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Skeleton
mineralized connective tissue (mostly bone; also dentin, cartilage and enamel/enameloid substances) plus ligaments, tendons and bursa
Bones
matrix of collagen fibers impregnated with hydroxyapatite crystals (calcium, phostphate and hydroxyl ions) from osteoblasts
Water and mucopolysaccharides
cementing subtance which binds the crystals to collagen
Osteon or haversian systems
concentric lamellae around a haversian canal containing an arteriole, venule, lymphatic and nerve fibers
Periosteum
dense fibrous membrane covering all bones except at articular surfaces
Spongy or cancellous bone
trabeculae (tissues) and marrow (reticulum of connective tissue with blood vessels and nerves; adipose and hemopoitietic tissue) interiorly lined with endosteum (thin connective membrane)
Dentin (teeth)
same constituents as dense and spongy bone, forms only in outer layer of dermis and is often coated with enamel
Dentinal tubules
odontoblasts retreat suring its formation and trail protoplasmic processes in canaliculi
Dentin prevalence
found in dermis of early verts, now only present in scales of ganoid/elasmobranch fish and in teeth
Acellular bone
no canaliculi or cell proccesses left after osteoblasts retreat (fibrous sheets of modern fish scales, cementum of vert teeth)
Blastema
aggregation of mesenchyme stimulated to differentiate into another tissue (bone, muscle, cartilage, etc.)
Membrane bone
deposited directly within membranous blastema without cartilaginous stage; no haversian canals
Dermal bone
membrane bone derived from skin dermis
Replacement bone
cartilage formed in advance of ossification and replaced with bone behind= bone growth
Endochondral ossification
deposited where hyaline cartilage already exists
Long bone
begins in the middle of shaft (diaphysis) and grows to both ends (epiphyses); ossification centers also soon appear in each epiphysis (epiphysial plates)
Epiphyses
usually ossify (stop growing) at sexual maturity
Cartilage
consists of sulfated mucopolysaccharides (from chondroblasts) deposited on preexisting collagenous matrix
Hyaline cartilage
least differentiated, precursor of replacement bone, remains only on articular surfaces
Fibrocartilage
thick, dense collagen bundles (intervetebral disks)
Elastic cartilage
contains additional elastic fibers (pinna of ear)
Calcified cartilage
deposition of calcium salts (jaws of sharks)
Homeostasis
hormonally controlled deposition/withdrawal of calcium and other mineral salts from bone in response to dietary intake and cellular demands
Resorption/replacement of bone and cartilage
leads to remodeling of growth demands and used to cope with mechanical stress
Tendons
connect muscles with bone
Ligaments
conect bone to bone
Aponeuroses
flat and very wide tendons and ligaments
Sesamoid cartilages or bones
mineralized tendons and ligaments
Arthrosis
meeting of two bones (joints)
Diarthrosis
freely movable, articular surfaces covered by hyaline cartilage, enclosed in fibrous capsule lined by synovial membrane that secretes a lubricatory fluid (elbow, knee)
Amphiarthrosis
limited movement, unitied by resilient fibrocartilage, fibrous joint capsule without synovial membrane (between vertebral centra)
Synarthrosis
immovable (skull) due to jagged seam (suture) which sometimes is obliterated during development (ankylosis)
Symphysis
bilateral bones immovably joined in body midline by pad or fibrocartilage (pubis)
Mineralized tissues and the invertebrates
matrix is collagen, crystals usually calcium carbonate
Regional components of the skeleton
axial and appendicular skeleton
Axial skeleton
notochord and vetebral column, ribs and sternum, skull and visceral skeleton
Appendicular skeleton
pectoral and pelvic girdles, skeleton of paired fins and limbs, skeleton of median fins of fishes
Heterotropic bones
develop in amniotes in areas of continual stress (human kneecap, steer os cordis, rodent baculum)
Vertebral column (fish)
intervertebral articulations permit only side to side movement
Vertebral column (land invasion of tetrapods)
development of dorsalventral flexibility, some side to side limitation, regional specialization along the column
Modern vertebrae
centrum (notochord in early otogeny), neural arch (snclosing spinal chord), hemal arch or chevron bones (amniotes) in tail (enclosing caudal artery and vein), apophyses (processes) from arches or centrum
Transverse processes (diapophyses)
articulate with ribs, some muscles attach (extend or flex vertebral column)
Pre and postzygapophyses
interlock and limit dorsoventral flexion
Parapophyses
lateral from centra (few tetrapods), articulate with capitulum of bicipital rib (unusual)
Hypapophyses
midventral (snakes)
Morphogenesis of vertebrae
central are developmentally intersegmental relative to the somites and myomeres except in the tail of some fishes and primative tetrapods (two centra per segment, diplospondyly)
The vertebral column of fishes
very divers between species, morphological specializations within a species: dorsal (trunk) and caudal (tail)
Shark vertebral column
notochord through vert column and restricted within centra; centra restricted at each end (amphicelous); vertebral canal (for spinal chord) = paired dorsal plates, dorsal intercalary plates and sometimes supradorsal cartilages; hemal arches = paired ventral plates and sometimes ventral intercalary plates
Fibroelastic ligament
overlies/connects neural spines in ALL fishes
Teleosts vertebral column
well ossified amphicelous vertebrae with remnant of notochord; connected by complex of collagenous and elastic ligaments (facilitate lateral undulation); neural spines often tall; variets of processes from arches and centra (unlike in tetrapods)
Evolution of terapod vertebrae
early tetrapods had amphicelous rachitomous (several bones per segment): hypocentrum, pair of pleurocentra, and a neural arch
Hypocentrum
u-shaped, anterior, cradled notochord
Change to modern amniotes (vertebrae evo)
increased pleurocentra and reduced hypocentra
Change from amphicelous
loss of concavity at one or both ends of centrum
Procelous (anurans, modern novian reptiles)
posterior convex buildup/coalescence of intercentrum (chordal cartilage)
Opisthocelous (salamanders)
anterior convexity
Acelous (mammals)
no concavity, independent fibrocartilaginous intervertebral disc with remnant of notochord (pulpy nucleus)
Reginal specialization in tetrapod columns
Sacral, cervical, thoracic, and lumbar
Sacral
to accept force transmitted through the pelvic girdle
Cervical
for increased mobility of head (more mobile joint with skull, shortened or eliminated ribs, more mobile intervertebral joints (neck)
Thoracic
long ribs restricted to region housing viscera (bony cage, assist external respiration)
Lumbar
between thoracic and sacral
Neck vertebrae (amphibians)
1 → 2 occipital condyles, first cervical vert lacks processes; leads to a dorsoventral rocking of the skull
Neck vertebrae (amniotes)
more cervical vert, first two modified for greater skull movement
Atlas
missing centrum (forms odontoid precess of axis), two superior articular facets meet with occipital condyles (1: reptile, 2: mammals) for ‘yes’, skull and atlas pivot on odontoid process (held in place by transverse atlantal ligament) for ‘no’ (no zygapophyses on atlas)
Heterocelous (birds)
caudal end of centra saddle shaped and accommodated by cephalic end of next vertebra; allows ‘yes’ and ‘no’ movements
Stabilizing the hind limbs
sacrum and synsacrum
Sacrum
ankylosed sacral vertebrae of mammals
Synsacrum (modern birds and armadillos)
ankylosed last thoracic vertebrae, all lumbars, sacrals, some caudals and their ribs, plus the pelvic girdle
Tail vertebrae
urostyle, pygostyle and coccyx
Urostyle (anurans)
from elongated perichordal cartilage at base of larval tail, grows and ossifies after loss of tail at metamorphosis
Pygostyle (birds)
fused caudal vertebrae at end of tail
Coccyx (apes, humans)
fused vestigial caidals
Ribs
formed intersegmentally like the centra vertebrae (by scleroblasts from two successive mesodermal somites)
Fish ribs
basal actinopterigian (Polypterus) and some teleosts have both dorsal ribs (between epaxial and hypaxial muscles) and ventral ribs (develop in myosepta, reach down just external to parietal peritoneum); most fish only have ventral ribs, some (sharks) only have dorsal ribs, some (skates) have none
Tetrapod ribs
most are bicipital with a dorsal head (tuberculum) and a ventral head (capitulum)
Tuberculum
articulates with transverse process
Capitulum
articulation site varies
Two parts of thoracic ribs in amniotes
Costal (adjacent to vertebra), sternal (more ventral, at least some articulate with sternum, may remain cartilaginous as in humans, then called costal cartilages)
Amphibian ribs (anurans and urodeles)
short and ankylosed to transverse processes in anurans
Amphibian ribs (apodans)
long thoughout vertebral column and important for locomotion
Nonavian reptile ribs
'conventional’ in lizards and crocodiles
Nonavian reptile ribs (turtles)
no cervical, those trunk vertebrae fused with costal plates of carapace
Nonavian reptile ribs (snakes)
no sternum, ligamentous connections with scutes (locomotion)
Bird ribs
functional ribs of thoracic basket are thin, flat, and bear uncinate processes; lightweight and sturdy structure for attachment of powerful flight muscles
Mammal ribs
recognizably confined thorax; ‘floating rib’ → costal cartilages (sternal ribs) fail to reach the sternum
Tetrapod sternum
serves as anteroventral site against which pectoral girdles of terrestrial vertebrates can be braced and pectoral flight muscles can be attached
Amphibian sternum
well differentiated only in anurans, consist of several linear segments or cartilage or replacement bone
Reptile sternum
well developed midventral endoskeletal plate (other than in turtles)
Flighted bird sternum
enormous keel or carina for muscle attachment
Terrestrial mammal sternum
Series of bony stenebrae with last segment bearing bony or cartilaginous xiphoid process
Neurocranium
part of skull that 1. protects brain and some sense organs, 2. arises as cartilage (from several centers that expand and unite), 3. usually replaced to some extent by bone
Parachordal
parallel anterior notochord (beneath mid- and hindbrain) → basal plate
Prechordal
anterior or notochord (beneath forebrain → ethmoid plate
Sense capsules
Olfactory, otic, and optic
Olfactory capsule
partially surrounds olfactory epithelium
Otic capsule
completely surrounds otocyst → inner ear
Optic capsule
forms around retina → sclerotic coat of eyeball (not connected to neurocranium)
Unions
ethmoid plate with olfactory capsules, basal plate with otic capsules, both plates complete the floor under the brain
Living agnathans neurocrania
embryonic components remain mostly independent throughout life