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Post-Cranial Skeleton Axial skeleton = vertebrae, sternum, & ribs; endochondral bone. Appendicular Skeleton Pectoral skeleton supports pectoral appendages (arms); dermal and endochondral bone. Pelvic skeleton supports pelvic appendages (legs); endochondral bone. Human Vertebra neural arch superior articular process transverse process body (centrum) inferior articular process spinous process (neural spine) facet for head of rib facet for tubercle of rib Human Vertebrae Intervertebral discs between (annulus and nucleus) Cervical Vertebrae – 7; #1 = atlas, #2 = axis; forked neural spine on #2-#7 Thoracic Vertebrae – 12 (ribs articulate) Lumbar Vertebrae – 5 (no ribs) Sacral Vertebrae – 5; fusion begins at 16-18 and usually completed at 25 Coccygeal Vertebrae – 3-5; may become fused (= caudal vertebrae) Human Vertebrae Intervertebral Disc Problems Atlas, Axis, & Other Cervical Vertebrae transverse foramen (vertebral artery) Thoracic Vert. Articulate w/ Ribs costal cartilage Human Ribs & Sternum Ribs – 12; on thoracic vertebrae True Ribs – 7; contact sternum directly False Ribs – 5; don’t contact sternum directly “False” / Vertebrochondral Ribs – 3; fuse to a single costal cartilage element Floating Ribs – 2; don’t contact sternum Sternum – segments fuse to form 3 bones; manubrium, body, xiphoid process (x.p. doesn’t articulate with ribs) Human Ribs Human Sacral/Coccygeal Vert. sacrum coccyx Vertebral Development Sclerotome = forms vertebral elements Myotome = forms muscle Both separated into segments when the somites form. Vertical myosepta exist between segments of myotome (derived from the somite) . The myotome is also divided by a horizontal septum. Neural & hemal arches are centered on the vertical myosepta. Vertebral Development Vertebrate Neurula (section) somite notochord gut intermediate mesoderm lateral plate mesoderm Vertebrate Embryo (section) neural crest cells sclerotome dermatome myotome splanchnic mesoderm somatic mesoderm gut intermediate mesoderm lateral plate mesoderm Vertebrate Embryo (section) neural crest cells sclerotome dermatome myotome splanchnic mesoderm somatic mesoderm gut intermediate mesoderm lateral plate mesoderm Vertebrate Embryo (section) pigment cell precursors sclerotome (neural crest) ganglionic neuron precursors dermatome myotome dorsal & ventral mesenteries (splanchnic mesoderm) (neural crest) gut coelom lateral plate mesoderm Vertebrate Embryo (section) vertebra horizontal septum dermatome myotome dorsal & ventral mesenteries (splanchnic mesoderm) gut coelom lateral plate mesoderm Vertebrate Embryo (section) vertebra horizontal septum dermatome myotome dorsal mesentery (splanchnic mesoderm) gut coelom lateral plate mesoderm Fish Section Comparative Vertebra Structure Hagfishes lack vertebral structures. Neural arches dorsal to notochord surrounds dorsal nerve cord. Neural spines project above neural arch Hemal arches ventral to notochord around blood vessels in “tail” Hemal spines project below hemal arch notochord Vertebral Centra Centrum = solid bone or cartilage vertebral “segment;” “replaces” part of notochord Intervertebral pads = separate centra; derived, in part, from the notochord Centra evolved at least three times in vertebrates (Chondrichthys, Tetrapoda, & Amia + Teleosts) Vertebral Regionalization Non-tetrapod vertebrates Trunk (Thoracic, Abdominal) vertebrae = anterior; often bear ribs, lack hemal arches Caudal vertebrae = posterior; rarely bear ribs, have hemal arches Tetrapods Have Ant. & Post. Articulatory Processes Cervical vertebrae = neck vertebrae Thoracic vertebrae = have ribs (Lumbar vertebrae = lack ribs) Sacral vertebrae = attached to pelvic girdle Vertebral Regionalization Bird Vertebral Regionalization Synsacral vertebrae = vertebrae attached to pelvic girdle (homologs of sacral, lumbar, some thoracic and some caudal vertebrae) Cervical: Atlas & Axis Atlas = first cervical vertebra in tetrapods; reduced centrum, large neural canal Axis = second cervical vertebra in amniotes (Lissamphibians have only 1 cervical vertebra, an atlas.); odontoid process (dens) for rotation of atlas, large neural canal Atlas/Occipital Condyle joint = tilting Atlas/Axis joint = rotation Vertebral Regionalization Group “fish”: hagfishes, lampreys, chondrichthyans, lungfish, actinopterygiians, Trunk Vertebrae many trunk (number greatly variable) amphibians 1 cervical, few to many trunk, 1 sacral mammals 7 cervical, 13-27 trunk (9-20 thoracic & 4-7 lumbar), 3-5 sacral, turtles, lizards & snakes, & crocodilians 7-9 cervical, few to many trunk (especially many in snakes), 2 sacral (0 in birds snakes) 11-25 cervical, 3-10 trunk, 10-22 synsacral Caudal (“Tail”) Vertebrae vary greatly in number, even within groups. Subphylum Vertebrata centra centra R S G AT centra; atlas; sacral vertebrae; articulatory processes axis, >1 sacral vertebrae (Amniota) Ribs Form within myosepta from cartilage. Dorsal ribs form where the myoseptum intersects the horizontal septum Ventral ribs form where the myoseptum intersects the coelom somatic mesoderm. HOMOLOGY CONTROVERSY Vertebrate Embryo (section) vertebra horizontal septum dermatome myotome dorsal mesentery (splanchnic mesoderm) gut coelom lateral plate mesoderm Ribs dorsal rib ventral rib Tetrapod Ribs tuberculum dorsal rib? or ventral rib? capitulum Sternum Sternum (midventral endochondral element) where ribs often articulate Evolved independently in most groups. Mammalian sternum composed of multiple ossified elements (stern - ebrae); manubrium = 1st; xiphisternum/xiphoid process = last Subphylum Vertebrata * ** 2 headed dorsal ribs, ventral ribs lost (Tetrapoda) Ribs (of some kind) (Gnathostomata) *segmental sternum **solid sternum Human Pectoral Skeleton Human Pectoral Girdle Clavicle – (collar bone) large, connects acromion process of scapula and superior sternum; dermal Scapula – (shoulder blade; endochondral Acromion process – contacts clavicle Coracoid process Glenoid fossa clavicle Scapular blade supraScapular spine sternal acromion notch Human Scapula acromion coracoid process coracoid process glenoid fossa scapular spine scapular blade acromion coracoid process Human Arm Bones Human Arm External styloid process of ulna lateral epicondyle of humerus olecranon process of ulna Human Hand (Manus) IV III II V I pisiform Human Hand (Manus) Bones Human Hand External Human Pelvic Skeleton Human Pevic Girdle Innominate Bone – pelvis; 3 pairs bones; connects to sacral vertebrae; endochondral Ilium – fused to sacrum Ischium – posterior & inferior Pubis – anterior & inferior Obturator foramen – btw. pubis & ischium Acetabulum – femur articulates Human Pelvic Girdle ilium sacrum coccyx pubis acetabulum obturator foramen ischium Human Pelvis External iliac crest (iliac spine) iliac crest (iliac spine) Human Leg Bones Human Leg External lateral epicondyle of femur patella medial epicondyle of femur head of fibula tibial tuberosity tibial shaft (“shin”) Human Pes Human Foot (Pes) Bones --Only Gnathostomes Have-Chondrichthyan Pectoral & Pelvic Pectoral girdle =cartilaginous scapulocoracoid Pelvic girdle = cartilaginous puboischiac bar Radials = support keratinous “fin rays” puboischiac bar scapulocoracoid radials Osteichthyan Pectoral Girdles Scapulocoracoid (or scapula & coracoid) = endochondral; articulate(s) with radials (also endochondral) Extensive, dermal pectoral girdle Posttemporal, supracleithrum, cleithrum, clavicle, and interclavicle (median & unpaired) Fin rays = dermal bony rods (support most of fin); modified lamellar bone “scales” Osteichthyan Pectoral Girdle Posttemporal Supracleithrum Cleithrum Clavicle Interclavicle Scapulocoracoid Lepidotrichia Radials Osteichthyan Pectoral Girdles Actinopterygiians - loss of ventral dermal bones (Teleosts & Amia lack clavicles & interclavicles. Gars lack interclavicles, but have clavicles.) Tetrapods - loss of dorsal dermal bones (lack posttemporal, supracleithrum, & cleithrum) Humerus = single radial articulates with “scapulocoracoid” in sarcopterygiians. (Glenoid fossa = articulation point) In Tetrapods - Humerus Radius & Ulna Carpals Metacarpals Phalanges Tetrapod Pectoral Appendages Scapula Glenoid fossa Humerus Ulna Clavicle Coracoid Carpals Interclavicle Radius Metacarpals Phalanges Amniote Pectoral Girdles Clavicles & interclavicle in turtle ventral shell. furcula = fused clavicles & interclavicle in birds. postcoracoid = endochondral ossification unique to mammals; fuses to scapula in therians (= coracoid process of scapula) Therian mammals lack interclavicle and coracoid. Echidna Skeleton Dermal Pectoral Girdle Group interclavicle clavicle cleithrum supracleithrum posttemporal reedfishes and sturgeons present present present present present gars absent present present present present teleosts & Amia absent absent present present present lungfishes present present present present absent amphibians present present absent absent absent monotremes present present absent absent absent Theria absent present absent absent absent turtles, birds, & lizards/snakes present present absent absent absent crocodilians present absent absent absent absent Endochondral Pectoral Girdle Group Chondrichthys reedfishes sturgeons & paddlefish, gars, & Amia teleosts lungfishes amphibians scapula present, fused to → present present, fused to → present present, fused to → coracoid present, fused to ← present present, fused to ← present present, fused to ← postcoracoid absent absent absent absent absent present present absent sometimes fused to → sometimes fused to ← monotremes present present present Theria present absent present, fused to scapula (= coracoid process) Reptilia present present sometimes fused to → sometimes fused to ← absent Subphylum Vertebrata **loss of interclavicle *loss of clavicles R S G AT * ** loss of cleithrum, supracleithrum, dermal & posttemporal pec. girdle (Osteichthys) (have “neck”) (Tetrapoda) pectoral & pelvic appendages (Gnathostomata) Subphylum Vertebrata humerus, (femur) (Sarcopterygii) radius, ulna, metacarpals, (tibia, femur, metatarsals), phalanges (Tetrapoda) Amniota posterior coracoid loss of interclavicle & anterior coracoid loss of clavicles furcula clavicles & interclavicle in ventral shell Osteichthyan Pelvic Girdles turtle articulates with plvic girdle in sarcopterygiians. (Acetabulum = articulation point) il is pu anterior ALL Endochondral Not in actinopts. = radials. 3 “bones” in tetrapods. Ilium = contacts sacral vert. Pubis = Anterioventral Ischium = Posterioventral Femur = single radial Tetrapod Pelvic Appendages Ilium Acetabulum Femur anterior Fibula Pubis Tibia Ischium Tarsals Metatarsals Amniote Pelvic Girdles In Archosaurs the bird pu anterior acetabulum is “perforated.” In birds the 3 pelvic bones is fuse (innominate bone); form a synsacrum. In mammals the 3 pelvic bones often fuse (innominate bone). il Subphylum Vertebrata ilium, ischium, & pelvic girdle pubis (Tetrapoda) lost; fused basal radials pelvic girdle Amniota synsacrum; tibiotarsus; tarsometatarsus perforated acetabulum (Archosauria) Tetrapod Digit Terminology Manus = “hand” Pes = “foot” Digits numbered anterior to posterior. Ancestrally 5 digits per limb in amniotes . Pollux = “thumb;” digit I of manus Hallux = “big toe;” digit I of pes Phalangeal formula = number of phalanges in each digit starting with digit I. (e.g., 2-3-3-3-3) Ancestral Tetrapod Manus distal carpals metacarpals ulnare ulna radius intermedium phalanges V IV 5 4 4 3 3 2 2 1 1 III II radiale centrales I Ancestral Tetrapod Pes distal tarsals metatarsals fibulare fibula tibia intermedium phalanges V IV 5 4 4 3 3 2 2 1 1 III II tibiale centrales I Human Manus Bird Manus & Pes Birds: much fusion of elements Only 2 free carpals Carpometacarpus = distal carpals + metacarpals of digits III & IV Tibiotarsus = tibia + proximal tarsals Tarsometatarsus = distal tarsals + metatarsals Amniote Manus & Pes Talus = tibiale + intermedium (called Astragalus in non-mammals) Mammals - Maximum of only 1 free centrale (humans = 0 in manus, 1 in pes) Pisiform = sesamoid ossification on manus Calcaneus = fibulare with posterior process; forms heel Genetralized Amniote Pes distal tarsals Calcaneus metatarsals phalanges V IV (fibulare) fibula tibia 5 4 4 3 3 2 2 1 1 III II Astragalus (tibiale + intermedium) centrales I Amniota calcaneus (= “heel” on fibulare); tibia-tarsal joint carpometacarpus; tibiotarsus; astragalus / talus tarsometatarsus (not an intratarsal joint) (= tibiale + intermedium)