1/9/2013. Divisions of the Skeleton: Topic 8: Appendicular Skeleton. Appendicular Components. Appendicular Components

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1 /9/203 Topic 8: Appendicular Skeleton Divisions of the Skeleton: Cranial Postcranial What makes up the appendicular skeleton? What is the pattern of serial homology of the limbs? Tetrapod front limb morphology Tetrapod hind limb morphology How do the limbs develop? What patterns of evolution have limbs undergone? Axial Appendicular What are the components of the appendicular skeleton? Transition from water to land What are some specializations of the limbs? Liem et al. Fig. 8-5, 8-8; Dean 895 Appendicular Components Appendicular Components Multiple elements In body proper Single long bone Many bones How have these bones developed? Modified from Liem et al. Fig. 9-7 Radius & Ulna Tibia & Fibula Autopodium Humerus Femur Zeugopodium Two long bones Variable elements (see later) Stylopodium Girdle Carpals, Metacarpals & Phalanges Tarsals, Metatarsals & Phalanges Liem et al. Fig. 9-0, 9-

2 /9/203 Starts with formation of _ What germ layers are involved? Early formation of (AER) Zone of cell proliferation Zone of differentiation Proximal structures develop first Zone of (ZPA) Expresses Sonic Hedgehog to define digit polarity P Martin; ncbi.nlm.nih.gov E Uthman; BioDes; cmrb.eu Fibroblast Growth Factor (FGF) Induces AER formation Proliferation of Mesoderm Formation of digits Bone Morphogenetic Protein (BMP) Involved in apoptosis Noggin gene Blocks apoptosis Limb element identity expression patterns Slight differences between front and hind limbs BioDes Hall, Fig

3 /9/203 Solid line indicates limb axis Gene expression data inform us about homology Are tetrapod digits homologous to radials or are they neomorphs? Liem et al. Focus 9-

4 /9/203 Front Limb & Pectoral Girdle Evolution Front limb differentiation Addition of elements Increased of elements Evolution of Front Limb & Pectoral Girdle Evolution _ Cleithrum, supracleithrum, interclavicle lost Clavicle remains _ Scapula and coracoid increase in size Mammal Coracoid lost in mammals Liem et al. Fig. 9-5, 9-6, 9-8 What has happened in terms of what elements are involved? Liem et al. Fig. 9-5, 9-9, 9- Front Limb & Pectoral Girdle Evolution Hind Limb Evolution Bony Fish Separation of front limb & girdle from the head skeleton Why? Very similar to front limb evolution Evolution of autopodium Evolution of digits Increased robustnesss Panderichthys Basal Amphibian Benton 997 pg 83; Benton Fig 4-2a; Pough et al 2004 Fig 2-2 Liem et al. Fig. 9-4, 9-6, 9-0; Hall 4-3 Pelvic Girdle Evolution How does this differ developmentally from the pectoral girdle? Bones are conserved in position Mammal Pelvic Girdle Evolution Transition in articulation of pelvic girdle Fishes: _ Tetrapods: solidly articulated with vertebral column Why? Protopterus (African lungfish) Acanthostega Liem et al. Fig. 9-4, 9-9, 9- Benton 997 Fig 3.22; Pough et al 2004 Fig 2-2a

5 /9/203 Evolution of Limbs Transition from water to land Obligate Aquatic What morphological changes are involved in this evolutionary transition? What are the functional reasons for each change? Terrestrial Evolution of Limbs What morphological changes are involved in this evolutionary transition? What are the functional reasons for each change? So, how does the function of front & hind limbs differ?

6 /9/203 Specialization of the Limbs Limbs to Fins Shortening of stylopodium & zeugopodium Fusion of digits (not phalanges) Specialization of the Limbs - Wings Digits -3 reduced Phalanges in digit 4 elongated Metacarpal 4 elongated Loss of digits & 5 Digits 2-4 reduced Carpometacarpus Elongation of radius & ulna Phalanges & metacarpals in digits 2-5 elongated Digit reduced Radius elongated Ulna reduced Storrs, 993 Liem et al. Fig. 9-2 Limb Specializations Limb posture and cursoriality _ _ _ Fusion of tibia & fibula Elongation and fusion of metatarsals Fusion of phalanges & loss of digits Limb Specializations A dog at full gallop with long strides How does this improve speed? What species have each posture? UMMZ Liem et al. Fig. 9-4 Limb Specializations Human skeletal adaptations for endurance running Narrow waist Large vertebral centra Relatively short Large Long Spring-like Short _ Relatively After Lieberman et al. 2009

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