Comparative Vertebrate Anatomy Lecture Chapter 7: Mineralized Tissue An Intro to the Skeleton The skeleton is composed of mineralize connective

ve tissue and of ligaments, tendons, and bursae. Bone: mineralized tissue for most part. But there is also dentin, cartilage, and enamel or enameloid substances. Osteoblasts: produce bones Odontoblasts: produce dentin Chondroblasts: produce cartilage Ameloblasts: produce enamels. These spxalized cells arise from less-differentiated scleroblasts that arise from the mesenchyme. Preliminary step in formation of skeletal tissues is the synthesis of collagen by fibroblasts

Canal + lamellae = osteon/haversian system. Blood vessels: configures haversian system and also branches and rebranches causing the system to do likewise.

Continuous with those in the bone marrow.

Shaft of long bones: haversian systems are more or less parallel to the long axis of the bone. (minimizes the likelihood of fracture under normal external stress) The lamellae of periosteal bone are formed by osteoblasts on the inner surface of the periosteum: dense fibrous membrane that covers all bones except at their articular surfaces. Haversian system: characteristic of amniotes Most amphi, a few reptiles, and some small insectivores and rodents lack them.

Bone

Collagen: proteinaceous fibril demonstable only by high power electron microscopy. Fibrils aggregate to form collagen fibers that are visible with light microscopy. The fibers form dense collagen bundles that are woven into a compact network of dense conn tissue (in dermis, tendons, and ligaments) Minerals are deposited in the network to form cartilage and bone

Spongy Bone Consists of bony trabeculae and bone marrow

Trabeculae: assemblage of beams, bars, and rods that form a rigid framework that provides maximum strength at areas of stress.

Composed of irregularly arranged without haversian systems.

lamellae

Marrow occupies the cavities between trabeculae. Consists of a reticulum of connective tissue fibers that support bV, nerve fibers, and adipose tissue (yellow marrow) Hemopoietic tissue (red marrow) in some bones produces RBC and some types of WBC. Endosteum: a thin connective tissue membrane lining the marrow cavities. Has many features of a periosteum and has the capacity to deposit bone or remodel it

Bone tissue is composed of a matrix of collagenous fibers.

Spaces between them are impregnated with hydroxyapatite crystals: composed of calcium, phosphate and hydroxyl ions Deposited under the influence of osteoblasts.

Cementing substance (composed of water and mucopolysaccharide): binds the crystals to the collagenous matrix.. Osteoblasts become trapped by the bone that they have laid around them. Osteocytes occupy tiny fluid-filled pools (lacunae) of interstitial fluid. Canaliculi: fluid-filled canal interconnecting the lacunae. Houses protoplasmic processes extending from the osteocytes. Fluid in lacunae and canaliculi contains calcium and phosphate ions which, depending on the serum calcium level, are being: deposited on the collagenous matrix or withdrawn from it

Flat bones: characterized by a core of spongy bone and marrow sandwiched between two layers of compact surface Ex. Ribs, scapulae and membrane bones of skull.

Dentin Has the same constituents as compact and spongy bone but odontoblasts are not trapped in lacunae during osteogenesis.

They retreat as they deposit dentin they are at its inner border. They leave behind protoplasmic processes in canaliculi

Compact Bone

consists of lamellae: collagenous bundles

layers

of

mineralized

Dentinal tubules: the canaliculi in dentin/ Extend all the way to the surface of the dentin. Forms in the outer layer of the dermis, just beneath the epidermis Frequently coated on its surface by enamel or enameloid. Enameloid of placoid scales is produced by odontoblasts very hard dentin. Dentis is found today only in scales of basal rayfinned and elasmobranch fishes and in teeth.

lamellae are concentric around a haversian canal. Contain an arteriole, a venule, a lymph vessel and nerve fibers.

Acellular Bone

impregnation of a collagenous hydroxyapatite crystals.

matrix

with

Osteoblasts do not only retreat as they deposit bone but also leave behind no processes or canaliculi. Acellular bone (aspidin): thin layer Constitutes the fibrous platelets of the flexible scales of modern fishes and the cementum of verteb teeth.

Difference is that in endochondral cargilage must be removed before bone can be deposited. Immediate result for both cases = formation of temporary spongy bone. The latter, in turn, is eroded and replaced by compact bone, spongy bone, or a marrow cavity depending on its location.

Term aspidin was first applied to acellular dermal bone found in fossil heterostracens Presence in teleosts is due to convergence.

Membrane Bone and Replacement Bone Before bone or cartilage can be deposited, a preskeletal blastema must develop

Blastema: any aggregation of mesenchyme that differentiates into some tissue such as muscles, cartilages and bones.

Once preskeletal blastema is formed some of mesenchyme cells become fibroblasts and secrete collagen other become either osteoblasts or chondroblasts and secret enzymes essential for formation of bone or cartilage.

Process of endochondral ossification of a typical long bone of a tetrapod limb commences midway in the shaft (diaphysis) of a miniature cartilaginous model of the bone-to-be and proceeds toward the two ends or epiphyses. When diaphyseal center becomes active, one or more endochondral ossification centers appear in each epiphysis. Additional cartilage continues to be deposited in advance of all zones of ossifcaiton, and rate of deposit equals rate of ossification = shaft continues to grow in length while being ossified.

Membrane Bone

bone deposited directly within a membranous blastema without having been preceded by a cartilaginous model intramembranous ossification gives rise: To certain bones of the lower jaw, skull, and pectoral girdles To dentin and other bone that forms dermis of the skin. To vertebrae in teleosts, urodeles, apodans only To a few miscellaneous bones. May be compact, spongy and lamellar or nonlamellar. It lacks haversian canals because of the arrangement of the bV that participates in its deposition. Dermal bone: any bone derived ontogenetically or phylogenetically from the dermis of the skin. Term specifies history and not its histological features. It is membrane bone Many, though not all, membrane bones are of dermal origin bones that constitute the dermatocranium, the bones that invest the Meckels cartilage, the membrane bones of the pelvic girdles phylogenetic derivatives of the integument. Not much bone remains in dermis of recent verteb but that which does, the bony plates and scales of early verteb, arises in situ.

As indiv grows older deposition of cartilage gradually slows eventually leaving only a narrow cartilaginous epiphyseal plate between the diaphysis and each epiphysis. Chondrogenesis continues within this plate fo a time and bone continues to elongate as epiphyseal plate are elaborating cartilage.

Birds and mammals epiphyseal centers cease elaborating cartilage after indiv attains sexual maturity the epi plates ossify, epi become sutured to the shaft and bone stops growing. Bones of other shape exhibit other combi of ossification centers. Growth in diameters of the bones take place when bone is deposited from the inner surface of the periosteum. While endochondral ossification is going on w/in a bone, periosteal bone is being deposited at the surface. As a result of this combines with continual resorption and remodeling bone reaches adult shape, size and proportions. Cart fishes either no longer have the genetic codes necessary for endochondral ossification or there is some factor that prevents gene expression for ossification.

Ectotherms this is throughout life.

Cartilage Cartilage resembles bone in that it is formed within a matrix of collagenous connective tissue and the cells lie in lacunae.

Replacement Bone Deposited where hyaline cartilage already exists. Existing cartilage undergoes degenerative changes and disappears. Process of endochondral and intramembranous ossification are the same in that both consist of

Intercellular matrix contains sulfated mucopolysaccharide Cartilage has no canaliculi demonstrable by light microscopy No blood vessels of its own.

Chrondrocytes: lacunar cells that are supplied with oxygen and nutrients diffused by the nearest capillaries adjacent to the cartilage. Cartilage formation is initiates when chondroblasts deposit their mucopolysaccharide on a preexisting collagen matrix.

Mesenchyme organizes a perichondrium (a bounding membrane) Additional cartilage is laid down within the blastema by the mesenchyme from the perichondrium and by fibroblasts and chondroblasts that have arisen by mitosis from parent cells within the developing mass. Chondroblasts become chondrocytes after being trapped in lacunae. Perichondium is constantly reconstructed as the mass enlarges and changes shape.

and replacement of these in a new alignment to provide a wider, higher skull with a cavity large enough to accommodate the growing brain. Cartilage and bones elsewhere are also subject to remodeling. Remodeling of bones also occurs in response to mechanical stress resulting from continuous use of inserting muscles or from weight bearing. Bone becomes thicker at sites where stress is high, then when stress is withheld. The arrangement of latticelike components of spongy bone bears a striking resemblance to lines of force generated by bones natural load.

Tendons, Ligaments, and Joints Tendons and ligaments consist chiefly of closely packed bundles of collagen.

Hyaline Cartilage: least differentiated variety and is the precursor of replacement bone Translucent because all components have the same refractive index to lightl Little hyaline cartilage remains after the growth of replacement bone has stopped. Then, hyaline cartilage are found chiefly on the articular sufaces of bones w/in joints of tetrapods. Most any other hyaline cartilage that forms is likely to be transformed into:

Tendons: connect muscles with bones and have a shiny appearance. Collagen bundles are so arranged as to offer a maximal resistance to the tensions created when a muscle contracts.

Where tendon is attached to muscle, collagenous bundles of tendon are directly continuous with those of the fibrous muscle sheath (epimysium) and where it attaches to cartilade or bone, they are continuous with the collagenous bundles of the perichondrium or periosteum.

Fibrocartilage: cart with exceptionally thick, dense collagenous bundles in the interstitial matrix interverteb discs of mammals Elastic cartilage: addiction to collagenous fibers are elastic fibers pinna of ear, walls of outer ear canal , epiglottis calcified cartilage: formed when calcium deposits are within the interstitial substance of hyaline cartilage or fibrocartilage mistaken for bone.

Ligaments: connect bone to bone. Arrangement of collagenous bundles is less regular but are directly continuous with those of the periosteum.

Aponeuroses: tendons and ligaments that are flat and wide.

The gala aponeurotica scalp.

mammalian

Skeletal Remodeling bones not only provide mechanical support for the verteb body but also constitute impt storage places for calcium and other mineral salts.

may also be used to refer to fibrous mesenteries or cords that support visceral organs or hold them n place. In some species, ligaments and tendons become mineralized. Sesamoid cartilages/bones: mineralized nodules in tendons or ligaments patella. Arthrosis: a joint; site where two bones/cart meet. The joint is a diarthrosis If freely movable in one or more planes and the articular surface are covered with hyaline cart If the joint enclosed in a fibrous capsule lined by a synovial membrane that secretes lubricatory fluid. Ligaments hold the bones in their proper alignment in the joint. Artic between upper and lower jaws od early bone fishes Same joints and hinge joint in elbow and knee of mammals. Amphiarthrosis: permits limited movement. Resilient fibrocartilage unites the components of the joint

Calcium: constantly being deposited and withdrawn in response to dietary intake and cellular demands.

When serum calcium are rising: excess calcium is deposited along with inorganic phosphate as hydroxyapatite crystals. When it is falling: calcium is withdrawn from skeleton and other depots thereby maintaining normal serum calcium level. Withdrawal regulated by parathyroid hormone and calcitonin. Deposit may not be under endocrine control.

Cartilage and bone are constantly being resorbed and replaced in another process skeletal remodeling the entire skull must be continuously remodeled by constant resorption of existing cartilage and bone

Fibrous joint capsules keep the bones properly aligned. There may or may not be a cavity No synovial membrane. Joints between centra of mammalian verteb

Synarthrosis: sutured joint Suture: irregular jagged seam at the junction of two bones that renders the joint immovable. Joints in roof of mammalian skull. Ankylosis: if suture between two bones become obliterated during development skull of birds, premaxilla and maxilla of human embryo Symphysis: joint in the midline of the body in which bilateral bones are separated by a pad of fibrocartilage and movement is restricted or impossible.

the gizzard of some doves tongue of atleast one species of bats gular pouch of SA lizard muscular diaphragm of camels syrinx of some birds, upper eyelids of crocodilians. Similar plate of fibrous tissue, tarsal plate of eyelid develops in human beings. Rostral bone develops in snout of swine and is used in routing soil. Cloacal bone develops in the ventral wall of cloaca of some lizards. All these including sesamoid bones are incidental bones located at foci of stress and lacking phylogenetical significance.

Pubic symphis of pregnant female mammal: becomes movable thru hormoneinitiated dissolution of the fibrocartilgae before onset of labor.

Mineralized Tissues and the Invertebrates Mineralized tissues are not limited to verteb. 2/3 of living species of animals that contain mineralized tissues are inverteb matrixz is collagen and it is as old as sponges but inorganic crystals are more often calcium carbonate than calcium phosphate. Cart is found among inverteb though it appears to be lacking in the outgroups of craniates (echinoderms and protochordates) Cart is present in hagfishes but bone dentin and enamel are restricted to verteb, Distribution suggests that cartilage might have preceded bone as a structural tissue in craniates. Regional Components of the Skeleton Skeleton may be divided regionally into the ff componenrts w/c include assoc ligaments and tendons: Axial Skeleton Notochord and verteb column Ribs and Sternum Skull and Visceral Skeleton Appendicular Skeleton Pectoral and Pelvic girdles. Skeleton of paired fins and limbs, Skeleton of median fins and limbs. Heterotrophic Bones

Comparative Vertebrate Anatomy Lec Chapter 8: Vertebrae, Ribs, and Sterna Major Components of Axial Skeleton: - Vertebral column - Skull - Ribs - Sterna - Ligaments Vertebral Column Keystone of the vertebrate skeleton Forms around, and sometimes invades, the embryonic notochord during ontogenesis A segmented, more or less flexible, arched rod flanked by axial musculature, To which the head is attached, From which the rest of the body is suspended in fishes From which the trunk is suspended between anterior and posterior limbs in tetrapods.

Heterotrophic bones: develop by endochondral or intramembranous ossification in areas subject to continual stress in amniotes.

Os cordis: interventricular septum of the heard of deer and bovines. Bacalum/os penis: septum bet spongy bodies of penis of dogs, basal primates, and other mammals. Os clitoridis: in many female mammals.

Forms in:

Provides a protective bony tunnel for the spinal cord, An essential architectural component of the axial locomotor apparatus in verteb other than agnathans.

Centra of urodeles and apodans: consist of perichordal membrane bone, chordal cartilage, and within the core, unaltered notochordal tissue Centra of anurans consist entirely of replacement bone deposited in a perichordal blastema; notochord disappears In forming centra, scleroblasts from the caudal half of one somite and the cephalic half of the next somite stream to an intersegmental location around the notochord to establish the perichordal blastema.

Central, Arches and Processes

Centra: occupy the position occupied early in ontogeny by the notochord. Neural Arches: perched on the centra; successive arches and their interconnecting ligaments enclose a long vertebral canal (neural canal) Neural canal: a long vertebral canal occupied by the spinal cord Hemal arches: also known as chevron bones i

Centra intersegmental Myomeres segmental

Inverted beneath the centra of the tail and houses the caudal artery and vein, consists of paired ventral plates

Transverse process (diapophyses): most common; Articulate with ribs that project laterad into the horizontal skeletogenous septum separating the epaxial and hypaxial muscles of the body wall. Serves as attachment for some of the muscles that extend or flex the verteb column. Zygapophyses: paired processes at the cephalic end of the trunk vertebrae (prezygaphpyses) and at their caudal end (postzygapophyses)

Diplospondyly two centra per segment chiefly in tetrapods may provide the tail with increased flexibility for locomotion generally, in fishes, flexibility of the vertebral column is facilitated by elasticity of the interverteb ligaments rather than by interverteb joints, as in tetrapods.

Vertebral column of fishes fish verteb are highly diverse

Exhibits only two major morphologic regions specialization trunk (dorsals) and tail (caudals) Living agnathans Strange vertebral column

of

Articular facets of prezyga face dorsad and articulate with the ventrad facing facets of postzyga interlocking arrangement limits dorsoventral flexion of the column in the region of the trunk. Tetrapods do not possess zygapophyses

the only skeletal elements are lateral neural cartilages, one or two pairs per body segment depending on the species Hagfishes lateral neural cartilages are limited to the tail Lateral neural cartilage fuse to form a single longitudinal dorsolateral cartilaginous plate perforated by foramina for spinal nerves May be vestigial vertebrae or primitive vertebrae or they may bear no phylogenetic relationship to vertebrae Sharks Notochord present throughout the length of adult vertebral column and is constricted within each centrum

Parapophyses: lateral projections from the centra of a few tetrapods, When present they serve as the articulation site for the bicipital rib Hypapophyses: prominent midventral projections from centra of snakes, Site for attachment for certain muscles and tendons

Morphogenesis of vertebrates Typical verteb arises from the mesenchyme cells that stream out of the sclerotomes of mesodermal somites, surround the notochord and neural tube and produce the blastema for a future vertebra. Chondroblasts: deposit a cartilaginous centrum and neural arch, and in the tail, a hemal arch. Result is a cartilaginous vertebra with the notochord constricted within each centrum. Later, except in cart fishes, the cartilage is generally removed and bone is deposited. Remnants of notochord may remain in the centrum to varying degrees depending on the species.

centra are amphicelous or concave at both ends and composed of chordal and perchordal cartilage. Vertebral canal consists of paired dorsal plates that constitute ta neural arch paired dorsal intercalary plates between the arches, in some species, paired wedge shaped supradorsal cartilage Squalus acanthias lacks supradorsals; their neural arch and intercalary plates are perforated by foramina for the dorsal and ventral roots of spinal nerves and for bV. hemal arches consists of paired ventral plates. Ventral intercalary plates: common between hemal arches.

Cartilaginous fishes: cartilaginous vertebrae are never replaced by bone Teleosts, urodeles and apodans: membrane bone rather than cartilage is deposited in the perichordal blastema but the arches arise from the cartilage Fishes and amphibians except anurans: not only is cartilage or membrane bone deposited around the notochord (perichordal cartilage/bone) but chondroblasts invade the notochord sheath and deposit cartilage in the sheath and sometimes in the notochord itself Chordal cartilage: found in the centra may either calcify or ossify

Fibroelastic ligament present in all fishes, connects the neural spines along the entire length of the vertebral column A few fishes have exceptional columns.

Dipnoans, Chondrostei and the coelacanth Latimeria develop no centra; notochord present and unconstricted;thick fibrous sheath contains little cartilage or bone. Associated with the notochord in each body segment are paired: Basidorsal

Basiventral Interdorsal interventral cartilages products of separate chondrification centers in the embryonic perichordal blastema

A modern adult verteb is a composite structure.

Holocephalans its vertebral column consists of thick rings of calcified cartilage that had been deposited in the notochord sheath; rings more numerous than body segments

Vertebrae of urodeles and apodans lack evidence of pleurocentra or hypocentra; phylogenetic history are debated Vertebrae are amphicelous Centra are concave at each end Vestiges of notochord persist in the interverteb joints.

Teleosts have well ossified amphicelous vertebrae. Core of each centrum is a dumbbell-shaped vacuole where the notochord previously existed Space between the successive verteb is occupied by a porous cartilagelike material that probably includes notochordal remnants. Centra and neural arches are interconnected by a complex system of collagenous and elastic ligaments Post trunk and tail: ligaments are frequently ossified to form long, delicate, bony rods.

In most other tetrapods, concavity has disappeared anteriorly, posteriorly, or at both ends ophistocelous, procelous, acelous.

Vertebrae of Anurans and modern non-avians procelous

Procelous evolved from the buildup of a hypocentrum of chordal cartilage between two centra and the eventual coalescence of the intercentrum with the centrum ahead of it. Salamanders

Neural spines are often very tall, and they are sometimes surmounted by supraneural bones.

Opisthocelous coalescence of the hypocentrum with the centrum immediately behind it

Elasmobranchs and numerous bony fishes have two centra and two sets of neural and hemal arches in each metamere of the tail Number of centra in those regions is double the number of myomeres and spinal nerves, a diplospondylous condition.

Acelous vertebrae of mammals have concavity at neither end.

Mammals have fibrocartilaginous intervertebral disc between successive vertebrae. Pulpy nucleus notochord. remnants of the embryonic

Amia- has two centra per body segment in the postanal portion of the vertebral column but only one bears arches The other is termed intercentrum though it may not be homologus with hypocentra of other fishes or tetrapods. Embryonic basidorsal and basiventral cartilages are incorporated in the centra, and the interdorsal and interventrals are incorporated in the intercentra. No movement possible n fishes between 1st verteb and skull. They are united by cartilage Caudal-most verteb are part of the skeleton and caudal fin.

Intervertebral joints permits more flexibility.

Regional Specializations in Tetrapod Columns Sacral vertebrae pelvic girdle to the hindmost trunk vertebrae Survival on land with its irreg contours and scattered obstructions to visions was facilitated in tetrapods by increased mobility of the head with its special sense organs, including eyes that can scan the horizon Small degree in amphibians devt of a somewhat more mobile joint between the first verteb and skull.

Innovation improved in amniotes shortening or eliminating ribs on some or of the more ant verteb and by increasing mobility of interverteb joints in that region neck Cervical vertebrae intervertebral joints in the neck region

Evolution of Tetrapod Vertebrae Columns of some rhipidistian fishes and early labyrinthodonts consisted of several bones per segments

Hypocentrum is a large median U-shaped anterior bone that cradled the notochord Pleurocentra small wedges of bone overlying the notochord dorsolaterally An independent left and right laminae of bone lateral to the spinal cord, that collecteively provided a neural arch. Each lamina rested in a notch between the pleurocentrum and the hypocentrum. Notochord was continuous throughout the length of the body but constricted at the level of each hypocentrum. Rachitomous consists of several pieces Later tetrapod vertebra except urodeles and apodans appear to be modifications of the rachitomous condition; changes characterized by increased prominence of pleurocentra and concominant reduction of hypocentra today, each centrum beings ossification at several paired and median loci in the perichondral blastema.

Thoracic vertebrae- houses the thoracic viscera and participates in external respiration Lumbar Vertebrae remaining vertebrae anterior to the sacral. Snakes-400 or more vertebrae Apodans 250 or more Urodeles 100 Awkward leaping anurans have the shortest columns and limited flexibility Turtles and birds cervical and caudal segments are flexible. Most trunk verteb are rigidly fused to the synsacrum in birds and to the carapace in turtles.

The Craniovertebral Junction and Neck Vertebrae

Modern Amphibians first and only cervical vertebrae lack processes, cranial end bears two smooth concave facets that articulate with the two occipital condyles

Absence of processes and nature of amphibian cranioverteb joint permites limited dorsoventral rocking of skull Two-facet artic is a change from earliest amphibians.

Amniotes have larger number of cervical vertebrae which provides them a long flexible neck. First 2 verteb are modified in such a fashion to permit more independent movement of the head. Atlas first vertebra and is ringlike bec. its centrum is missing

Superior articular facets found in cephalic end of atlas one or two deep cavities for articulation with the single occipital condyle of reptiles or with the two of mammals. Condyloid joints. Except in snakes, centrum of atlas has become the odontoid process of the axis (2nd verteb)

Last thoracic verteb + all lumbars + all sacrals + first few caudals + ribs = synsacrum w/c fuses with the pelvic girdle A compact pelvis is formed that provides a rigid brace for the stance of birds. The thoracic verteb anterior to synsacrum also unite more or less completely so there is little flexibility in the avian backbone = minimizes number of muscles needed to keep streamlined body during flight = reducing cost in terms of energy expended. Archeopteryx Trunk verteb not fused Pelvic girdle was proportionally smaller Weight when standing was counterbalanced to a considerable degree by long tail. Armadillos also have synsacrum.

Tail Vertebrae: Urostyle, Pygostyle, Coccyx Caudal verteb in early tetrapods probably numbered more than 50

Process projects forward to rest on the floor of the atlas where in mammals, it is held into place by a transverse atlantal ligament

In modern number is more reduced and highly variable Toward end of tail arches and processes become shorter and rudimentary until vertebrae consist solely of small cylindrical centra. Anurans

Proatlas resembling a neural arch and derived from a pair of bilateral cartilaginous blastemas first appeared in basal amniotes and was present in synapsid ancestor of mammals. Flexible neck is a valuable asset gathering food, avoiding enemies, extends horizon, increases number of degrees in its arc. Flexibility of neck is exceptional in birds because of the nature of articulations of their verteb. caudal ends of centra are saddleshapes having a convexity in the RL axis and a concavity in the dorsoventral axis.

Urostyle: found at the end of vertebral column. Develops from a continuous elongated perichondral cartilage at the base of the larval tail, and it grows and ossifies after tail is lost Composed of postsacral verteb Reptiles (lizards) Many lizards, when captured by the tail break off to the end distal to the site of capture and scurry away. Then the tail regenerates

Heterocelous enable both lateral and dorsoventral flexion of the neck. Turtles also have a uniquely flexible neck. Ball-and-socket joints between successive procelous centra enable the entire head and neck to be completely retracted into the shell by flexing the neck into several folds dorsoventrally or sidewise. Turtles have only 8 cervical verteb. Remainder of the column of birds and turtles, except a short portion of the caudal verteb, is totally rigid. Mammals almost always have 7 cervical verteb

Autonomy: implemented by a zone of soft tissue that divides each tail vertebra into cephalic and caudal sections, the location being at level of myoseptum. Archeopteryx Had a long tail Modern birds still have a tail but not prominent.

Exception: three edentates (two sloths and Great Anteater six, eight, or 9) manatees (6) marsupial moles, armadillos andcetaceans (shortened an more or less fused together) Length of the centra determines neck length in mammals.

Pigeons have 15 caudal verteb 5 are ankylosed with the synsacrum, six are independent, last four are fused to form the pygostyle: skeleton of the visible part of the tail, develops as four separate centra. Mammals 3-50 caudal verteb. Tail of sperm whales 24 Apes and humans 4 or 5 vestigial caudal verteb Caudal verteb lack arches but have rudimentary transverse proceses.

Stabilizing the Hind Limbs Sacrum and Synsacrum Sacral verteb bear stout transverse processes that are strong enough. Ankylosed to the distal ends of the transverse processes may be equally stout ribs. Amphi have one sacral verteb Reptiles including most birds and opossums have 2 Mammals have 3-5 Some perissodactyls- up to 8 Edentates- 13

Last three or four diminish progressively in size and in humans, they fuse with another to form coccyx (25y/o) Centra of coccyx is still unidentifiable but last one is a mere nodule of bone. Adult rhesus monkeys have a prehensile tail the length of w/c averages nearly 50% of monkeys sitting height.

Sacral verteb of mammals usually ankylose to form a single bony complex, the sacrum Sacral verteb do not usually differentiate in tetrapods that lack hind limbs. Modern birds.

Ribs Ribs articulate with verteb and extend into the body wall. The rib is formed intersegmentally by scleroblasts from two successive mesodermal somites in the same manner as centra.

Ruijin et al: confirmed a sclerotomal origin for the entire rib but have noted a necessary interaction between sclerotome and myotome for the successful development of a rib.

Most bony ribs are of endochondral origin. Gastralia: so-called abdominal ribs of some reptiles are not ribs, but riblike membrane bones that may be remnants of ancestral bony dermal exoskeleton. Fishes

Basal actinoptrygian Polypterus and some teleosts have two set of ribs, dorsal and ventral, assoc with each trunk verteb. Dorsal Ribs: pass laterad intot he horizontal skeletogenous septum that separates the epaxial and hypaxial muscles.

Ventral ribs: develop in myosepta and arch ventrad in the lateral body wall just external to the parietal peritoneum as in the mammalian thorax. each pair of ventral ribs, or their basal stumps, approach one another beneath the Centrum & unite to form the hemal arches of the tail. Most fishes have only ventral ribs. Sharks and few others have only dorsal ribs. Skates, chimaeras, & a few unusual teleosts such as sea horses have no ribs. living agnathans do not have ribs, Fish ribs strengthen the myosepta to w/c the longitudinally disposed locomotor muscles of the myomeres are attached. Tetra pods Early tetrapods -> ribs were assoc with vertebrae from the atlas to nearly the base of the tail. They later became restricted in length and in number, the short ones often ankylosing with transverse processes.

Skeletal enclosure for viscera of amniotes is provided by column, costal ribs, sternal ribs, sternum. No skeletal cage in amphibians. Amphibians. All ribs of anurans and urodeles have become very short. Anurans ankylosed to transverse processes. Ribs of apodans quite long Present in assoc with all except the first verteb and few of the more post ones. Snakes play vital role in locomotion. Among amphi, only ribs of anurans are not bicipital. Non Avian Reptiles Lizards and Crocodilians Long ribs on many of the trunk verteb and short ribs in most of necks Sphenodon have long ribs in most of trunk and ribs cont to tail. Geckos have ribs assoc with every cervical verteb More spexalized lizards usually lack ribs in axis and atlas. A half dozen or more of the post ribs of trunk of Draco are elongated.

They can be rotated outward to elevate a patagium (broad fold of body wall skin) whent hen becomes a winglike nenbrane on which lizard soars; not in use patagium folded at sides of body.

The long ribs of the ant thoracic region acquired a ventral anchorage, the sternum. Typical long tetrapod ribs occupy same position as the ventral ribs of fishes encircle coelomic cavity rather than extending laterally within the horizontal skeletogenous spetum Tetrapod rib and ventral ribs of fishes are not homologous structures. Most tetrapods are bicipital (having dorsal head) or tuberculum (having a ventral head) and the capitulum. Tuberculum articulated with extremity of a transverse process Capitulum articulated with hypocentrum. Hypocentrum reduced in size and articulation site of capitulum changed:

Turtles No cervical ribs Ribs of trunk fused with costal plates of carapace. Two sacral ribs not fused with carapace short, and their expanded distal ends are ankylosed to the ilia of pelvic girdle. Brace the girdle against the verteb column. Slender, riblike processes may be assoc with some of the caudal verteb. Snakes

Birds

Have long, curved ribs starting at 2nd verteb and cont to tail. No sternum for ribs to attach to. Ventral ends of ribs have ligamentous conn with integ scutes. Ribs participate in locomotion.

On two adj demifacets: one on the caudal end of a centrum and other on cephalic end of the next more post centrum. With a parapophysis In a facet on single centrum Site changes along length of trunk.

First two pairs of ribs artic with the last two cervical verteb at base of the neck. Generally movable and lack sternal segments The next 5 pairs are thoracic ribs with bony sternal segments. Thoracic ribs major part of thoracic basket. Ribs are thin, flat, and borad and most of them bear broad uncinate processes thin broad ribs and uncinate process = lightweight but sturdy thoracic body wall skeleton for attachment of powerful flight muscles.

In some mammals tuberculum becomes reduced to a vestige in one region of the column or another. Some other tetrapods, crocodilians trend toward fusion of the two heads until only tuberculum remained. Typical thoracic ribs consist of two parts:

Costal rib: adjacent to verteb Sternal rib: more distal and more ventral.

May remain cartilaginous (humans) costal cartilages.

Ribs caudal to chest cavity ankylosed to underside of synsacrum. Ribs are confined to trunk in adults. Developing embryos exhibit transitory uncinate processes are also found in some lizards and were present in some early labyrinthodonts. Mammals Recognizable ribs in mammals are generally confined to thorax. 9 pairs 24 pairs; 12 pairs are common.

Where the number is larger than 10, the rest are floating ribs costal ribs fail to reach the sternum Humans and apes other than orangutans have 12 pairs of thoracic ribs Apes have addl 2 pairs of lumbars. Humans freq have an addl rib (cervical or lumbar) as an anomaly. Studies of mammalian embryos reveal that two heads of a vestigial bicipital rib develop in assoc with each cervical verteb. One of heads is attached to transverse process; other to a centrum. Transverse foramen: foramen created by two head on each cervical verteb. Arteriovertebral canal: bony tunnel provided by a consecutive foramina. Transmits a vertebral artery from source at base of neck to brain. Vertebral artery exists in other reptiles but is not enclosed in a canal.

Sternum of archaeopteryx was small and lacked a keel, an indication that first birds were not strong flyers. Mammals

Composed of a series of bony segments sternebrae and is comparatively narrow comparison to that of reptiles. The last sternebra, the xiphisternum, cartilaginous or bony xiphoid process. bears

or in a

Marine mammals short sternum, only a small number of ribs reach sternum, an indiv sternebrae tend to be fused.

Condition in cetaceans correlated with the galloping mode of swimming necessitated by lack of functionally competent anterior flippers. Amniote Arises as paired mesenchymal bars that later unite and undergo chondrogenesis.

Tetrapod Sternum Sternum is a tetrapod structure and predominantly amniote. Of endochondral origin Presence, size, and anatomical relationships are correlated with the extent to which the forelimbs are employed in locomotion. Serves as base against which pectoral girdles and ribs are braced and as anatomical origin of ventral muscles of forelimbs. Largest in pterosaurs, craniate birds, bats which require strong flight muscles. Has acquired addl roles in some tetrapods such as assisting in breathing movements in birds. Amphibians Well differentiated only in anurans. Whether or not Necturus has a sternum depends on ones definition. Consists of more or less isolated nodules of cartilage assoc with the linea alba immediately caudal to the coracoid cartilage of pectoral girdle and extending into proximal ends of adjacent myosepta. Small but discrete sternal element is present in other salamanders Apodans have no sternum. Lizards Agile climbers and runners. Strong pectoral girdles are firmly braced against a substantial shield-shaped sternum of cartilage or replacement bone. Crocodilians Sternum is a simple cartilaginous plate attached to the procoracoid of the pectoral girdle. Caudally, continuous with an expanse of mineralized fibrous membrane incorporating gastralia. Turtles No sternum. 2 halves of pectoral girdles are untied by fibrous ligaments or by a cartilage. Birds

Presternal and suprasternal (mammals): also develop

blastemas

Presternal: contribute to the manubrium Suprasternal centers sometimes do and may rise to independent suprasternal ossicles. Some humans have suprasternal ossicles but are not found unless there is an occastion to xray the sternclavicular joint.

Sternum have developed an enormous keel or carina to which the massive pectoral flight muscles are attached. Development of carina is an instance of convergence evolution.

Comparative Vertebrate Anatomy Lec Chapter 9: Skull and Visceral Skeleton SKULL 1. Neurocranium or primary brain case 2. Dermatocranium 3. Splanchnocranium/Visceral Skeleton 1 & 3 of early craniates were formed in cartilage & are ossified to various degrees pattern still seen in development of craniates. membrane bones were added to the integument surrounding the skull and jaws. the upper jaw is visceral skeleton derived from the 1st visceral arch NEUROCRANIUM also called the endocranium, chondrocranium, or primary braincase protects the brain and certain special sense organs arises as cartilage (parachordal & prechordal cartilages) and is then replaced partly/wholly by bone (except in cartilaginous fishes) in gnathostomes: neurocrania starts out as several independent cartilages that later expand and unite into one cartilaginous braincase Cartilaginous Stage neurocranium commences as a pair of parachordal cartilage and prechordal cartilage underneath the brain Parachordal cartilage: parallel the anterior end of the notochord beneath the midbrain & hindbrain Prechordal cartilage/trabeculae cranii: develop anterior to the notochord beneath the forebrain Parachordal cartilage expand across the midline toward each other and unite. Basal plate: broad and cartilaginous; combination of notochord & parachordal cartilages Ethmoid plate: prechordal cartilages expand & unite across the midline at their anterior ends Sense capsules While prechordal and parachordal cartilages are forming, cartilage also appears in 2 other locations:

Olfactory (nasal) capsule: partially surrounding the olfactory epithelium. Incomplete anteriorly because water (in fishes) or air (in tetrapods) must have access to the olfactory epithelium. Otic capsule:completely surrounding the otocysts, which is the developing inner ear. Walls of both capsules are perforated by foramina that transmit nerves & vascular channels Optic capsule: forms around the retina Not the orbit or skeletal socket: where eyeball lies. Sclerotic coat of the eyeball Optic capsule is fibrous in mammals. Cartilaginous/bony plates often form within the sclerotic coat (an ancient condition ostraco, placo and basal osteich) optic capsule does not fuse with the neurocranium, eyeball is free to move independent of the skull therefore is not part of the neurocranium Completion of the Floor Walls & Roof Ethmoid plate unites w/olfactory capsules Basal plate unites w/otic capsules w/c are lateral to hind brain Ethmoid & Basal plate expand toward each other to form a floor to which the brain rests on hypophyseal fenestra: located at the midline between the 2 plates accommodates the hypophysis & internal carotid arteries (en route to the brain) the hypophyseal fenestra later reduces down to a pair of foramina transmitting the arteries. Further development: of the neurocranium involves: Construction of cartilaginous walls around the brain and as a primitive condition in some craniates, a cartilaginous roof (tectum) w/1 or 2 fenestra Cranial nerves and blood vessels are already present at this time Cartilage is deposited in such a manner to leave foramina for these structures. Foramen magnum: largest foramen located in the rear wall of the neurocranium As the brain, nerves, vessels, & sense organs grow, the neurocranium goes through remodeling (cartilaginous braincase, olfactory epithelia, & an inner ear). mesenchyme of the neurocranium comes from: Prechordal cartilage from neural crest ectoderm that streams ventrad in from of the developing eyestalks (optic stalks) that connect thre rbain with the dev eyeball. Mesenchyme that formes the parachordal cartilages, occupular and encapsulates the inner

ear comes from the sclerotome (epimeric mesoderm) Source of mesenchyme that forms the remaining bulk of the neurocranium is derived from the neural crest

cartilage: chondrosteans, basal neopterygians other than gars, dipnoans

Cartilaginous Neurocrania of Adult Craniates Living Agnathans: several cartilaginous components remain more/less independent throughout life an olfactory, optic, and otic capsule, basal plate, & notochord (not fused with the basal plate) are identifiable & also several other cartilages of unknown homology are present roof above the brain is unchondrified fibrous Cartilaginous Fishes (Squalus acanthias): typical of cart fishes constitutes a high water mark in dev of neurocrania because embryonic components unite to form a boxlike chondrocranium (cartilaginous braincase) Basic components have almost lost their identity. Fully developed walls In gnathostomes: we see a post occipital wall for the first time. cartilaginous roof last portion of the roof to chondrify is an area just behind the rostrum young skulls unossified fenestra otic capsules fused into the posterolateral walls of the braincase olfactory capsules fused to it anteriorly notochord is visible on ventral aspect as a ridge extending cephalad from the base of the foramen magnum sella turcica (cartilaginous pocket beneath the brain): holds the hypophysis neurocranium projects forward beyond the olfactory capsules as the rostrum occipital condyles on each side of the foramen magnum site of immovable artic between occipital region of the neurocranium and the first vertebra. endolymphatic fossa: on the posterodorsal aspect has 2 pairs of foramina that house endolymphatic duct: open to surface of the head perilymphatic duct +Chodrichthyes: well-developed, totally cartilaginous adult, devoid of any dermal bones Bony Fishes Cartilaginous neurocranium in embryonic structures of teleosts & tetrapods replaced partly/wholly by endochondral bone as development progresses)

Neurocranial Ossification Centers Process of endochondral ossification w/in neurocranium occurs more or less simultaneously at numerous separate ossification centers. Specific number of centers varies among species but four regional grps are universal Occipital Centers cartilage surrounding the foramen magnum may be replaced by as many as four bones: Basioccipital bone: produced by one or more endochondral ossification centers ventral to the foramen magnum; beneath the hindbrain 2 Exoccipital bones: produced by centers in the lateral walls of the foramen magnum Supraoccipital bone: above the foramen,

Mammals: all 4 bones may unite to form an

occipital bone

Modern Amphibians: one or more may remain

cartilaginous although they were bony in stem amphi Tetrapods: neurocranium articulates w/1st vertebra by one/two occipital condyles Amphibians: single condyle on the basioccipital bone Modern Amphibians & Mammals: 2 exoccipitals Living Reptiles: single condyle

Sphenoid Centers synapsid & reptilian lineages: ossification of the sphenoid region occurs independently Basisphenoid bone (anterior to basioccipital): ossified embryonic cartilaginous neurocranium underlying the midbrain and pituitary gland Presphenoid bone (in mammals): ossifies anterior to the basisphenoid Basioccipital & sphenoid bones create a bony platform for the brain Sidewalls in the sphenoid region: above the basisphenoid; formed by an addl presphenoid ossification in mammals. Laterosphenoid bone (in archosaurs): forms the lateral ossification of the sphenoid region Orbitosphenoid bone (in archosaurs): a separate interorbital septum Alisphenoid bone (in some mammals): helps to form the lateral wall but derived from the palatoquadrate cartilage not the neurocranium Sphenoid bone (in mammals): combination of the basisphenoid, presphenoid, & alisphenoid bones Pituitary rests in the sella turcica of the basisphenoid region no replacement bones develop above the brain Ethmoid Centers

anterior to the sphenoid includes ethmoid plate & olfactory capsules of the 4 ossification centers, the ethmoid tends to remain cartilaginous in tetrapods from apmphibians to mammals. basal tetrapods: no ethmoid centers Mesethmoid bones (in amniotes): contribute to cartilaginous nasal septum of birds & mammals Turbinal bones (conchae): anterior portion of interorbital septum; in the walls of nasal passageways of most reptiles, birds, & mammals Cribriform plate (in mammals): perforated by olfactory foramina that transmit bundles of olfactory nerve fibers from the olfactory epithelium to the brain Sphenethmoid (in anurans): sole bone arising in the ethmoid & sphenoid regions Ectethmoid develops in the lateral walls of the nasal passageway of Sphenodon not all cartilages in the nasal passages are from ethmoid ossification centers (i.e.winglike alar & sesamoid cartilages that keep the walls from collapsing against the nasal septum during inhalation)

Primary Palate bones: dermal bones

Opercular Bones A question of Homology homologues sometimes have diff names in diff species dermal bones of placoderms are not homologous to those in bony fishes roofing bones in mammals can be traced through basal tetrapods to their lobe-finned outgroups the frontal bone is a new structure among lobefinned fishes problem of naming roofing bones in ray-finned fishes; 2 possible hypotheses: Ray-finned & lobe finned fishes share ( via their common ancestor) parietal & postparietal bones w/the frontal being unique to later sarcopterygians The roofing bones evolved independently & are not homologous (& should be given distinct names).

Otic Centers otic capsule replaced by the prootic, opisthotic, & epiotic bones one/more of these can fuse with nearby replacement/membrane bones opisthotics fuse w/exoccipital (in nonavian reptiles) Periotic/petrosal bone: combination of prootic, opisthotic, & epiotic (birds and mammals) Temporal bone: the Periotic fuses w/squamosal Six ossification centers have been described in the otic capsule of a human fetus DERMATOCRANIUM Membrane bones of the skull collectively constitute the dermatocranium. earliest vertebrates most of the body was encased in dermal armor (i.e. ostracoderms) armor varied widely in extent to which it covered the body, have gone through expansion & reduction of dermal armor (small scales to large scales) Modern vertebrates reverses the trend of expansion & reduction less complete on the head region bony plates present in the skin of the head these bones in the skin are also part of the skull neurocranium come from endoskeleton dermatocranium/dermal bones come from exoskeleton basic structure

Roofing Bones early patterns seen in rhipidistian fishes & inherited almost wholly by labyrinthodonts provide shield over the brain & special sense organs w/openings for the nares, paired eyes, & a median (parietal) eye. In rhipidistians, a series of paired and unpaired scalelike bones extended along the middorsal line from the nares to occiput overlying the brain, olfactory capsules, and any other components of the neurocranium that developed in that area. in labyrinthodonts, the unpaired bones were missing & were replaced by paired bones (nasals, frontals, parietals, & postparietals) in modern amphibians, postparietals have disappeared as independent bones but may have coalesced with endochondral supraoccitals that appeared first in early reptiles. bones that form a ring around the orbit: lacrimal, prefrontal, postfrontal, & jugal bones at the posterior angle of the skull: intertemporal, supratemporal, tabular, squamosal, & quadratojugal abyrinthodonts developed a longer snout- this could be correlated w/altered methods of gathering/capturing food or manipulating it in the mouth Marginal Bones (dermal bones of the upper jaw) palatoquadrate cartilages, the only upper jaws that cart fishes develop, are the embryonic precursors of the upper jaw of bony vertebrates: in vertebrates, their upper jaws become ensheathed by tooth-bearing dermal bones: premaxillae & maxillae- both become part of the margins of the dermatocranium the completed upper jaw complex of bony verteb becomes part of the skull. Primary Palatal Bones

Roofing bones: bones that form above and

alongside the brain and neurocranium Marginal bones: dermal bones of the upper jaw

The primary palate: roof of oropharyngeal cavity (fishes) oral cavity (basal tetrapods) cartilaginous floor of the neurocranium (sharks) Membrane bones (vertebrates): applied to the underside of the neurocranium & to any part of the palatoquadrate (upper jaw) cartilages major components of the primary palate in rhipidistian fishes & early tetrapods, these membrane bones were unpaired parasphenoid (beneath sphenoid region), paired vomers (beneath ethmoid) & paired palatines, ectopterygoids, & pterygoids teeth-bearing bones (primitive) internal nares pierced the palate anteriorly a primary palate still exist in all tetrapods but w/modifications tetrapods w/a secondary palate have the primary palate in the roof of the nasal passageway.

Polypterus retain primitive features of Paleozoic

fishes: neurocranium well-ossified and is overlaid in the middorsal line by paired nasals, parietals, & postparietals of dermal origin lateral to these is a linear series of small, mostly unnamed bbony dermal plates. premaxillae&maxillae: overlie endochondral palatoquadrate of the 1st visceral arch preopercular bone: overlies the site of articulation of the upper&lower jaws other opercular bones complete the dermatocranium laterally gular bones: pairl and primitive; of dermal origin lie in the opercular membrane

Opercular Bones flaps of tissue that arises as an outgrowth of the hyoid arch & extends caudad over the gill slits membranous in holocephalans absent in elasmobranches stiffened by squamous plates of dermal bone in bony fishes different opercular bones: large opercular smaller preoperculars (articulation of the upper & lower jaws suboperculars interoperculars gular bones: lie in the opercular membrane in the floor of the opercular chamber in some basal bony fishes represented by branchiostegal rays located in the branchiostegal membranes in more specialized ray-finned fishes & dipnoans Neurocranial-Dermatocranial Complex of Bony Fishes combination of the neurocranium & dermatocranium Basal Actinopterygians sturgeons and spoonbills have a skull that justifies the name Chondrostei Sturgeons: neurocranium remains almost completely cartilaginous throughout life only traces of endochondral ossification in the neurocranium of sturgeons are isolated sites in the otic capsules & sphenoid bone where it contributes to the walls of the orbit. Elsewhere, neurocranium remains cartilaginous. Spoonbill: unusual bill is an extension of cartilaginous rostrum numerous overlying dermal bones hide the cart neurocranium and the cartilaginous components of the upper jaws and hyoid arch.

Basal Neopterygians (Amia & gars) Skulls bear little resemblance to the skulls of most modern fishes. Dermal bones are grooved & pitted They can be characterized equally well as skull bones or dermal scales Cheek plates of gars: compelling evidence that the dermatocranium may have been derived from early dermal scales Neurocranium of Amia remains highly cartilaginous Bony neurocranium (gars)

E. Jarvik: detailed account of anatomy of skull of

Amia Teleosts skulls are highly specialized & architecturally diversecorrelated w/diverse feeding habits of the group w/c includes species that harvest plant tissue or capture animal life in every conceivable niche. Combination of highly maneuverable jaws&palates Largest # of bones in any modern vertebrate skull Skull of carp: is compressed laterally and vaulted dorsally Maxillae, pre, dentary, articular, quadrre, and symplectic assoc with jaws and hyoid arch Posttemporal is the dorsalmost segment of the pectoral girdle and has been secondarily incorporated with the skull Mesethmoid, epiotic, and pterotic incorporate some membrane along with unusual replacement bone. Branchiostegal rays in paired branchiostegal membranes beneath gill chamber take place of gular bones Neurocranium fully ossified except olfactory capsules in cyprinids (carps) neurocranial ossification is less and islands of cartilage on surface of the skull are unossified regions of the neurocranium palate reflects that of ancestral bony fishes: vomer, parashpenoid, & pterygoid bones names given to bones of skull have been assigned on basis of loc, shape or other char.

Dipnoans did not experience burst of speciation changes in skull have been more conservative dermal bones increased in number and decreased in size Cartilaginous neurocranium the dermatocranium of lungfishes has evolved from a large # of scalelike dermal bones in fossils to a few broad bony plates in modern species palate accommodates nasal canal openings into the oral cavity just behind the mouth Neurocranial-Dermatocranial Complex of Modern Tetrapods Amphibians Skulls are modified from those of labyrinthodonts, though still flattened (platybasic) Neurocranium incomplete dorsally Much remains cartilaginous except in apodans rigid skulls of apodanscorrelated w/burrowing replacement bones in anurans & urodeles: sphenethmoid 2 prootics 2 exoccipitals w/condyles columella(stapes): skeletal rod abutting against otic capsule in amphibians middle ear ossicle that conducts sound waves from an eardrum to the otic capsule incomplete dermatocranium bones around the orbit have been lost except the lacrimals & prefrontals in basal urodeles bones of the temporal region have been lost (intertemporals, supratemporals, tabulars, & postparietal)loss of these bones leave the otic capsules (prootic) exposed dorsally and laterally. Only squamosal and sometimes a quadrotojugal remains Premaxillae and maxillae: usually present to ensheath the upper jaw cartilages at the margin of dermatocranium in perennibranchiates, sphenethmoid fails to ossify & maxillae may fail to develop primary palate is altered in anurans&some urodeles because of large palatal spaces that evolved beneath the orbit reduced palatines to transverse splinters that brace the upper jaw against the palate anteriorly reduced pterygoids to a pair of bipartite bones that brace the upper jaw against the braincase posteriorly can enable the eyeball to retract until the membranous floor of the orbit bulges into the oral cavity in urodeles, parasphenoid has become exceptionally broad and palatal vacuities are lacking in some orders. Nonavian Reptiles Primitive features Skulls of stem reptiles were little changed from those of labyrinthodonts.

Some primitive features are still present in reptiles. well-ossified neurocranium single occipital condyle large complement of membrane bones median eye housed in a parietal foramen (in rhynchocephalians & lizards) Major changes (inherited by birds) appearance of temporal fossae partial/complete secondary palate fully ossified neurocranium of an adult alligator consists of: two exoccipital bones supraoccipital basioccipital bearing a single condyle for articulation with atlas. Laterosphenoids Ethmoids Several otics that arise from separate ossification centers in the otic capsules. None of the otic capsules are on the surface. Some are coalesced with adj occipital Chiefly cart neurocranium of a juvenile Sphenodon is seen in situ It will ossify further with age, but more cart remains in adult sphenodons and lizards than in any other reptiles. crocodilians retain the largest # if membrane bones turtles have the most enigmatic (mysterious) skull

Temporal fossae (t.f.) temporal fossa: deep opening in the temporal region bounded by 1/more bony arches Stem reptiles: had none (retention of primitive cond) anapsid (no tf) lacking temporal arches Today: among living reptiles, turtles are anapsid. Synapsida (1 lateral tf): developed a single lateral temporal fossa surrounded by the postorbital, squamosal, and jugal bones,. last 2 forms an infratemporal/zygomatic arch synapsid skull transmitted to mammals. zyg.arches human cheek bones.

Archosaurs, snakes, and predecessors of Sphenodon

lizards had superior and inferior temporal fossa diapsid: 2 fossae lower arch: zygomatic arch of mammals

upper arch: supratemporal (arch between two

fossae) consists of parts of the postorbital and squamosal bones. euryapsid (1 t.f. dorsally located): ichtyhsaurs & plesiosaurs (both extinct) crocodilians and Sphenodon still have diapsid skulls. But modern lizards have lost part or all of the lower arch, and snakes lost both of it. loss heft a cavernous void in the posterolateral walls of squamate skulls. modified diapsid skulls. Reduction or loss of arches along with the acquisition of intracranial joints facilitated

cranial kinesis: movement of one secion of a skull independent of other.s. Ichthyosaurs and plesiosaurs had one temporal fossa, dorsally located, that resembled the dorsal fossa of diapsids in its anatomic relation. Euryapids: instance of evolutionary convergence. All euryapsid reptiles are extinct. Temporal region of a turtle skull is an enigma. Absence of temporal region is a primitive condition. But there has been extensive loss of dermal bones and excavation of temporal region dorsal to otic capsule: condition more pronounced in some family Supratemporal, tabular, and postparietal bones are missing Postorbitals and postfrontals: united Parietals give impression of having receded from rear provide space & surfaces for accommodating adductor muscles to operate the lower jaws of amniotes labyrinthodonts and extinct basal reptiles: adductor mandibulae was confined to cramped qtrs internal to the temporal region of the dermatocranium at best, this muscle enabled bony fishes and early tetrapods to seize food, bite and close the mouth and food was swallowed whole. Provide access to surface of dermatocranium so that one portion of adductor, the temporalis muscle, could acquire more leverage by spreading upward onto the temporal region of the skull. Provide zygomatic arch to which another portion of the adductor, the masseter, could acquire an anchorage. Powerful adductors + other muscles of mandibular and hyoid arches = complex side-to-side, forwardbackward, and rotary chewing movements

Cranial Kinesis/kinetism movement of a functional part of a skull independent of other parts made possible by intracranial joints between the 2 parts (teleosts, lizards, snakes, & birds have these joints examples: most of them move the upper jaw and palate as a unit independent of the neurocranium when they open their mouth some teleosts even two sides of upper jaw can be moved movements cannot be made by crocodilians, frogs, salamanders, mammals. correlated primarily w/food-getting & manipulation of food enables some squamates to open their mouth wide enough to swallow prey larger than their own head. Palate and temporal region of squamates and some species of birds have become so modified for feeding. parasphenoid of the primary palate is not kinetic Birds skull is fundamentally reptilian w/modification correlated w/flight, feeding habits, & larger brain. Some of the roofing bone has been lost and remaining dermal bones are much thinner. Most of the sutures have been obliterated 2 functional regions of the skull: solid bony neurocranium & dermatocranium (rear) houses brain, olfactory organs, eyeball and equilibatory and hearing complex, protects all structures needed for input and processing information. beak & palate (front): food procuring and handling area post component is highly vaulted, bulging outward and arching upward alongside the greatly enlarged brain. Orbits are situated where they achieve maximal arcs of visibility, and they reflect size of the massive eyeballs. parietal foramen has closed neurocranium incomplete dorsally

Secondary Palates Secondary plates: horizontal partition that partially/completely divides the primitive oral cavity into separate oral & nasal passageways displaces internal nares (posterior choanae) caudad verteb with 2ndary palate, primary palatal components remain in roof of nasal passageway no parasphenoid bone is present in mammals

well-ossified neurocranium w/ a cartilaginous

Crocodilians: palatal processes of the premaxillae,

maxillae, palatine, & pterygoid bones meet in the midline to form a long bony secondary palate w/internal nares. reptiles: secondary palate is incomplete, therefore the respiratory airstream is channeled in a palatal fissure (fairly deep longitudinal groove) in the roof of the oral cavity palatal folds: fleshy borders of the fissure mammals: secondary palate extends all the way to the pharynxsoft palate lacks bone

olfactory capsule & the mesethmoid component of the interorbital septum contains 1 occipital condyle modified diapsid skull, the arch between the superior and inferior temporal fossa having been lost. Preorbital fossa, separated from the orbit by the lacrimal bone, is present. Infratemporal arch, consisting of primitive complement of jugal and quadrotojugal bones, is intact but very slender. Kinetic palate resembles that of squamates except that the ectopterygoids have been lost. immobile parasphenoid bone fused to the basisphenoid

not all bird skulls are equally kinetic (ex. woodpecker)

temporal complex consists of: numerous

Mammals major features that differentiate mammalian skulls from other amniotes: emergence of dentary as sole bone of the lower jaw altered site of articulation of the lower jaw w/the braincase alterations in the secondary palate 3 bones in the middle ear differ from members of reptiles in their modifications in the temporal region of the dermatocranium. synapsid skulls and skulls have become increasingly doomed as the cerebral hemispheres have ballooned dorsally, laterally, and caudad. incomplete dorsal neurocranium: soft spots called fontanels (newborn humans) enable to be molded during delivery through the birth canal. 1 or more bregmatic bones may ossify in the frontal fontanel in some species; single bone may develop as an anomaly in humans antiepileptic bone- Paraclesus -- served as a pop-up valve for relieving pressure w/in the cranial cavity basioccipital, basisphenoid, and presphenoid bones form a floor on w/c he brain rests exoccipitals, alisphenoids and lateral extensions of parasphenoids from partial side walls. A supraoccipital completes a bony ring that resembles a vertebra. ethmoid cartlages & bones house the olfactory epithelium, underlie the olfactory bulbs, and make up much of the nasal septum (mesethmoid) w/c are perforated by foramina for olfactory nerve bundles ossification centers in the otic capsules unite solidly to form a petrosal bones beneath the overgrown temporal lobes of the mammalian cerebral hemisphere often incorporated into a temporal bone complex each exoccipital bone bears a condyle sequence of centrumlike basioccipital, basisphenoid, and presphenoid bones and their dorsal wings resembles a series of three successive vert that are incomplete dorsally. inspired Goethes Verteb Theory of Origin of the Skull dermatocranium is represented in mammals by paired premax, max, jugals, nasalas, lacrimals, and squamosals; by paired or unpaired frontals and parietals; and by an unpaired interparietal a postparietal was present in Homo erectus and is still present in some human populations -> called an inca bone premax are not identifiable in adult human skulls because they unite with the maxillae early in embryonic life zygomatic arch varies from massive to slender, depending on the magnitude of force exerted by the masseter. Become incompelete or delicate in insectivores

components of intramembranous and endochondral origin: squamous portion: squamosal of lower tetrapods. tympanic bulla: consists of 2 parts: tympanic (surrounds eardrum as bony ring) said to have derived from angular bones of nonmammalian amniotes. entotympanic (cartilage replacement ossificationa new structure in some mammals) petrous portion (ossified otic capsule) fused prootic, ophisthostic, & epiotic mastoid portion (endochondral origin -- new in mammals) mastoid process petrotympanic bone: combined petrous & tympanic portions

syloid process: dorsal segment of hyoid arch

skeleton coalesces in some species with the temporal bone squamosal bone has become a new site of articulation of the lower jaw with the skull in mammals. Shift from quadrate-articular joint of other verteb is correlated with expansion of dentary bone during mammalian evolution air-filled cranial sinuses: found w/in the maxilla, frontal, sphenoid, & ethmoid bones an unpaired vomer lies at the base of the nasal septum septum consists of a mesethmoid bone and more or less cartilage. nasal process of the palatine in the lateral wall of the nasopharynx: contributes to the orbit wall palatal process of palatine bone: contributes to secondary plate pterygoids reduce to small,winglike pterygoid processes of the sphenoid complex serve as anatomic origin of the powerful pterygoid muscle. Parasphenoid and ectopterygoids have been lost. Bony part consists of palatal processes of premax, max, and palatine bones. Caudal to the bony part is a membranous soft palate. Mammals have three pairs of scroll-like turbinal bones (nasal conchae) inferior (maxillary) concha is an independent scroll middle & superior conchae are appendages of the ethmoid bone two lower conchae covered with olfactory epithelium scroll-like structure of conchae increases the surface area of these epithelia. Most reptiles other than turtles have one pair of turbinals and birds have two.

features unique in mammals: ossified posterior tips of the palatoquadrate & Meckels cartilages in the middle ear cavity which serve as ear ossicles w/the columella

Reduction in # of Cranial Bones during Phylogeny # of indiv bones, especially membrane bones in the mammalian skull, has tended to be reduced during evolution of tetrapods. reduction is a result of fusion of adjacent embryonic ossification centers, phylogenetic loss of ossification centers, and obliteration of sutures in young animals membrane bones often unite w/adjacent replacement bones to make a single bone w/dual history postfrontals and supratemporals sometimes unite with replacement bones of the otic capsule to form spehnotic and pterotic bones. Squamosal unites with otic and other elements to contribute to temporal bones. Mammalian interparietal, a membrane bone, may unite with supraoccipital. Unions reduced number of bones in skulls.

Squalus acanthias: generalized vertebrate except that it lacks bones. Visceral skeleton: Consist of cartilages in each pharyngeal arch, and median basihyal, and basibranchial cartilages in the pharyngeal floor. Skeleton of each arch conforms fairly closely to a basic pattern, and all but the first and last in Squalus support gills. First arch and, to a degree, the second are modified for procuring food. Skeleton of the mandibular arch consist of two cartilages on each side: palatoquadrate cartilage-dorsally; Meckels cartilage-ventrally Left and right palatoquadrate cartilages meet in the midline dorsally to form the upper jaw, Meckels meet in the midline ventrally to form the lower jaw. Slender Labial cartilage unknown significance; extend from the angles of the mouth into a position where lips would be Skeleton of the hyoid arch consists of: paired hyomandibular cartilages- dorsally;

gill-bearing ceratohyals laterally at the angle of the mouth: Meckels cartilage and
palatoquadrate articulate with one another and with the hyomandibular cartilage in a movable joint united by ligament dorsal end of the Hyomandibula is bound by ligaments to the otic capsule and suspends the jaws and the entire branchial skeleton from the neurocranium hyostylic jaw suspension

VISCERAL SKELETON or splanchnocranium the skeleton that develops within the pharyngeal arches.

in fishes: skeleton of the jaws and gill arches in tetrapods: skeleton has become modified to perform new functions on land blastemas that give rise to the visceral skeleton comes from the neural crest and they first produce cartilage Later, the cartilage might by partly or wholly replaced by bone. Only in the first arch it is ensheathed by dermal bone.

Sharks No bone forms Visceral skeleton is seen it its primitive role, that of supporting the jaws and gills.

Jaw Suspension in Fishes jaw-hyoid complex of fishes must necessarily be braced against some support. The nearest is the braincase. Elasmobranch and most bony fishes -- the hyomandibular is braced against the otic capsule, and the posterior end of the palatoquadrate cartilage is braced against the hyomandibula hyostylic jaw suspension or hyostyly Amphistyly seen in more primitive condition of older sharks; the hyomandibula and one or more processes of the palatoquadrate are braced independently against the braincase Autostyly oldest; found in very early shark independently evolved with modifications in chimaeras and lungfishes the palatoquadrate is attached to the neurocranium, and the hyomandibula plays no role in jaw suspension structural details of jaw suspension among fishes, ancient and modern, vary greatly, and an expanded terminology hjas been devised for this variants. Anatomic relationship of palatoquadrate to the hyomandibula and to the neurocranialdermatocranial complex is correlated with the diet of the species.

Bony Fishes Visceral skeleton: Resembles that of shark in basic morphology Difference: the embryonic palatoquadrate and Meckels cartilage in bony fishes become invested by membrane bones during development, the hyoid skeleton may consist of a greater number of segments, and the embryonic cartilages of gill arches are replaced by bone Except mammals: the site of articulation between the upper and lower jaws of bony fishes and of all other vertebrates is the same as in the cartilaginous fishes Posterior ends of palatoquadrate cartilages articulate with the posterior ends of Meckels cartilage Fate of the embryonic palatoquadrate: Cartilage becomes ensheated by two membrane bones, premaxilla and maxilla The palatal portion (portion in roof of oropharyngeal cavity) develops 2-3 sites of dermal ossification including palatine and ectopterygoid bones Posterior tip of the palatoquadrate cartilage ossifies to become a quadrate bone Entire complex of ensheathing and replacement bones unites with the dermatocranium to one degree or another, depending on the species. Articular bones ossified Meckels cartilage at their caudal ends Dentary and Angular the remainder of the cartilage then becomes invested by several membrane bones Mandible of Basal Amia consist of: dentary, angular, surangulars, prearticulars, and 4 pairs of coronoids All but the angular and surangular bear teeth Symplectic and interhyal ossification centers in the hyomandibular cartilage Symplectic usually articulates with the quadrate bone or with the quadrate and lower jaw, where it participates in cranial kinetism. Hyomandibula lost in fishes. Epihyal ossification center in the ceratohya Bony gill arch consist of 4 segments: Hypobranchial Ceratobranchial Epibranchial Pharyngobranchial Only middle of 2 segments bear gills. Pharyngeal borders of branchial arches are covered with bony dermal plates having flattened or pointed denticles. Size and arrangement of plates and denticles are correlated with the nature of diet and the method of food handling in the pharynx.

Feeding mechanisms: They had wide mouths, hinge of jaws was far back under the skull and the upper jaw was fused with the braincase incapable of independent movement Mandible was simply lowered and then snapped shut on prey in manner of some sharks. The portion of prey that was inside the orobranchial chamber was swallowed. Appearance of type of kinestism seen in teleosts more complex mechanical means of procuring food. The large adductor mandibulae muscle inserted farther and farther forward on the mandible, an additional array of jaw muscles developed, and the mouth become narrower and more oval Effect of these changes was that the jaws of the most highly specialized telosts can be thrust forward and employed in feeding by inertial suction. Mechanism works in herbi and carni vorous fishes. Herbivorous fishes: feeding by inertial suction. feeds on algae attached to submerged objects; abduction of the hyomandibula expands the orobranchial chamber, lower jaws fall open, and the opercular cavity is then expanded creating forward suction In zooplankton: slowly drawn to the mouth. the lower jaw is then raised, the upper jaw remains briefly protruded, and the hyomandibula is adducted, which compresses the chamber two phases of the pharyngeal pump, expansion and compression, occupy together about 600 milliseconds. By manipulating the hyoid skeleton and jaws appropriately protruded jaws can be directed upward for collecting food on surface or downward for food on the substrate. Predatory fishes: similar mechanism works. When hyomandibula is drawn forward, symplectic forces the kinetic elements of the upper jaw and palate to slide forward, and the premax and dentary bones close on the prey. A process of maxilla becomes attached by a ligament to the dentary movement of one participates in displacing the other. Snapping and biting has not gone out of style. Onehalf or more of the teleosts have nonprotrusive jaws. Theory has been advanced that vertebrate jaws are former gill arches that become modified Basis for this idea: Hyoid arch does not contain demibranch in Squalus Spiracle is a pharyngeal slit as are all gill slits. Nerves and arteries of the mandibular and hyoid arches most anterior of a series that is repeated in each gill arch. Theory is an interesting speculation

Living Agnathans Visceral skeleton: Unlike that of jawed fishes. No identifiable palatoquadrate cartilages, Meckels cartilages, hyoid caritilages, or branchial arch cartilages Lingual cartilage - v-shaped bearing horny teeth and located in the floor of the buccal cavity moves in and out of the buccal funnel, serving as a rasping tronguelike organ Operated by protractor and retractor muscles attached to a narrow, segmented, immobile basal plate cartilage lying immediately beneath it. No evidence that any of the cartilages associated with the feeding apparatus are derived from ancestral visceral arches. Lingual cart and assoc cartilages are correlates with the unique method of feeding characteristics. Rest of the pharyngeal skeleton consists of a branchial basket w/c is simply a fenestrated framework of cart immediately under the skin. Caudal end of it in lamprey is assoc with the level of heart. Tetrapods Visceral skeleton underwent profound changes: Some previously functional parts have been deleted Those that persisted formed new and sometimes unexpected functions Larval frogs: have 6 pairs of visceral cartilages (last 4 bears gills) Hypobranchial plate: where cartilages of the gill arches meet ventrally During metamorphosis, 3rd 5th 6th visceral cartilages recess. Hypobranchial plate enlarges and along with 1st basihyal becomes incorporated into the body of the hyoid: Latter of it becomes: Ceratohyal cartilage of 2nd arch: becomes a slender anterior horn (cornu) of the hyoid apparatus (a broad cartilaginous and bony plate in the buccopharyngeal floor) Ceratohyal cartilage of 2nd arch becomes a slender anterior horn or cornu of they hyoid apparatus Cartilage of 4th becomes posterior horn

Embryonic Meckels cartilage of amniotes may

continue to grow and become a prominent core of cartilage within the mandible Cartilages become ensheathed by dermal bones that included a dentary, angular, surangular, splenial, coronoids, and prearticular Modern amniotes had a smaller numner, and mammals have only the dentary. In bony fishes: the posterior ends of each Meckels cartilage ossifies to become an articular bone at the hinge of jaws In mammals: it becomes malleus, another middle ear ossicle

Expansion of the dentary and a new jaw joint in mammals Temporal fossae enabled the adductor mandibulae muscles of synapsids to enlarge, subdivide, and acquire anatomic origins on the temporal region of the skull and zygomatic arch Increased mass of these new muscles was accompanied by expansion of the dentary bone and formation of a ramus on w/c the temporalis muscle inserted. Other dermal bones of mandible disappeared Articular bone became an ear ossivle Mammals were left with a mandible consisting of two dentaries 2 sites of articulation of the lower jaw with the skull Articular against the quadrate (primitive one) Condyloid process of the dentary against squamosal (new one) Eventually, articular and quadrate were captured by middle ear cavity and only the new articulation remained in mammals. Shape, slant, and relationships of condyloid process of mandible differ with demands made on it by the feeding mechanism of the various mammalian orders. Ear ossicles from the hyomandibula and jaws Hyomandibular cartilage of sharks is interposed between quadrate cart and otic capsule (houses inner ear) In an autosylic jaw hyomandibular can be dispensed with dipnoans persisted in tetrapods but became disassociated from the quadrate and attached to the tympanic membrane Otic capsule houses the inner ear; became surrounded by a portion of the first pharyngeal pouch middle ear cavity Columella stapes; 1st ear ossicle that came from the hyomandibula Conducted soundwaves from eardrum across the middle ear cavity to the inner ear. When dentary bone of therapsids acquired articulation with squamosal, articular and quadrate bones were freed to perform a new function or to disappear. Caudal end of Meckels cartilage malleus of mammals

As a result, the visceral skeleton initially

adapted for branchial respiration becomes converted to one adapted for life on land.

Fate of the palatoquadrate and Meckels cartilage Quadrate remains as the site of articulation of the cranium with the lower jaw in all nonmammalian tetrapods Incus quadrate bone in mammals; middle ear ossicle

 Embryonic Meckels can be seen projecting into

an area where middle ear cavitation is proceeding and its cartilaginous post tip can be seep to separate, ossify, and become the malleus Evidence that the quadrate bone of synapsids became the incus of the middle ear of mammals: Articular and quadrate bones have been articulating in a diarthrosis Articular has separated from the lower jaw and is now the malleus Quadrate has disappeared from the upper jaw Articular still articulates in a diarthrosis with a bone (incus) C. Reichert proposed the theory that mammalian ear ossicles are derived from the jaws E. Gaupp included the origin of the stapes from the dorsal tip of the hyoid arch

Tympanohyal dorsal most and ends in a

notch in the tympanic bulla In rabbits: the anterior horns are short ones (lesser horns); tympanohyal is represented by a slender stylohyal bone embedded in the tendon of insertion of the posterior belly of the stylohyoid muscle In humans: anterior horns are equivalent to ceratohyals, are lesser horns; unossified stylohyoid ligament represents epihyal and stylohyal segments; tympanohal is attached to the temporal bone = styloid process Hyoid anchors the tongue of tetrapods;

skeleton for the buccopharyngeal pressure

Amniote hyoid Consists of body in the pharyngeal floor just anterior to the pharynx and of and 2 or 3 horns (cornua) in the pharyngeal walls. Hyoid of anurans: derived at metamorphosis from basibranchial and hypobranchial cartilages, from the skeleton of hyoid arches, and from larval gill-bearing arches In amniotes: derived from homologous anlagen; cartilages of the 2nd arch becomes anterior horns, and 3rd arches or 4th becomes additional horns In lizards and birds: entoglossus (an elongated bony process) extends from the body forward into the long darting tongue In some male lizards, a similar process extends caudad into the dewlap or gular pouch. Snakes have no hyoid branchial skeleton is vestigial. Entoglossus and caudal horns of woodpeckers is a remarkable tool for impaling grubs From the base of the entoglossus, two long flexible caudal horns loop caudad then dorsad around the occipital region of the skull and forward under the scalp all the way to the Lore space between the eyes and the bill where both horns dip into the right nasal passageway. They then continue forward in the nasal canal a short distance father and terminate. The termination od both horns in a single nasal passageway is surprising and lacks an explanation. When impaling grubs, the horns are straightened by a muscle that shoots the tongue into the prey Elastic recoil immediately returns tongue to the mouth. In mammals: has cranial horns from the 2nd arch and caudal horns from the 3rd In dogs and cats: the cranial horns are the longer ones (greater horns) and composed of 4 segments

pump used in respiration in anurans; provides attachment for some of the extrinsic muscles of the larynx, has subtle effects on lower jaw movements, and site of attachment of muscles that participate in swallowing associated hypobranchial and branchiomeric muscles of amniotes approach the hyoid from many directions lower jaw, larynx, sternum, clavicle, the temporal region of the skull, and elsewhere. These muscles stabilize the hyoid in a single position or move it forward, backward, up or down.

Laryngeal skeleton Cricoids and arytenoids cartilages replacing bones (in tetrapods) Mammals: have thyroid cartilages or bone in addition Thyroid cartilage arise from mesenchyme of the 4th and 5th pharyngeal arche Cricoids and arytenoids cartilages probably products of 5th arch Because the caudal end of the pharyngeal arch series has been subject to reduction during evolution, it is not surprising that problems are encountered in relating the caudalmost laryngeal cartilages to specific arches. PERSPECTIVE It is evident that the visceral skeleton is a functional complex that was associated primitively with feeding and branchial respiration. Tetrapods: it has been modified in part for transmission of airborne sounds, for attachment of tongue muscles, and for support of vocal cords.

Know by heart TABLE 9.4 (p. 196) Know FIGURES o 9.2 o 9.4 o 9.6 o 9.8 o 9.9

o o o o o o

9.10 9.18 9.19 9.20 9.32 9.41