 types: cervical trunk sacral caudal All bear ribs 4 types: thoracolumbar (new) 5 types: thorax & lumbar lose ribs Amphibians have a single cervical vertebra (L., cervix 5 neck), called the atlas, which allows the head limited mobility . The trunk vertebrae are essentially alike. A single sacral vertebra (so called because this part of the body of domestic animals was offered in sacrifices) and pair of ribs articulate with an expanded pelvic girdle and transfer body weight to the hind legs. Weight transfer to the pectoral girdle of nearly all tetrapods is not by bone but by muscular connections between the trunk skeleton and the scapula. Caudal vertebrae follow the sacral vertebra. Reduced hemal arches are present on the caudal vertebrae that lie behind the cloaca. Ribs and zygapophyses are usually absent in caudal vertebrae. The caudal vertebrae become progressively smaller toward the tail tip, and the hemal and neural arches gradually disappear. 13

Reptiles Five vertebral regions, cervical, thoracic, lumbar, sacral and caudal. The Alligator has 8 cervical, 11 thoracic, 5 lumbar, 2 sacral (fused) and 40 caudal vertebrae. Moveable, double-headed, ribs are borne on the thoracic vertebrae. Ribs, if present on the lumbar vertebrae are fused. Snakes may have as many as 500 vertebrae. Both thoracic and lumbar regions bear ribs. The Turtle vertebral column has 8 cervical, 10 trunk, 2 sacral and 16 to 30 caudal vertebrae. The first caudal as well as all the sacral and trunk vertebrae are fused with dermal bone to form the carapace. The ribs are expanded and fused to the inner surface of the costal plates of the carapace. The ribs are single headed. in the Reptiles the two anterior cervical vertebrae are specialized. The first cervical vertebra, the Atlas, is ring-like and lacks a centrum, and articulates with the occipital condyles of the skull. The second cervical vertebra, or Axis, has an anteriorly projecting process, the odontoid process, which fits into the cavity of the Atlas, acting as a pivot in turning the head. Birds Flying and bipedal walking. Rigidity of the vertebral column is achieved by the fusion of many vertebrae. The cervical vertebrae number 13 to as many as 25, and have great flexibility, due to the heterocoelous centra. There are 5 thoracic vertebrae, but the last one is fused into the synsacrum, and the first four are fused together. The last thoracic, all the lumbar, the 2 sacral and several caudal vertebra all fuse to form one bone, the Synsacrum. This in turn is fused to the pelvic girdle. There are several free caudal vertebrae then the tail ends in an enlarged Pygostyle, which represents several fused vertebrae. The ribs bear posteriorly projecting Uncinate processes, each being ankylosed to the next posterior rib, the distal region of the ribs are joined to the sternum via sternal processes. 14

Fig. 8.17 Mammals Note the five vertebral regions - cervical, thoracic, lumbar, sacral and caudal. In the cat there are 7 cervical, 13 thoracic, 7 lumbar, 3 sacral and 4 to 26 caudal vertebrae. Only the thoracic vertebrae bear ribs. The ribs are double headed. The three sacral vertebrae fuse to form one bone, the sacrum, with which the pelvic girdle articulates. Between the vertebra are intervertebral cartilages (disks), which are composed of fibers and notochord remnants. 15

SUMMARY 1.The axial skeleton consists of the notochord, vertebral column, ribs, median fins (in fishes), and sternum (in terrestrial vertebrates). It forms the fundamental framework of the body and transfers body weight to the girdles and appendages in tetrapods. Rib movements help ventilate the lungs in most tetrapods. 2.Vertebrae develop in an intervertebral position by the condensation of mesenchyme of sclerotomal origin. The mesenchyme first chondrifies in the neural and hemal arches, and, in most species, the cartilage is later replaced by bone. The centra develop in different ways in different taxa. 3.The vertebral axis of a fish is an integral part of its locomotor apparatus. It resists compression of the body and converts the contraction of segmental longitudinal muscles into lateral undulations. Because all parts of the vertebral column have nearly the same function, there is little regional differentiation. The first vertebra articulates with the skull, and the caudal vertebrae bear hemal arches. 4.The notochord persists in the adult of living jawless craniates, lungfishes, sturgeons and paddlefishes, and some other species that do not move rapidly through the water. Contemporary sharks and their allies have cartilaginous centra that are partly calcified; advanced actinopterygians have well-ossified centra. 5.Although the vertebral axis participates in lateral undulations in some amphibians and early reptilomorphs, it is primarily a supporting beam. It must resist torsion and bending in the vertical plane so that these low-slung animals do not drag on the ground when they walk 6.Ribs were well developed in basal tetrapods and attached by two heads to most of the vertebrae. In contemporary amphibians they are very short 7.The vertebral axis of an amphibian is stronger and more complex than in a fish. A well-developed cervical vertebra (the atlas) permits movements between the head and trunk, a sacral vertebra and pair of ribs articulate the vertebral column with the pelvic girdle, and caudal vertebrae lie in the tail. 8.Evolutionary trends that began in the axial skeleton of amphibians continue in

reptiles. Intercentra are lost in most contemporary species, and the definitive centrum develops from the expanded pleurocentra. The neck is longer and more mobile, and there are more cervical vertebrae. At least two sacral vertebrae and their ribs articulate with the pelvic girdle. Tail length varies. 9.The anterior trunk ribs of living reptiles connect by flexible costal cartilages with the sternum in most species. Abdominal ribs may be present caudal to the sternum. 16

10.The structure of the axial skeleton of birds correlate with their specializations for flight and bipedal walking. The long and flexible neck allows the head and bill to be used in many manipulative functions that the specialized forelimbs can no longer do. Many trunk vertebrae are fused to form a firm fulcrum for wing action; others have united with the sacral vertebrae to form a solid synsacrum to which the pelvic girdle unites. The terminal caudal vertebrae have united to form a pygostyle, to which large tail feathers attach. 11.The sternum of birds is ossified and usually bears a large keel that increases the area for the attachment of flight muscles. 12.Mammals are active, agile vertebrates with strong but flexible axial skeletons. Most have seven cervical vertebrae, of which the first two (atlas and axis) are specialized to allow for free movement of the head. The thoracic region contains the lungs and transfers body weight to the pectoral girdle. Three or more sacral vertebrae usually unite, forming a strong sacrum. Tail length and functions vary among mammalian taxa 17