phylogenetically oldest: notochord (fibrous tube but hollow and full of water)
functionally replaced and regionally displaced by vertebral column
protect nerve cord
resist axial compression
muscle attachment (myosepta muscle connection)
suspend body (on land)
protect thoracic cavity (protect lungs)
aid in lung ventilation (breathing)
Vertebrae
neural arches phylogenetically first to appear, protect never chord
hemal arches in gnathostomes, protect blood vessels
centra in most gnathostomes displace notochord
two per body segment (one often lost or fused)
Vertebral articulation:
Amphicoelous(dish on both sides/both ways) - centra concave on both ends, fishes, some lizards
acoelous: centra flat ended mammals
procoelous/ opisthocoelous: centra form ball and socket joints (lissamphibia, reptiles)
heterocoelous: saddle-shaped joints between centra turtles, birds (thumb joint can be moved up/down and left/right)
Vertebrae parts
vertebrae offset with respect to myomeres
ribs and spines in mysepta, centra & arches bridge
ventral ribs (hemal arches) in fishes line peritoneum
dorsal ribs in transverse septum (epaxial/ hypaxial)
Vertebral Parts
arches and centra may be seperate elements or fused
Apidospondly - primitive fishes and tetrapods
Holospondly (hollow+solid): most extant vertebrates
most loose inter neural
Vertebral Parts
development of intercentrum and pleurocentrum has been influential in early tetrapods phylogeny
pleurocentrum most developed in amniotes
lissamphibia difficult to place
Vertebral embryology
mesenchymal cells of the primary sclerotome migreate to notochord ->
condesne between primary somites as secondary sclerotome
thought that each primary somite contributes to both anterior and posterior secondary sclerotome
Vertebral embryology
in primitive fish embryology, the sclerotome condenses into four parts pairs of cartilages, and this woud account for the four elements per vertebrae seen across many taxa
Vertebral Embryology
not so clear in teleosts, where centra directly replace the notochord
first the notochord sheath is replaced by the chondrocytes then replacement bone
then further direct ossification is added
Vertebral Embryology
not so clear in tetrapods where schlerentome surrounds the notochord to form a continuous though constricted tube
the tube becomes segmented
arches grow from the tube
Vertebral Differentiation
trunk and caudal vertebrae with little differentiation
except tail of teleost where vertebral elements become highly modified
Vertebral Differentiation
Atlas: nodding at occipital condyles
Axis: rotation with the atlas
cervical : ribs in sauropsids foramina for cervical ganglia
trunk: with true ribs, false ribs (doesnt connect to stirnum), or floating ribs, may be divided into throacic and lumbar
Vertebral Differentiation
Atlas: nodding at occipital condyles
Axis: rotation with the atlas
cervical : ribs in sauropsids foramina for cervical ganglia
trunk: with true ribs, false ribs (does not connect to sternum), or floating ribs, may be divided into thoracic and lumbar
sacral: fused to pelvic girdle - not fish
caudals - last
Appendicular Skeleton
pectoral and pelvic girdles - fish
medial fins and supports - fish
pectoral and pelvic girdles - mammals
limbs (chyridia)
Paired Fins
girdle in body wall, made of endochondral (replacment bones) and dermal elements
supports an endochondral fin base made of basal and radial pterygiophores
that supports dermal fin rays and fin webs - modified scales on top
Paired fins
Archipterygial (wings/fins)- with radial pterygiophores on either side of the basals
Metapterygial - with radial peterygiophores on anterior side of basals (later specialization)
Paired fins Origin
an early theory posited that the paired appendicular skeleton is derived from the gill arches - this doesnt explain the pelvic girdle - must be another theory
and the basals from gill rays
archipterygial fin would be primitive (gill arch theory - fins are modified gill arches)
paired fin origin
fin fold alternative hypothesis tat the paired fins are out grwoths of the body wall
metapterygial fin is primitive
Paired fin Origin
gene expression findings support fin fold hypothesis
TBX gene in amphioxsisexpressed ventro-latterally
two TBX genes in sharks + vertebrates (exactly where pelvic and pectoral girdles are
Sonic Hedgehog (shh) gene expressed in sarcopterygii extends length of limbs (expressed for complexity)
paired fin origin
Hox (found along the chromosome, anterior/posterior) gene expression shows that digits in tetrapods are derived from pterygiophores posterior to the basal pterygiophores axis
Pectoral Girdle
mainly dermal bone in actinopterygii and primitive sarcopterygii
attached to head, no mobile head allowed
incorporates some dermal bones in bony fish
Pectoral Girdle
reduction of dermal bony elements in tetrapods
often only clavicle or interclavicle remains of both completely lost
Pelvic Girdle
NEVER dermal elements
single element in fishes
does not attach to vertebral column
three ossifications in tetrapods - illium - connects with sacrum, ischium and pubis
Pelvic Girdle
ischium and oubis elongated in archosaurs
bipedalism
two patterns - lizard hips/ bird hips
Heterotopic Bones (occur in unusual places)
not part of the regular skeleton
Sesamoid Bones - aid tendons sliding over joints (knee cap)
Heart bone in deer and bovids
Eyelid bone in some archosaurs
Os penis (baculum) and Os clittoris in manu mammals including primates but lost in humans often used for identification