Comparative Anatomy of the Vertebrates:

[CHAPTER 8:

Vertebra, Ribs, and Sterna]

Chondrichthyes Cartilaginous vertebrae are never replaced by bones Teleosts, urodeles and apodans Membrane bone rather than cartilage is deposited except in arches; CHORDAL cartilage Anurans Replacement bone; notchord disappears

OUTLINE: I. Vertebral Column Morphogenesis of Vertebrae The Vertebral Column of Fishes Evolution of Tetrapod Vertebrae Rachitomous Vertebrae Types based on Shape Regional Specialization Cervical Vertebrae Sacrum and Synsacrum Tail Vertebrae II. Ribs Fishes Tetrapods III. Tetrapod Sternum Development of Amniote Sternum I. VERTEBRAL COLUMN: - Segmented, flexible, arched rod flanked by axial musculature; axial locomotor apparatus - Provides a protective bony tunnel for the spinal cord - In Fishes: articulations between vertebrae permit only SIDE-TO-SIDE flexibility (lateral undulation) - In Tetrapods: provides DORSOVENTRAL flexibility and a strong bowlike arch for suspending the trunk above the ground; functional separation of muscles in locomotion from ventilation (regional specialization) Centra - Occupy the position of the notochord Arches - Encloses a long vertebral or neural canal ~ Hemal arches or chevron bones in amniotes Processes - Transverse processes (diapophyses) articulate with the ribs; attachment for muscles - Zygapophyses are paired processes in the trunk vertebrae, chiefly in tetrapods; LIMITS dorsoventral flexion of the column - Parapophyses are lateral projections that serves as articulation site for the capitulum - Hypapophyses are midventral projections that are sites of muscle and tendon attachment Tetrapod tails have no zygapophyses thus flexible. Morphogenesis of Vertebrae 1. Sclerotome of mesodermal somites surround the notochord and neural tube 2. Sclerotomes produce blastema for a future vertebra 3. Scleroblast from caudal end of one somite + cephalic end of the next, stream INTERSEGMENTALLY (myomeres: segmentally)

Diplospondyly - In some fishes; two centra per segment; provides increased flexibility for locomotion The Vertebral Column of Fishes Living Agnathans ~ LATERAL NEURAL cartilages (limited to the tail in hagfishes) Sharks ~ constricted notochord ~ chordal and perichordal cartilage ~amphicelous ~ dorsal plates, dorsal intercalary plates, and wedge-shaped supradorsal cartilages Teleosts ~ Remnants are porous cartilagelike material ~ Centrum has dumbbellshaped vacuole ~ amphicelous ~ elastic ligaments; supraneural bones

Dipnoans, Chondrostei, and Latimeria ~ notochord unconstricted ~ develop no centra ~ amphicelous ~ basidorsal, basiventral, interdorsal and inerventral cartilages

Intercentrum - The other vertebrae in two centra per body segment (but only one arch); not homologous to hypocentra below - Incorporated with the interdrosal and interventrals Evolution of Tetrapod Vertebrae Rachitomous Vertebrae: 1. Hypocentrum or intercentrum; large, median, U-shaped anterior bone that cradled the notochord 2. Pleurocentra paired small wedges of bone overlying the notochord dorsolaterally 3. Laminae of bone rested in a notch between pleuroc. and hypoc. providing a neural arch Notochord is constricted at the level of hypocentrum. TREND: increase prominence of pleurocentra and concomitant reduction of hypocentra

Comparative Anatomy of the Vertebrates:

[CHAPTER 8:

Vertebra, Ribs, and Sterna]

Amphibians ~ One sacral vertebra Reptiles ~2 Birds ~2 ~ Synsacrum that provides axial rigidity for streamline body during flight Mammals ~ 3-5 ~ Sacrum ~ Armadillos have a synsacrum

Types of Vertebrae based on Shape: Amphicelous )_( ~ fishes Heterocelous Saddle-shaped ~ birds

Procelous )_) Evolved from the buildup of hypocentrum and coalition of intercentrum and centrum ~ anurans and modern nonavian reptiles Opisthocelous (_( Evolved from the coalition of hypocentrum and centrum behind it ~ salamanders (urodeles) Acelous |_| Intervertebral discs has in its core the remnant of embryonic notochord = PULPY NUCLEUS ~ mammals

Sacral vertebrae do not differentiate in tetrapods that lack hind limbs. Tail Vertebrae Anurans ~ urostyle: develops from a continuous elongated perichordal cartilage at the base of the larval tail Lizards ~ exhibits autonomy (to break off) of soft tissues that divide each tail vertebrae Birds ~ pygostyle: skeleton of the visible part of the tail; 4 Mammals ~ coccyx (4-5): tailbone

Regional Specialization in Tetrapod Columns 1. Sacral vertebrae 2. Cervical vertebrae 3. Thoracic vertebrae 4. Lumbar Vertebrae Snakes have almost no regional specialization. Longest columns Snakes Shortest columns Anurans Cervical Vertebrae

Some monkeys have PREHENSILE tail (capable of wrapping around an object.) Apodans have NO postanal tail! II. RIBS

Amphibians ~ only ONE cervical vertebra that lacks processes; preventing dorsoventral rocking ~ two-facet articulation

AMNIOTES ~ atlas (ringlike) and axis (with odontoid process); more independent movement of the head

Intersegmental development; same as vertebra but the development requires interaction of sclerotome and myotome Endochondral

Gastralia Proatlas - Resembling neural arch, interposed between the skull and atlas in crocodilians, Sphenodon, and spiny hedgehogs Birds ~ Exceptional flexibility of the neck; heterocelous Turtles ~ Uniquely flexible neck; ball-and-socket joint in the procelous centra Abdominal rib-like membrane bones in some reptiles; remnants of dermal exoskeleton

Fishes ~ Dorsal ribs separate the hypaxial and epaxial muscles ~ Ventral ribs arch ventrad in the lateral body and stop short of the midventral line Sharks: dorsal ribs only Most teleosts: ventral only Skates, chimaeras, sea horses and agnathans have no ribs

Mammals ~ always 7 cervical vert. except in sloth, Great Anteater, and manatees ~ transverse atlantal ligament holds the odontoid process in place Sacrum and Synsacrum Have stout transverse ankylosed stout ribs processes with

TETRAPODS ~ Ribs = Ventral ribs of fishes ~ Early to Late: Ribs became restricted to thoracic only ~ BICIPETAL: dorsal tuberculum and ventral capitulum ~ Two parts: costal rib and a more ventral sternal rib

Comparative Anatomy of the Vertebrates: Anurans and Urodeles ~ no such skeletal cage ~ very short and ankylosed to the transverse process ~ Ribs of anurans are NOT bicipetal

[CHAPTER 8:

Vertebra, Ribs, and Sterna]

Apodans ~ no such skeletal cage ~ ribs are quite long for locomotion

*Reptiles: no sternum in snakes and turtles *Ribs of cobras and Draco causes the skin folds to spread + inflation of lungs Uncinate process Provide birds with a lightweight but sturdy thoracic body wall skeleton for attachement of flight muscles In some lizards and Labyrinthodonts

In mammals, when the number of ribs is larger than 10, the rest are "floating. Note: o o Transverse foramen formed by two heads of the rib; in mammals and in birds Arteriovertebral canal tunnel formed by consecutive transverse foramen

III. T ETRAPOD STERNUM Tetrapod structure and predominantly amniote Endochondral

Functions: 1. A base braced by the pectoral girdles and ribs 2. Serves as anatomical origin of ventral muscles of the forelimbs 3. Largest in carinates, bats, pterosaurs (convergent evolution) for the attachment of flight muscles 4. Assists breathing movements In terrestrial mammals, sternum is composed of a series of bony segments = STERNEBRAE. In marine mammals, sternum is small and fused; adaptation: galloping mode of swimming Development of Amniote Sternum 1. Paired mesenchymal bars unite and undergo chondrogenesis 2. In many mammals, presternal and suprasternal blastemas also develop. 3. Presetrnal blastema = manubrium Suprasternal blastema = suprasternal ossicles