10 - skeleton ii

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    Lecture 10The Dermatocranium and Axial SkeletonKK Chapter 7,8; H&G Chapter 8,9

    The dermatocranium changes relatively little from early bony fishes through to

    mammals. Many of the changes involve simplification by loss of bones,

    especially the back of the skull, in the transition from fish to tetrapod.

    (Fish have no necks, tetrapods do!)

    However, the dermatocranium changes more among teleost fishes where

    adaptations for feeding cause extensive modifications. We will not cover

    these changes.

    We will start by looking at the skull ofAmia, a primitive bony fish, and followthe changes to mammals.

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    Amia KK 7.23, H&G 8.11

    Amia has a skull typical of early Osteichthyes, and not unlike

    Crossopterygii. Many of the dermatocranium elements are homologouswith those of tetrapods. Early tetrapods lack the opercular bones and the

    bones connecting the pectoral girdle to the head and in that location they

    have instead a flexible neck. The bones shaded in pink are part of the

    pectoral girdle. The red line marks the part of the skull that is lost with the

    gills.

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    Dermal Bones of a Tetrapod

    The major difference relative

    to Amia is the loss of bones

    at the back of the head. Thistrend will continue with loss

    of postparietals and

    temporals, most of the orbital

    series, and simplification of

    the lower jaw.

    KK 7.10 H&G 8.6

    TopView Palate

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    Early Amphibian (KK 7.37, H&G 8.12) Early Reptile (KK 7.33)

    Turtle (KK 7.37, H&G 8.15)

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    The dermal skull roof changes relatively little in tetrapods,

    except for appearance of openings in the temple. In many

    mammal and birds the temple opening converges with the orbit.

    KK 7.34,H&G 8.18

    What are

    euryapsids?

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    Temporal Openings in Tetrapods

    Temporal openings allow for muscles closing the jaws to bulge, andprovide better points of attachment.

    Note the zygomatic arch. Some mammals have a sagittal and/or nuchal

    crest on the top or back of the head to provide even better attachment

    points.

    KK 7.35, H&G 8.17 Frontal sections.

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    The temple opening in

    humans. Humans are

    among the mammals

    (apes, artiodactyls)

    where the temple

    opening is not

    confluent with the

    orbit.

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    From Internal Nares to Secondary Palate

    Choanate vertebrates (Sarcopterygii, tetrapods)

    have a connection between the external naris

    and the mouth. In mammals, this connection

    between respiratory and digestive pathways

    moves to the back of the mouth because of the

    hard and soft palate.

    early reptile (Parareptilia)

    KK 7.57, H&G 8.19

    KK 7.27

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    The Secondary Palate

    The secondary palate is formed by

    folds in the marginal bones of the

    palate (premaxilla, maxilla and

    palatine).

    The space between the original andsecondary palate in mammals

    contains turbinate bones.

    The secondary palate is also found in

    turtles and crocodiles.

    Why did this evolve? And what arethe implications for the evolution

    of endothermy?

    Palatine fissures?

    KK 7.58, H&G 8.19

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    Review: the skull of the

    opossum, Didelphis

    KK 7.51

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    The Axial SkeletonKK Chapter 8, H&G Chapter 9

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    Notochord

    Vertebral Column

    Ribs, sternum, gastralia

    Median Fins (caudal, dorsal, anal)

    Parts of the Axial Skeleton

    Embryological origins

    notochord, schlerotome andmesenchyme

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    The Notochord

    found in all Chordata

    found in all vertebrate embryos, plus adults of

    Agnatha, Placodermi, Acanthodii, Holocephali, some

    Actinopterygii (not teleosts), many Sarcopterygii, and

    the first Amphibia mechanically important for swimming

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    Centra in bony fish provide the same mechanicalproperties as the notochord.

    KK 8.19

    intervertebral

    ligament

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    Fish with notochords have bony or cartilaginous pieces (neural or hemal

    arches or spines) associated with the notochord. Fish with vertebrae tend

    to have biconcave (amphicoelous) centra. Tetrapod centra tend to be

    acoelous (platyan); more stout. Procoelous or opisthocoelous centra may

    be found where more flexibility is required, e.g. tail or neck.

    KK 8.1, H&G 9.4

    KK 8.4, H&G 9.2

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    Vertebrae of Fish

    The ribs are between the muscle blocks (myomeres) of the fish trunk.

    The dorsal ribs follow the horizontal septum between the epaxial and

    hypaxial muscles.

    KK 8.6

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    Development of the Vertebral Column

    Vertebrae arise

    from thescherotome, and

    begin as two

    elements per

    segment. In most

    cases they fuse to

    make a singlecentrum, but do so

    in an inter-

    segmental fashion.

    Where they

    remain separate,

    they are calledpleurocentrum and

    intercentrum.

    KK 8.10 & 8.13, H&G 9.3

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    Evolution of the vertebra in the Amphibia

    A labyrinthodont with equal inter-and pleurocentra (above). In the

    lineage leading to amniotes, the

    pleurocentra dominate.

    KK 8.24, 8.26. H&G 9.7

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    Caudal Fins

    The tail fin issupported by

    flattened neural

    and especially

    hemal arches.

    The tail of fishes

    with homocercal

    tails begin as

    heterocercal.

    KK 8.20,H&G

    9.14

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    The impact of terrestrial life on the axial skeleton

    Centra expand, fuse and replace the notochord.

    Intervertebral joints and processes

    (zygapophyeses) develop that restrict or facilitate

    movement depending on the location Processes to increase the leverage of muscles

    Connection to limb girdles.

    More flexible relationship with the head via

    cervical vertebrae + atlas (and axis).

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    Tetrapod

    Vertebrae

    Zygapophyses resist the tendency of

    the trunk to sag, and ribs now have two

    attachments to the vertebral column

    and meet at the ventral mid-line

    (sternum).

    Tetrapods do not have dorsal ribs, but

    their ribs often have two heads and

    attach to the sternum.

    KK 8.7, H&G 9.1

    R i l S i li ti f V t b

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    Regional Specialization of Vertebrae

    cervicals - small and flexible, with ribs small or absent

    thoracics - ribs, no hemal arches, reach sternum in tetrapods

    lumbars - no ribs, very stout

    sacrals - with heavy sacral ribs attaching to pelvic girdle

    caudals - flexible, with ribs small or absent. May be hemal

    arches, intercentra.

    R i l S i li ti f V t b

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    Regional Specialization of Vertebrae

    Fishes:thoracics and caudals only

    Amphibia:atlas, 1 cervical,thoracics, 1 sacral, andcaudals Reptilia:atlas, axis, cervicals,thoracics, 2+ sacrals, caudals

    Aves:atlas, axis, cervicals,thoracics, synsacrum,pygostyle

    Mammalia:atlas, axis, cervicals,thoracics, lumbars, 3+ sacrals,

    caudals

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    Atlas and Axis

    In amniotes, the first twovertebrae (atlas and axis) are

    specialized for nodding the head

    (occipital condyles articulate with

    the atlas) and rotating the head

    (atlas rotates on the axis on the

    axis).

    The latter joint is actually

    between the pleurocentrum of the

    atlas, now fused to the axis, and

    the intercentrum of the atlas.

    KK 8.27, H&G 9.10

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    Bull Skeleton