January 16, 2014 VERTEBRAL COLUMN, RIBS, & STERNUM Endoskeleton a. Somatic axial and appendicular skeletons b.

. Visceral cartilage or bone associated with gills and skeletal elements (such as jaw cartilages) derived from them Axial Skeleton elements along the mid-sagittal axis of the body; includes notochord, vertebrae, ribs, sternum, and median fins Description forms around and sometimes invades the embryonic notochord during ontogenesis. forms in the lateral body walls forms in the ventral body walls (tetrapods) Embryonic Origin mesenchymal sclerotome of epimere scleroblasts migrating laterally into myotomes mesenchymal tissues in ventral midline

Established as: 1. Hyaline Cartilage replaced by endochondrial ossification 2. Intramembranous Bone Formation direct ossification Vertebral Column Keystone of vertebrate skeleton Segmented, flexible, arched rod to which head is attached and the rest of the body is suspended Main Functions: 1. Protect the spinal cord 2. Provide support to internal organs via dorsal mesentery 3. Locomotion - via trunk muscle attachment Fish Surrounded by environment for forward locomotion and buoyant effect on body Articulations between vertebrae permit only side-to-side flexibility of column

Type of Skeleton Vertebral Column Ribs Sternum

Interocoelus outpocketing of mesoderm Fate of the Mesoderm I. Dorsal or Paraxial Mesoderm epimere or somites a. Myotomes skeletal muscles b. Sclerotomes vertebral column and ribs c. Dermatomes dermis of skin II. Intermediate Mesoderm mesomere; kidney tubules, longitudinal ducts of urogenital system III. Lateral-plate Mesoderm hypomere a. Somatic mesoderm + ectoderm : somatopleure b. Splanchnic mesoderm + endoderm : splanchnopleure c. Coelom cavity between somatic and splanchnic mesoderm Skeletal Development (Multiple Cellular Origins) 1. Paraxial Mesoderm/epimere Somite, sclerotome Axial skeleton (e.g. vertebra) 2. Lateral Plate Mesoderm/hypomere Appendicular skeleton (e.g. limbs) 3. Neural Crest Head skeleton

Tetrapod Land is beneath the animal not surrounding the body Trunk is suspended between the forelimb and the hindlimb Dorsal-ventral flexion correlated with regional specialization of vertebral column Basic Parts of a Vertebra 1. Centrum vertebral body Occupy position occupied early in ontogeny by notochord Large solid disk that takes compressive forces during body movement 2. Arches

Neural arch above the centrum Haemal arch beneath the centrum in tail vertebrae - chevron bones (amniotes)

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Process or Apophyses Projections from arches and centra Some give rigidity to the column Articulate with ribs Serve as sites for muscle attachment

Types of Apophyses or Processes 1. Diapophysis (transverse process) Most common Attachment for muscles Articulate with ribs that project laterally into the horizontal septum that separates epaxial and hypaxial muscles 2. Zygapophyses (only in tetrapods) Paired processes at cephalic end of the trunk (pre-zygapophysis articular facet face dorsad) and at caudal end (post-zygapophysis ventrad facing) interlocking arrangement limits dorsoventral flexion of column Parapophysis Lateral projections from centra of few tetrapods Articulation site for capitulum of bicipital rib Hypapophysis Prominent midventral projections in centra of snakes and other amniotes Attachment for certain muscles and tendons

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MORPHOGENESIS OF VERTEBRAE Migration of sclerotomes Differentiation of sclerotomic segments each segments differentiated into a. Cephalic part (less condensed) b. Caudal part (more condensed) Development of Vertebra Sclerotome cells surround notochord on both sides o Cranial loosely arranged cells o Caudal densely packed cells Each vertebra is derived from 2 sclerotome segments o Caudal (dense) cells from a cranial sclerotome o Cranial (loose) cells from the next caudal sclerotome Intervertebral disc o Located between vertebra o Forms at the interface between loose and dense cells (center of sclerotome) Development of Intervertebral Discs 1. Densely packed cells move cranially to the middle part of each segment. 2. Formation of the peripheral part annulus fibrosus

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Enclosed notochord expands and undergoes mucoid degeneration Formation of the central part nucleus pulposus

The Vertebral Column of Fishes Highly diverse due to enormous species that evolved Variations in morphology due to developmental factors o How scleroblasts invade and thicken notochord o Extent notochord persists within adult column o Number of chondrification and ossification centers in perichordal blastemas o Extent which separate centers remain independent or coalesce For single species, 2 major morphologic regions of specialization: a) Trunk (dorsals) b) Tail (caudals) 1. Primitive Fishes Vertebrae are uniform Little regional differentiation whether cartilage or bone Has no hard skeletal parts but dermal bone elements in the head region Early vertebrates (ostracoderms) had so much dermal bone they were called armored fishes Agnathans Only skeletal elements associated with the notochord are lateral neural cartilages.

2. Development of the Body of Vertebrae 1. Caudal part remained fused with cephalic part adjacent to it to form mesenchymal centrum. 2. Notochord degenerates and disappears when surrounded by vertebral body. Development of Neural Arch 1. Sclerotomic tissues migrate backward from both sides of the centrum and surround the neural tube. 2. Neural spine forms at the meeting point of the neural arch. 3. Sclerotomic tissue also extends laterally from both sides of the centrum to form 2 processes: (a) costal ventral & (b) transverse dorsal Comparative Features Notochord Stiffness: flexion of body Vertebrae Greater strength Levers for muscles 3.

*Notochordal remnants remain in vertebral column in all vertebrate classes. *In mammals: vestige persists in intervertebral discs as a pulpy nucleus

Cartilaginous Fish Notochord present throughout adult and constricted within each centrum Centra (composed of chordal and perichordal cartilage) are concave at each end (amphicoelus) Each rib articulates with a transverse process of a centrum Vertebral canal Paired dorsal plates that constitute a neural arch Paired dorsal intercalary plates between arches Paired wedge-shaped supradorsal cartilages (in some species)

Exceptional Vertebrae : Chondrostei & Coelacanth No centrra Notochord present and unconstructed, its thick fibrous sheath contains little cartilage or bone Associated structures (basidorsals, interdorsals, basiventrals, interventrals) products of separate chondrification centers. 5. Exceptional Vertebral Column : Holocephalan (extinct) Thick rings of calcified cartilage deposited in notochord sheath Vertebral rings more numerous than body segments 6. Bony Fishes or Teleosts Well ossified amphicoelus vertebrae Core of centrum includes remnants of notochord *Monospondylous vertebra one centra/segment *Diplospondylous vertebra two centra/segment e.g. tail of some fishes and of few primitive tetrapods *Intercentrum in fish Amia no neural arch *Embryonic basidorsal & basiventral incorporated in centrum (w/ neural arch) *Embryonic interdorsal & interventral incorporated in intercentrum (w/out neural arch) Evolution of Tetrapod Vertebrae Vertebral column of modern tetrapods: centrum and neural arch Vertebral column of earlier tetrapods: consisted of several bones or segments Hypocentrum/Intercentrum: large, median, U-shaped piece 2 Pleurocentra: smaller; overlying notochord dorsolaterally ; notochord is constricted at the level of each hypocentrum Rachitomous Vertebra consists of several pieces

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Amphicoelus Concave at both ends Present in most fishes and in few salamanders (Nectutus) and caecilians Procoelus Concave in front and convex in back Present in anurans and modern reptiles Opisthocoelus Convex in front and concave in back Present in most salamanders Acoelus Flat ended Present in mammals Heterocoelus Saddle-shaped centra at both ends Present in birds

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*Pleurocentra (2 smaller) to increase *Hypocentra (single and bigger) to decrease Types of Vertebrae

Regional Specialization in Tetrapod Columns 1. Limbs push against the earth Purely for locomotion Supporting the body above graound 2. Opposing physical force exerted by the earth in response to push of the posterior appendage. Transmitted via pelvic girdle to hindmost trunk presently sacral 3. Survival on land with irregular contours and obstructions to vision (rocks and vegetation) Facilitated by increased mobility of head with special sense organs (e.g. eyes to scan the horizon) Mobile joint between skull and first vertebra Anterior vertebrae: shortening or eliminating ribs and increase the mobility of intervertebral joints Vertebra/Vertebral Description Region Atlas/ Atlas + Axis Improve mobility of skull on rest of the vertebral column Cervical Neck vertebrae, make head mobile relative to body Thoracic Vertebrae of the thorax (chest) provide attachment for ribs Lumbar Vertebrae of the abdominal region, have transverse processes Sacral Form sacrum which attaches pelvis to the axial skeleton Caudal Forms the tail Vertebral Column of Tetrapods Cervical Region o Amphibians Single cervical vertebra (atlas) Allows little head movement o Reptiles Increased numbers of cervical vertebrae (usually 7; atlas and axis first 2) Increased flexibility of head Ball and socket joints between procoelus vertebrae enable head and neck to retract o Birds Atlas-axis complex Largest number of cervcal vertebrae (usually 12; as many as 25 in swans) Heterocoelus vertebrae allow for a 180-degree head turn o Mammals Usually 7 cervical vertebrae Dorsal Region

Dorsals name given to vertebrae between cervicals and sacrals when all articulate with similar ribs (e.g. fish, amphibians, and snakes) Crocodilians, lizards, birds and mammals have ribs confined to the anterior region of the trunk. Thoracic vertebrae with ribs Lumbar vertebrae without ribs Sacrum and Synsacrum Sacral Vertebrae have short transverse processes that brace the pelvic girdle and hindlimbs against the vertebral column o o o Amphibians Living reptiles and most birds Most mammals : 1 sacral vertebra : 2 sacral vertebra : 3-5 sacral vertebra

Sacrum single bony complex consisting of fused sacral vertebrae found when there is more than 1 sacral vertebra Synsacrum found in birds produced by the fusion of last thoracics, all lumbars, all sacrals and first few caudals fused with pelvic girdle provides rigid support for bipedal locomotion Caudal Region o Primitive Tetrapods 50 or more caudal vertebrae o Present-day Tetrapods Number or caudal vertebrae is reduced Arches and processes get progressively shorter (the last few caudals typically consist of just cylindrical centra) o Anurans Unique terminal segment called the urostyle (section of unsegmented vertebral column probably derived from separate caudals of early anurans) o Birds Last 4 or 5 caudal vertebrae fuse to form pygostyle o Apes and Humans Last 3-5 caudal vertebra fuse to form coccygeal (tail bone)

Flexibility of neck o 15-20 vertebrae (largest number of cervical vertebrae) o Heterocoelus centra (saddle-shaped at both ends) o Atlas-axis comples

Mammals Regional specialization (cervical, thoracic, lumbar, sacral and caudal vertebrae) Epiphyses 7 cervical vertebrae as a rule Approximately 20 trunk vertebrae (thoracic and lumbar) Swingers and jumpers (thoracic and lumbar proportions) Characteristics Body Amphibia Gravity and limbs Vertebrae: cervical, trunk, sacral, caudal Single cervical vertebra unfused to skull and allows dorsoventral mobility of head an advantage on land trunk vertebrae Development of fused sacral (single) vertebrae allows more rigid attachment of pelvic girdle to axis of trunk Reptiles Cervical (3-7) Smalle, wide side to side Short, projects directly posteriorly Triangular Contain foramena Sup facet sup-pos Inf facet inf-ant Flex and extension, lateral flex, rotation Thoracic Larger than cervical, heart-shaped, bears two costal demifacets Long, sharp, projects inferiorly Circular Bear facets for ribs (except T11-T12) Sup facets posterior Inf. facets anterior Rotation, limited lateral flex (ribs), flext and extend prevented Lumbar Massive, kidney shaped Short, blunt, projects directly posteriorly Triangular Thin and tapered Sup facet postmedian Inf. facet antlateral Flex and extend, lateral flexion, rotation prevented

Spinous Process Vertebral Foramen Transverse Process Sup. and Inf. Art. Process Movements

Terrestrial Cervical, thoracolumbar, sacral, caudal vertebrae Distinct neck st nd Atlas and axis 1 & 2 cervical vertebra Stronger limbs Tremendous variation o Snakes: 400 vertebrae with no specialization o Apodans: 250 vertebrae o Urodeles: 100 vertebrae

Ribs May be short, cartilaginous, or bony Articulate medially with vertebrae and extend into the body wall A few teleosts have 2 pairs of ribs for each centrum of the trunk (dorsal rib separates epaxial and hypaxial muscles) Most teleosts ventral ribs only Sharks dorsal ribs only Agnathans no ribs

Birds

Rigid back o Synsacrum Number of fused trunk vertebrae (last thoracics, all lumbars, all sacrals and first few caudals) and fused with pelvic girdle

Tetrapods Ribs usually articulate with vertebrae in movable joints Early Tetrapods

Ribs articulated with every vertebra from the atlas to the end of the trunk Modern Tetrapods Long ribs are limited to the thoracic region Thoracic Ribs most composed of a dorsal element (vertebral rib) and a ventral element (sternal rib) Sternal Rib may be ossified (birds) or remain cartilaginous (mammals) - usually articulate with sternum (except floating ribs) Uncinate Process found in birds - provides rib-cage with additional support Amphibians Short in anurans and urodeles Long in apodans Ribs are reduced/lacking Frogs only have sacral ribs

Birds

Costal/vertebral dorsal Sternal ribs ventral Uncinate processes for muscle attachment

Mammals Thoracic (12 in humans; 13 in cat) True ribs False ribs Floating ribs Thoracic vertebrae have 2-part ribs Mammals lack ribs in the lumbar and caudal regions Sternum is a series of separate bones (jointed rod) In humans, sternums is a 3-part plate

Reptiles Typically short in neck and long in trunk Turtles: thoracic ribs are long and fused to carapace : sacral ribs unite with the pelvic girdle : have no sternum Snakes: long and curved Crocodiles: gastralia are not ribs *Monocipital in snakes and monotremes *Bicipital - dorsal head: tuberculum DT - ventral head: capitulum VC

The Thoracic Cage The skeleton of the chest o Supports the thoracic cavity o Consists of: a. Thoracic vertebrae b. Ribs c. Sternum (breastbone) The rib cage o Formed of ribs and sternum Ribs (1-7) True Ribs o Vertebrosternal ribs o Connected to the sternum by costal cartilages Ribs (8-12) False Ribs o Do not attach directly to the sternum o Vertebrochondral ribs(ribs 8-10) Fuse together Merge with cartilage before reaching the sternum o Floating or vertebral ribs (ribs 11-12) Connect only to the vertebrae and back muscles Have no connection with the sternum

Tetrapod Sternum Sternum Strictly a tetrapod structure and primarily an amniote structure Amphibians No sternum in early amphibians and among modern amphibians Only anurans have one Amniotes Sternum is a plate of cartilage and replacement bone Articulates with the pectoral girdle anteriorly and with a variable number of ribs o Amphibians Frogs and toads poorly ossified Urodeles flimsy if present Caecilians none Reptiles Lizards large Crocodiles large cartilaginous plates Snakes none Turtles none needed Birds Keel Carina For muscle attachment Ribs meet sternum Large and ossified sternum with prominent keel Mammals Segmented sternum Xiphoid Process caudal most segment

Rib Structure Shaft Head posterior end; articulates with vertebrae Tubercle articulates with vertebrae Costal cartilage hyaline cartilage